PN os) PON AEROS 8 L RSSSANGOR On PO Se RA So = a SO aa Gon ray V SMITHSONIAN “ores \ISCELLANEOUS COLLECTIONS @ao00000G5> {1S OBSERVATIONS, RESEARCHES, “SnVERY MAN IS A VALUABLE MEMBER OF SOCIETY WHO, BY f N’’—SMITHBON AND EXPERIMENTS, PROCURES KNOWLEDGE FOR ME (PUBLICATION 3063) GITY OF WASHINGTON PUBLISHED BY THE SMITHSONIAN INSTITUTION 1930 } R A 9 2 2 a 2 » ao = 5 ae ® a J Oo al 2 a » <4 wo b a oI o = a 5 2.) Large dorsal valve. Plesiotype. U. S. Nat. Mus., Cat. No. 78377. 2. (>2.) Ventral and-dorsal valves. Plesiotype. U. S. Nat. Mus., Cat. No. 78378. Mesonacts fremonts (Walcott) 5.0. saceee oo sts se ela ee eetene eleanor 6 Fic. 3. Fairly well preserved cephalon. The course of the posterior facial suture, the marginal suture, the occiptal and intergenal spines, and the general shape of the cranidium, together with the slightly advanced position of the genal spines, are clearly shown. Plesiotype. U. S. Nat. Mus., Cat. No. 78370. 4. Another cephalon in which the anterior margin on the left is disturbed by the peculiar slickensiding in the fossils from this locality. Size and position of the eyes and character of elabeller furrows are well shown. Plesiotype. U. S. Nat. Mus., Cat. No. 78380. 5. A third, less complete cephalon, well preserved on the right side, showing particularly the posterior facial and intramarginal sutures as well as the striations on the rim. Plesiotype. U. S. Nat. Mus., Cat. No. 78381. ‘ 6. Small cephalon with left eye practically complete. Plesiotype. U. S. Nat.’ Mus., Cat. No. 78382. Mould of portion of cephalon and thorax. Note extra width of third segment. Plesiotype. U. S. Nat. Mus., Cat. No. 78383. 8. ( 4.) Enlargement of genal angle of specimen illustrated in preceding figures, showing striated rim and course of the intra- marginal suture across the genal angle. 9. The associated hypostoma, referred to the species. Plesiotype. U. S. Nat. Mus., Cat. No. 78384. N * All figures natural size unless otherwise stated. SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLE eSiiINO em 2qmibtee nit Cambrian Fossils from the Mohave Desert. SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL 8 eNO 2a P io Cambrian Fossils from the Mohave Desert. NO. 2 CAMBRIAN FOSSILS FROM MOHAVE DESERT—RESSER 13 — DESCRIPTION OF PLATE 2 NESONMAGUSEANSOLEIISmIMe Wie SPECIES 4 sierc ssa aco eroielais seisteiore elnta a.nenid esis el «sins 8 Fic. 1. Poorly preserved carapace giving some idea of the shape and general characteristics of the species. Cotype. U.S. Nat. Mus., Cat. No. 78386. 2. Well preserved cephalon beside the hypostoma of Mesonacis fre- monti. Cotype. U.S. Nat. Mus., Cat. No. 78387. 3. Small cephalon illustrating the size and position of the eyes and the glabella. Note the occipital spine. Cotype. U. S. Nat. Mus., Cat. No. 78388. 4. A larger head in which the full size of the advanced genal spines is shown. Cotype. U. S. Nat. Mus., Cat. No. 78380. IMCS OnUGISTOIUSLOVEHSUSsMEW ES PECIESinc uaa ales stisice 2c siemie|ses «le eieie @ cists eicle sora 7 Fics. 5,6. Cephalon and enlarged (4) view of the glabella showing the surface features. Cotype. U. S. Nat. Mus., Cat. No. 78390. 7. Another cephalon with a fairly complete glabella. Cotype. U. S. Nat. Mus., Cat. No. 78301. 8. Fairly complete cephala of this species and of MM. insolens, show- ing the different angles at which the genal spines arise. Cotype. U. S. Nat. Mus., Cat. No. 78392. WESONGGIS MELONI ONNIALCOLL) lene rrale cai cjehereteere sic tetere ees Olors oie in Sinise eee eos se 6 Fic. 9. Portion of the thorax near the posterior end. Plesiotype. U. S. Nat. Mus., Cat. No. 78385. 14 SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL. 81 DESCRIPTIONJOER PEATE Pacdumias clarkiz, new SpeCieSa.4 on eae ates Oe ee ee 9 Fic. 1. A specimen preserving the major portion of the thorax. Note the narrow rim and the median ridge between it and the glabella. Two other cephala occur on the same piece of rock. Cotype. U. S. Nat. Mus., Cat. No. 78393. [ 2. Larger cephalon with eyes and glabella well preserved. Cotype. U. S. Nat. Mus., Cat. No. 78394. Paedumias nevadensis) (NWValcott)= donee ee eee 9 Fic. 3. Small, but fairly complete cephalon. Plesiotype. U. S. Nat. Mus., Cat. No. 78395. 4. Large cephalon showing the intergenal spines. Plesiotype. U. S. Nat. Mus., Cat. No. 78396. 5. Another large cephalon, somewhat crushed, causing this specimen to resemble P. clarki. .Plesiotype. U. S. Nat. Mus., Cat. No. 78307. 6. Smaller cephalon with the glabella better preserved. Note the occipital spine. Plesiotype. U. S. Nat. Mus., Cat. No. 78308. 7. Cephalon complete on the left side, showing position and size of the genal spine. Plesiotype. U. S. Nat. Mus., Cat. No. 78399. Mesonacis: fremonti, (Walcott) scceeeers aot ee eee ee ere 6 Fic. 8. Enlargement (x 4) of the left rear quadrant of the hypostoma shown on plate 1, figure 9. Plesiotype. U. S. Nat. Mus., Cat. No. 78384. Dolichometopus. 2 lodensis, (Clara aes ee eee eee 10 Fic. 9. The smaller shield shows the general characters of this species. The larger, less complete carapace is referred to Dolichometo- pus productus. Holotype. U. S. Nat. Mus., Cat. No. 78400. SMITHSONIAN MISCELLANEOUS COLLECTIONS VOES Sil), NOS 25) PE. 3 Cambrian Fossils from the Mohave Desert. . , even. j Tas, f 4 : : tt pe é i z 1 . ¥ ‘ \ j % 1 ; Z 4 - f ot 3 We q Oa ‘ ‘ a roK . t ¥ ‘ i } : { S , J pes ‘ ‘ it , a ‘ R s 4 $e r sel J E i . “ . ; i ; ; ( 4 ms . \ t 4 4 . " : ‘ ; . v ie ( ; j é s en \ ee y \ : “ 4 A z . t eh TNS a os “ 4 , om i ‘ j , SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLUME 81, NUMBER 3 MORPHOLOGY AND EVOLUTION OF THE ered HEAD AND ITS APPENDAGES BY R. E. SNODGRASS Bureau of Entomology (PUBLICATION 2971) CITY OF WASHINGTON PUBLISHED BY THE SMITHSONIAN INSTITUTION NOVEMBER 20, 1928 = SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLUME 81, NUMBER 3 MORPHOLOGY AND EVOLUTION OF THE INSECT HEAD AND ITS APPENDAGES Bi R. E. SNODGRASS Bureau of Entomology (PUBLICATION 2971) GITY OF WASHINGTON PUBLISHED BY THE SMITHSONIAN INSTITUTION NOVEMBER 20, 1928 pve] 2 ys a 2 e °o E £ oa 2 pe R °o a xs a » BALTIMORE, MD., U. S. A. MORPHOLOGY AND EVOLUTION OF THE INSECT HEAD AND ITS APPENDAGES By R. E. SNODGRASS BurREAU OF ENTOMOLOGY CONTENTS PAGE Um teOdUGtTO Mae eee renee teri, oh evar oate th Seer ce Ba hua es eh gomeubense 2 Pee olutonmotatherantinopodmneadinrm ay cee ne tres oe ea ana are 2 Cephalizationtemaeicis noc eo esos tee ne a ash tte vcard wears ony 3 Development of the body if segmented animals................... 12 Mhewprotocephalomancreces cs sci sta coche eile ct seiciely steals sere te 19 Wine Geinmivahne eredamopyodl Ineetlsooccccoosasaescnacnsnoceeb onsdopor 2 MenGeneqalestnictunerommne wnSecty Cadman ate tol eles eine eieler 33 Ativan leteeral: eanosiellies aoe ced Sad oddatkm, aes dot Onc sen Semon too cnr 34 AMitem al teiatmratclere pip la tayiexcees eee aes eee ener neers a arate eyetar Al MBC MSEOITIOM LIME eres one ey aier se eet esha! Biba erences 2 Malren Hn pOphahvaDsmcrias fers hres oe ee ok eter teal e in foyard ok 45 MHS LIOTSULITIIER Set rote oes ois See OG nets cro eetoc neaser ol cbsespors 50 tie hesneaduacappendagesemreemecemre ace ee te ne rete et ye i ro 5 Pblemamntenitnc cm werner tetcr ck aetete rien Ge ora AP ua en ee cra cee peloiate eye tena ok yers 57 shempostantennalmappendases. ca ac ie.5 see Rice veucsacierre ayo eboeyeiiena = 59 Mihiewan athiclbampemmaee sic. + Macc ir ote cer casiwaer met ttye alas eoers Sseillesanete: Sox's 60 Miiveestaaalm cli Lesimpetne tem eet teresa gers cia hota ceca PTR noes Hcl ee atc eomch a 62 Aliinematitss tural aera nie eh reteset hoe ys eter ge ied saruniic. Na eislovel 74 Milfemsecond eniascallacee asia orto ie ee ele cio examen ctety Ti Morphology of the gnathal appendages.........:.......-2+++0+- 79 EN ee Sinn yO LetIMIPOLLAM Ey POMS). 0.2. !naiis tae a beer Sem as cme gametes Fs = 90 WerhigetheadsotcamenassnOppenn ts tsea.cr saljca tsa tncy- co cieiela a= oti e ee Mth oe wes _Lm pe el ell C ant) §D Fic. 22.—Arthropod embryos showing relative development of the trito- cerebral appendages. A, embryo of a crayfish, Astacus (Potamobius) astacus (from Reichen- bach, 1877). B, embryo of a spider, Trochosa singoriensis (from Jaworowski, 1891). C, embryo of a spider, Angelena labyrinthea (from Balfour, 1880). D, embryo of an apterygote insect, Anurida maritima (from Wheeler, 1893). An, anus; Ant, antenna; rAnt, first antenna; 2Ant, second antenna; Ch, chelicera; //I, tritocerebral segment; rl, first leg; Li, prothoracic leg; Lim, labrum; Md, mandible; Mth, mouth; Pdp, pedipalp; Pi, pit on head region; Punt, postantennal appendage; Prc, protocephalon. belong to the second, or deutocerebral, segment of the protocephalon, the other appendages to the gnathal segments. In many insect em- bryos there is present a pair of small lobes on the third protocephalic segment, which lobes are unquestionably rudiments of the tritocere- bral appendages. Preantennal appendages have been reported in Scolo- pendra and in the phasmid insect, Carausius (fig. 14 A, B Prnt). As already pointed out, there is some reason for regarding the crustacean eye stalks as being the appendages of the preantennal segment, though the true status of these organs has not yet been demonstrated. NO. 3 INSECT HEAD—SNODGRASS 57 The eye stalks of the decapod crustaceans arise from the ends of a transverse ridge on the top of the protocephalon, and project later- ally from under the base of the rostrum, the latter being a process of the anterior edge of the carapace, and, therefore, from the tergum of the mandibular segment. Each eye stalk (fig. 17 B) consists of two movable segments, a narrow basal one forming a short peduncle, and a large terminal one capped by the hemispherical compound eye. Schmidt (1915) enumerates ten individual muscles for each eye stalk in the crayfish, the basal segment being provided with muscles arising on the head walls that move the appendage as a whole, while muscles from the basal segment move the terminal eye-bearing segment. The eye muscles are innervated by an oculo-motor nerve arising from the brain near the base of the sensory optic nerve. THE ANTENNAE The insect antenna is typically a many-jointed filament. Usually the first two basal segments are differentiated from the rest of the Fic. 23.—The antenna. A, diagram of typical segmentation and articulation of an insect antenna. B, head of a chilopod, Scutigera forceps, dorsal, showing dorsal articulation of antennae, and origin of antennal muscles on walls of cranium. Ant, antenna; as, antennal suture; E, eye; , articular pivot of antenna; Pdc, pedicel; Scp, scape; Fl, flagellum. shaft (fig. 23 A). The first segment serves to attach the antenna to the head, and, being often, thicker and longer than the others, forms a basal stalk, or scape (Scp), of the appendage. The second segment, or pedicel (Pdc), is short, and in nearly all insects contains a special sensory apparatus known as the organ of Johnston. The part of the antenna beyond the pedicel is termed the fagellui or clavola (Fl). The flagellum may be long and tapering and made up of many small segments, or it may be abbreviated, and reduced even to a single seg- ment. The scape is set upon a small membranous area of the head wall, sometimes depressed to form a cavity, or antennal socket. } : ; sen ati A 58 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 81 The head wall surrounding the antennal base is strengthened by an internal ridge, the line of which is marked externally by a suture (fig. 23, as), setting off a circular, marginal rim known as the antennal sclerite. Usually a pivot-like process (7) from the rim of the sclerite forms a special support and articular point for the base of the scape, and allows the antenna a free motion in all directions. In its single point of articulation with the head wall, the antenna resembles the maxilla, or the mandible of those apterygote insects in which the jaw does not have a double hinge with the cranium. In most pterygote insects the antennal pivot is ventral or postero-ventral in position, relative to the base of the antenna (fig. 23 A), while the single mandib- ular or maxillary articulations are dorsal. The ventral position of the antennal articulation might be supposed to have shifted during the forward and upward migration of the appendage from its primitive ventral and postoral situation; but in Japya the antennal pivot is dorsal, as it is also in the Chilopoda (fig. 23 B, 7). Each antenna is moved by muscles inserted upon the base of the scape. The origin of the antennal muscles in adult pterygote insects is commonly on the dorsal, or dorsal and anterior arms of the ten- torium (fig. 38 D, DT, AT), but in the caterpillars (fig. 50 B, C, E, F) and in some coleopteran larvae, the antennal muscles arise upon the walls of the epicranium. The cranial origin of the muscles is prob- ably the primitive condition, for, as already shown, the tentorium belongs to the gnathal segments only. The attachment of the anten- nal muscles on the tentorium, therefore, appears to be a secondary condition that has resulted from the migration of the muscle bases to the dorsal tentorial arms when the latter make contact with the dorsal wall of the head. In Crustacea and Chilopoda the antennal muscles have their origin on the head wall. In Scutigera (fig. 23 B) a dorsal set to each antenna arises on the dorsal wall of the cranium mesad and posterior to the antennal base, and a ventral set arises on the lat- eral walls below the antenna, and below the eyes. The insertion points of these muscles, distributed on three sides of the articular pivot (7), allow the muscles to act as levators, depressors, and rotators of the appendage. The part of the insect antenna distal to the scape is moved by muscles arising within the scape and inserted on the base of the pedicel (fig. 23 A). The segments of the flagellum in insects, however, so far as known to the writer, are never provided with muscles, and their lack of muscles suggests that the flagellum is a single segment secondarily subsegmented, corresponding with the flagellum of a crustacean antenna (fig. 24 B, Fl), which is a many-jointed dacty- lopodite. In the Myriapoda, however, all the antennal segments may NO. 3 INSECT HEAD SNODGRASS 59 be individually provided with muscles (Scolopendra, S pirobolus). The first antenna, or antennule, of the crayfish, according to Schmidt (1915), has paired antagonistic muscles for each of its first three proximal segments, and the third segment contains a single reductor ‘nserted on the base of the dorsal branch of the flagellum, but other- wise none of the flagellar segments is provided with muscles. The Arachnida and Xiphosura lack antennal appendages in the adult stage. Croneberg (1880) describes a pair of head lobes in the arachnid embryo, which he says fuse into a median rostrum in the mites and in the higher arachnids, and which he believes represent the antennal appendages. Jaworoski (1891) likewise describes in the embryo of a spider, Trochosa singoriensis, a pair of lobes situated before the chelicerae, which he claims are rudiments of the antennae (fig. 22 B, Ant), but he says the lobes disappear during later develop- ment. THE POSTANTENNAL APPENDAGES The pair of postantennal appendages on the tritocerebral segment of the head, known also as the antennae (Crustacea), second antennae, premandibular appendages, and intercalary appendages, are at best rudimentary in all insects. According to Uzel (1897), two small lobes in the adult head of Campodea, lying between the labrum and the maxillae, in the space left free by the retracted mandibles, are the tritocerebral appendages; the writer has found a pair of small papillae in Dissosteira between the bases of the mandibles and the angles of the mouth (fig. 42 B, Pnt) that might be vestiges of these organs. Otherwise tritocerebral appendages are known in insects only as evanescent rudiments in the embryo (fig. 22 D, Pnt). In the Myria- poda, likewise, the postantennal appendages are lacking, or possibly are present as temporary premandibular lobes on the head of the embryo (“rudiments of lower lip” in Geophilus, Zograf, 1883). In the Crustacea, on the other hand, the appendages of the tritocerebral segment, though sometimes reduced or lacking, are commonly highly developed, biramous organs, the second antennae, or “the antennae ” according to the terms of carcinology. In the decapods each appen- dage consists of a two-segmented base (fig. 24 B, Prtp), of a large, one-segmented exopodite (Exp), and of a long, slender endopodite (Endp), of which the terminal segment is the many-jointed flagellum (FI). The exopodite is independently movable by abductor and ad- ductor muscles arising in the second segment of the base. In Xiphosura and Arachnida, the chelicerae (fig. 24 A) are gen- erally regarded as the appendages of the tritocerebral segment. Their 60 SMITHSONIAN MISCELLANEOUS COLLECTIONS voL. 81 rudiments in the embryo of a spider (fig. 22 C, Ch) bear a relation- ship to the head so similar to that of the tritocerebral rudiments in the insect embryo (D, Pnt), that the identity of the two sets of organs can scarcely be questioned. Holmgren (1916), furthermore, claims that the histology of the arachnid brain shows that the chelicerae are innervated from the tritocerebral region of the brain. If this homology is correct, there is no reason for calling the tritocerebral appendages “second antennae ’”’ except in the Crustacea. The arachnid chelicera is a uniramous organ, that of a scorpion (fig. 24 A) having three well-developed segments. Fic. 24.—Postantennal appendage of adult arthropods. A, chelicera of a scorpion, left, ventral view, showing uniramous structure and three segments. B, second antenna of a decapod crustacean (Spirontocaris groenlandicus), left, ventral view, showing biramous structure, consisting of two-segmented base (Prtp) bearing an exopodite (Exp) and an endopodite (Endp). THE GNATHAL APPENDAGES There can be no doubt that the gnathal organs—the mandibles, the first maxillae, and the second maxillae—constitute a distinct group of appendages in the eugnathate arthropods. The mandibles are the most highly modified of the gnathal appendages, and, in most cases, their structure has lost all resemblance to that of the more generalized insect maxillae. A maxillary appendage, therefore, should be studied first as affording a better example of the basic structure of the gnathal organs, and, in insects, the first maxilla preserves most nearly the primitive structure, since the second maxillary appendages are united to form the labium. The first maxilla of an insect with typical biting mouth parts, of which the roach offers a good example (fig. 25 A), consists of a basal stalk, two terminal lobes, and a palpus. The base is divided into a proximal cardo (Cd), suspended from the head by a single point of articulation (e), and a distal stipes (St). The cardo and stipes are freely flexible on each other by a broad hinge line, and their planes may form an abrupt angle at the union, but neither has an inner wall, NO. 3 INSECT HEAD—SNODGRASS O1 the two being merely strongly convex sclerites set upon the mem- branous lateral wall of the head, and their cavities are a part of the general head cavity. Only the terminal maxillary lobes and the palpus are free parts of the appendage. The lobes arise from the distal end of the stipes, one, the Jacinia (Lc), being internal, the other, the galea (Ga) external. The galea is also anterior to the lacinia (or dorsal to it in insects with the head flattened and held horizontal). The 1 LW , Fic. 25—Maxilla of Periplaneta. A, left maxilla, posterior (ventral) surface. B, internal surface of cardo. C, right maxilla, anterior (dorsal) view, showing muscles. Cd, cardo; e, articulation of cardo with cranium; fga, flexor of galea; ficc, cranial flexor of lacinia; fics, stipital flexor of lacinia; ft, femoro-tibial joint of palpus; Ga, galea; J, promotor of cardo; KLcd, adductor of cardo (origin on tentorium); AZLst, adductor of stipes (origin on tentorium) ; Lc, lacinia; O, levator of palpus; Plp, palpus; rplp, first segment of palpus; Q, depressor of palpus; g, submarginal suture (and internal ridge) near inner margin of stipes; v, internal ridge of cardo; St, stipes; 7, depressor of fourth segment (tibia) of palpus; ”, depressor of fifth segment (tarsus) of palpus. galea is usually a soft lobe; the lacinia is more strongly chitinized, and ends in a strong incisor point provided with one or more apical teeth curved inward. Both lobes are movable on the end of the stipes : the galea can be deflexed, and the lacinia can be flexed inward. The palpus (P/p) arises from the lateral surface of the stipes, a short distance proximal to the base of the galea. The palpus of the roach is five-segmented. The musculature of the maxilla (fig. 25 C) comprises muscles that move the appendage as a whole, and muscles that move the terminal 62 SMITHSONIAN MISCELLANEOUS COLLECTIONS voL. 81 lobes and the palpus. The first group includes a tergal muscle (/) arising on the posterior dorsal wall of the head, and two sets of sternal muscles (KLed, KLst) arising on the tentorium in most insects, or on the homologous hypopharyngeal apodemes in some apterygote insects (fig. 30 B, HA). The single tergal muscle (fig. 25 C, J) is inserted on the proximal end of the cardo just before the articulation of the latter with the head (c) ; it is probably a promotor, serving to swing the appendage forward. The sternal muscles (i. e., the tentorial or hypopharyngeal muscles) consist of two large flat bundles of fibers, one group (KLcd) inserted on the inner face of the cardo, the other (KLst) on an internal ridge of the stipes near the mesal border of the posterior face of the latter (A, q). These muscles are the ad- ductors of the maxilla; the fibers of the cardo muscle arise anterior (or dorsal) to those of the stipes muscle and cross them obliquely. The muscles of the movable parts of the maxilla include muscles of the galea, the lacinia, and the palpus. The galea has a single muscle (fig. 25 C, fga) arising on the posterior wall of the stipes and inserted on the posterior rim of the base of the galea; it is a reductor in as much as it serves to flex the galea posteriorly (or ventrally). The lacinia has a large flexor (fics) arising in the base of the stipes, and a second muscle (fcc) arising on the posterior dorsal wall of the cranium. In the roach these two muscles are inserted by a common broad tendinous base on the inner proximal angle of the lacinia; in other insects they usually have separate insertions (fig. 30 B, fcs and flcc, fig. 40B, 14, 15). The palpus is provided with two muscles (fig. 25 C,O,Q), both of which arise within the stipes and are in- serted on the base of the first segment of the palpus (A, rplp). The two palpus muscles are more distinct in most other insects than in the roach (fig. 31 A, B, C, E), and since one is dorsal and the other ven- tral, relative to the morphologically vertical axis of the maxilla, they are clearly a levator and a depressor, or abductor (O) and adductor (Q), of the palpus. The muscles within the palpus vary somewhat in different insects. In the palpus of the roach, a levator of the second segment arises in the first, where also a long depressor of the fourth segment (7) has its origin. A depressor of the terminal segment (1/7) arises ventrally in the penultimate segment. THE MANDIBLES The most generalized mandibular appendage in the arthropods, i. e€., one corresponding most closely in structure and musculature with a typical maxilla, is to be found, not in the insects or crustaceans, but in the myriapods, and best developed in the Diplopoda. On. INSECT HEAD—SNODGRASS 63 The diplopod mandible consists of a large basal plate, which appears to form an extensive part of the lateral head wall (fig. 17 K, Md), and of a movable terminal lobe mostly concealed in the normal con- dition by the gnathochilarium (Gch). The basal plate is subdivided into several regions, but particularly there is a proximal piece (fig. 26-A, Cd) and a distal piece (St), separated by a line of flexibility. The proximal piece is loosely articulated to the head wall by a single point on its dorsal posterior angle (a). The entire mandibular base is slightly movable by its membranous union with the head, but it is not of the nature of a free appendicular structure, since it has no inner wall—it is merely a convex plate in the lateral wall of the head, but Fic. 26.—Mandibles of Myriapoda. A, right mandible of a diplopod, Thyropygus (Spirostreptus), dorsal, showing large dumb-bell adductors (KL, KL) from opposite mandibles, united by median tendon (k). B, left mandible of a chilopod, Scutigera forceps, lateral view. C, right mandible of Scutigera, dorsal, somewhat diagrammatic. a, articulation of mandible with cranium; BP, basal plate of mandible; Cd, “cardo” of mandible; flcc, cranial flexor of lacinia; flcs, stipital flexor of lacinia; J, promotor of mandible; J, remotor of mandible; k, median tendon of mandibular adductors; KL, mandibular adductors, united by median tendon in diplopod (A, k) to form dumb-bell muscle; Lc, lacinia; St, “ stipes ” of mandible. separated from the cranium by a membranous suture. The free ter- minal lobe of the mandible is a strongly chitinized, jaw-like structure with a proximal molar area and terminal incisor point (fig. 26 A, Lc). It is hinged by a dorsal articulation at its base with the end of the basal plate. So closely do the parts of the diplopod mandible (fig. 26 A) re- semble the cardo, the stipes, and the lacinia of an insect maxilla (fig. 25 A), that the imagination at once sees in the diplopod jaw an appendage similar to the maxilla, lacking only a galea and a palpus. That the fancied resemblance is real is easily demonstrated by a study of the musculature. The musculature of the diplopod mandible consists of muscles that move the appendage as a whole, and of muscles that move the lacinial 5 \ \ | ‘ { i 64 SMITHSONIAN MISCELLANEOUS COLLECTIONS voL. 81 lobe. As in the insect maxilla, the muscles that move the entire organ include a tergal promotor and a group of ventral adductors. The promotor (fig. 26 A, /) arises on the wall of the cranium dorsal and posterior to the articulation of the basal plate with the head. It is inserted on the dorsal (anterior) margin of the distal division (St) of the basal plate, and in its point of insertion alone does this muscle differ from the promotor of the insect maxilla, which is inserted on the edge of the cardo (fig. 25 C,/). Functionally, however, the two muscles are the same, and a shift in the point of attachment is not a morphological difference. The adductor muscles of the diplopod mandible consist principally of a great mass of fibers (fig. 26 A, KL) filling the cavity of both divi- sions of the basal plate (Cd and St). These muscles are clearly the homologues of the adductors of the cardo and the stipes in the insect maxilla (fig. 25, KLed, KLst), which have their origins on the ten- torium, or on the hypopharyngeal apodemes. In the diplopod mandi- bles, however, the fibers of the adductor muscles converge medially from each jaw upon a large, tough, transverse ligament (fig. 26 A, rk), and the two conical fiber masses, together with the connecting liga- ment, form a great dumb-hell-shaped muscle uniting the two man- dibles. The two sets of fibers pull against each other to close the jaws. Clearly, the inner ends of these muscles have become detached from the hypopharyngeal apodemes, and the fibers from opposite sides have been united across the middle of the head by means of a transverse ligament. There is also, however, a small group of adductor fibers to each mandible (not seen in the figure) that still retains a connection with the corresponding apodeme of the hypopharynx. Besides the mandibular muscles, other muscles have their origin on the transverse ligament, including muscles to the gnathochilarium, which is either the united second maxillae, or the combined first and second maxillary appendages. In the Diplopoda, therefore, the ventral adductors of all the gnathal appendages have lost their sternal connections by their de- tachment from the hypopharyngeal apodemes. This is a specialized condition, and the ligamentous bridge on which the muscles arise has no relation to the insect tentorium. The muscles of the free terminal lobe of the diplopod mandible (fig. 26 A, Lc) include a muscle inserted directly on the base of the lobe (fics) arising within the stipes (St), and a large cranial muscle (ficc) arising on the dorsal wall of the head and inserted by a strong, chitinous apodeme on the inner basal angle of the lobe. These muscles correspond exactly with the lacinial flexors of the insect maxilla, one of which (fig. 25 C, fics) arises within the stipes, the other (fcc) on the NO. 3 INSECT HEAD—SNODGRASS 65 dorsal wall of the cranium. In most insects the second muscle is in- serted, as in the diplopod, on a chitinous apodeme from the inner angle of the lacinia (fig. 30 B, fcc). There can be little question, therefore, that the single lobe (Lc) of the diplopod mandible is the lacinia, and that the jaw of the Diplopoda has a structure identical with that of the insect maxilla, except for the lack of a galea and a palpus. The mandible of the Chilopoda is more specialized in structure than is that of the diplopods, but in its musculature it is in some respects more generalized. In Scolopendra, Lithobius, Scutigera, the jaw is slender and greatly elongate. In Scutigera (fig. 17 G, Md) its taper- ing base is exposed on the side of the head where it is articulated to the cranial margin (a), but in Scolopendra (fig. 21 B) the end of the mandible is buried in a pocket of the head wall lying mesad of the base of the maxilla (/*C). The long basal plate of the chilopod jaw is undivided (fig. 26 B, BP), and is articulated to the head wall by its apical point (a). In some chilopods there is an anterior articulation between the mandible and the suspensorial plate of the hypopharynx, but this articulation is a mere contact between external surfaces. As in the diplopods, the basal plate has no inner wall. The distal part of the mandible is a free lobe (Lc) movable on the base, but not so definitely hinged to the latter as is that of the diplopod mandible. The musculature of the chilopod mandible is practically alike in both the Pleurostigma and the Notostigma, and is essentially the same as in the diplopods, though the muscles differ in relative size. The basal plate is provided with both tergal and sternal muscles. Of the former, there are two sets of fibers, one inserted on the dorsal (an- terior) edge of the proximal part of the plate (fig. 26 B, C,/), the other (J) on the ventral (posterior) edge; both have their origins on the dorsal wall of the cranium. These muscles apparently serve to rotate the mandible on its long axis, and they probably act as pro- tractors where the mandible is capable of a longitudinal movement ; but clearly the first would be a promotor, and the second a remotor in an appendage with primitive relations to the head. The sternal muscles of the mandible consist of a conical mass of adductor fibers (fig. 26 B,C, KL) spreading upon the inner wall of the basal plate from their median origin (fig. 21 B, AL), which is on the ligamentous bridge uniting the two apodemes of the hypopharynx (fig. 21 A, C, 7). The adductors of the chilopod mandibles are unquestionably homo- logues of the dumb-bell muscle of the diplopods. The condition of the mandibular adductors, therefore, is more primitive in the Chilo- poda, for here the muscles retain their connections with the sternal, hypopharyngeal apodemes. 66 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 81 The movable terminal lobe of the chilopod mandible (fig. 26 B, C, Lc) is provided with the same muscles as is the corresponding lobe of the diplopod mandible (A, Lc) and the lacinia of the insect maxilla (figs. 25 C, 30 B, Lc). The muscle from the lobe to the basal plate in the chilopod jaw is very large (fig. 26 B, C, fics), suggesting that of Japyx (fig. 30 B, fics), and is composed of two groups of fibers. The cranial muscle (flcc) arises by a broad base on the dorsal wall of the head, and is inserted on a slender apodeme from the inner angle of the lobe. In the chilopod mandible, therefore, there is a basal plate (fig. 26 B, BP) corresponding with the cardo and stipes of the insect maxilla, but not divided as in the diplopods, and a free terminal lobe (Lc) which represents the lacinia. In retaining the connection of the adductor muscles with the hypopharyngeal apodemes, the chilopod mandible preserves the primitive condition shown by the maxilla of Japyx (fig. 30 B). In the Crustacea and Hexapoda, the mandible, or the jaw part of the mandibular appendage, which may bear a palpus, consists of a single piece. Whatever may be the primitive elements that have entered into its composition, these elements are fused into a solid gnathal organ. There are, hence, never muscles entirely within the mandible, except those that pertain to the palpus, when a palpus is present. The mandibular musculature consists exclusively of the muscles that move the appendage as a whole, and these musé¢les cor- respond with the muscles of the basal plate of the myriapod mandible, or with those of the cardo and stipes of the insect maxilla. In the phyllopod crustacean Apus, the large mandibles (fig. 27 A, Md) hang vertically from the wail of the mandibular segment (JV). Each is a strongly convex, elongate oval structure, attached to the lateral membranous wall of the head by most of its inner margins, leaving only a ventral masticatory part projecting below as a free lobe. A single, dorsal point of suspension (a) allows the base of the man- dible to turn on its vertical axis, or to swing inward and outward as far as the membranous lateral head wall will permit. The musculature is correspondingly simple: two dorsal muscles from the tergum of the mandibular segment (/V’) are inserted on the! base of the jaw, one (7) on the anterior margin, the other (J) on the posterior margin; the hollow of the mandible is filled with a great mass of fibers (KL) which converge upon a median transverse ligament (k) that receives likewise the muscles from the opposite jaw. Here, then, is a ventral dumb-bell adductor, as in the diplopods, and two dorsal muscles, which may function either as productors and reductors, or as anterior and posterior rotators. It is not clear as to what constitutes the mechanism NO. 3 INSECT HEAD—SNODGRASS 67 of abduction in appendages with this type of articulation and mus- culature. The Apus type of mandible is probably characteristic of most of the more generalized Crustacea; it is present also in some of the = = = B HA Fic. 27—Mandibles of Crustacea and Apierygota. A, mandibles of Apus longicaudata (phyllopod), anterior. B, mandibles of Spirontocaris groenlandicus (decapod), anterior. C, mandibles of Heterojapyx gallardi (apterygote insect), anterior (dorsal). D, mandibles of Nesomachilis maoricus (apterygote insect), posterior. a, articulation of mandible with cranium, or with wall of mandibular seg- ment (JV); HA, hypopharyngeal apophysis; J, promotor of mandible; J, remotor of mandible; k, median tendon of mandibular adductors of dumb-bell muscle (KL or KLk); KLk, fibers of mandibular adductors united by tendon (k); KLt, fibers of mandibular adductors retaining origin on hypopharyngeal apophyses (D, HA); m, suspensory tendons of mandibular adductors; Md, mandible; PcR, posterior cranial ridge; tf, branch of labral muscle attached on mandible. decapods (Virbius, Spirontocaris). In Spirontocaris (fig. 27 B), the median ligament (k) of the dumb-bell adductors (KL) is connected with the hypodermis of the dorsal wall of the body by a branched arm (m) on each side. As before pointed out, however, the adductor liga- 68 SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL. 81 ment is in no sense to be homologized with the tentorium as developed in some of the higher crustacea and in the pterygote insects. Each mandible of Spirontocaris is provided with two dorsal productor muscles (/), but a reductor was not observed. Spirontocaris preserves the primitive single dorsal point of articulation of the mandible (a) with the wall of its segment. In higher decapods, the amphipods, and the isopods, where the mandible may have a double hinge with the . wall of the head, the musculature of the organ is modified in a manner Hy to be described later. ne The simple mechanism of the mandible of the higher pterygote in- sects is well understood ; the complicated musculature of the mandible in Apterygota has been given scant attention, and the derivation of the pterygote jaw mechanism from that of the Apterygota has been almost ignored. Borner (1909) has given the first comparative account of the mandibular musculature in the more generalized insects, and has pointed out certain points of similarity with the musculature of higher crustaceans. He did not, however, carry his comparisons to the myriapods, and thereby missed some fundamental relations. The mandibles of the Machilidae will serve best as an example of the more generalized apterygote jaw. The mandible of JMachilis or of Nesomachilis (fig. 27D, Md) is surprisingly similar in form to that of the crustacean Apus (A), except that it has a long incisor point in addition to a broad molar iobe. In this latter character the machilid jaw resembles the mandibles of some of the decapod crus- taceans, such as Sprirotocaris and Virbius, as has been pointed out by Crampton (1921b). The mandible of Machilis is suspended by a single dorsal point of articulation (a) against the lateral wall of the head. The cavity of the elongate base of each organ 1s filled by a mass of muscle fibers (AK Lk), and these fibers from the two mandibles con- verge upon the ends of a common transverse tendon () that passes through the base of the hypopharynx. Here, in an insect, therefore, i we find the same type of dumb-bell adductor uniting the two mandibles | as occurs in the Diplopoda and in lower Crustacea. In Machilis, however, there is a second and larger set of adductor fibers (KL?) vi which has its origin on the hypopharyngeal apodemes (HA). Machi- iH) lis, therefore, in the possession of two differentiated sets of mandibular i adductor fibers, combines the primitive condition of the chilopods Ki with the specialized condition of the diplopods and lower crustaceans. The tergal musculature of the mandible in Machilis is simple, con- sisting of an anterior promotor (J) and a posterior remotor (J). The two muscles are disposed exactly as in Apus (A), and are in entire NO. 3 INSECT HEAD—SNODGRASS 69 conformity with the tergal musculature of the basal plate of the jaw of Scutigera (fig. 26 B, C, J, J) and other chilopods. The machilid type of mandibular musculature appears to be char- acteristic of most apterygote insects except the Lepismatidae. In Japyx and Campodea, the bases of the elongate mandibles and maxillae are deeply retracted into the head above the labium, and the edges of the labium are fused to the postgenal margins of the head, so that the distal edge of the labium appears as the ventral lip of a pouch containing the other gnathal appendages and the hypopharynx. The mandibles of Heterojapyx (fig. 27 C, Md) are simple, slender organs, each consisting of a long, hollow basal piece, and of a more strongly chitinized free terminal lobe with a toothed incisor edge. The proximal tapering end of each jaw is set off from the rest by a thick internal ridge, superficially suggesting the division of the maxillary base into cardo and stipes ; but the “ mandible gives rigidity instead of flexibility. The two mandibles of Heterojapyx are connected by a large dumb-bell adductor muscle (KLk), the spreading fibers of which fill the basal cavities of the organs. Besides this muscle there are also sets of ventral fibers (K Lt) to the mandible that arises on the hypopharyngeal apophyses. The tergal muscles of the mandibles are large: they include for each jaw an anterior muscle (/) arising against a dorsal cranial ridge (PcR), and a wide fan of posterior fibers (J) arising along a median coronal ridge. Because of the retraction of the mouth appendages, the hypo- pharyngeal muscles of the mandibles (AK Lt) would appear to function as protractors, and the tergal muscles as retractors ; but the former are clearly the hypopharyngeal adductors of Machilis (D, KLt), and the latter the tergal promotors (J) and remotors (J). A _ peculiarity noted in Heterojapyx, if the writer observed correctly, is the attach- ment of a branch of the retractor of the labrum (ft) on the base of the mandible. In the Collembola, which also have retracted mandibles and max- illae, the mandibular musculature would appear, from Folsom’s (1899) account of Orchesclla cincta, to be of the same essential nature as that of Japyx. Folsom enumerates ten muscles for each mandible of Orchesella, but they all fall into three groups according to their origins, namely, muscles arising on the walls of the head, muscles arising on the “tentorium’”’ (hypopharyngeal apodemes), and fibers from one mandible to the other. The second and third groups constitute the adductors of the jaw; their fibers are inserted, Folsom says, on the inside of the lateral wall of the mandible, and most of them have their origin on the “ tentorium,” but a few of the division’ in the Japyx 70 SMITHSONIAN MISCELLANEOUS COLLECTIONS voL. 81 fibers, he adds, ‘‘ pass under the tentorium and become continuous with similar fibers from the opposite mandible.” Folsom, it will be noticed, says the adductor fibers connecting the mandibles pass be- neath the tentorial arms. In Japyx the tendon of the dumb-bell muscle distinctly lies dorsal to the hypopharyngeal apodemes. In Machilis the apodemes are so loosely connected with the base of the hypo- pharynx and so strongly united with the lateral inflections of the head wall, that in dissections their hypopharyngeal connections are easily lost, and the impression is given that the ‘tendon of the dumb-bell 1Ant Mx 2Mx A Fic. 28.—Head of Gammarus locusta (amphipod crustacean). A, lateral view of head, showing tergal abductors (J) and adductors (J) of left mandible, and base of ventral adductor (KL). B, postero-ventral view of back of head, showing origin of ventral adductors (KL) on posterior tentorial bar (PT). TAnt, first antenna; 2Ant, second antenna; J, abductor of mandible; J, dorsal adductor of mandible; AL, ventral adductor of mandible; Lm, labrum; Md, mandible; M/dP/p, mandibular palpus; 1M, first maxilla; 2M, second maxilla ; iMxp, first maxilliped; pt, posterior tentorial pit; PT, transverse posterior tentorial bar. adductor lies ventral to the apodemes. It does lie ventral to the sus- pensorial plates uniting the apodemes with the lateral walls of the head, but it passes anterior, 7. e., dorsal, to the ends of the apodemes themselves. Folsom’s statement, therefore, should be verified, for a discrepancy in the relations of the parts in question seems hardly permissible if we are dealing with homologous structures. The mandibles of the Protura, as described by Berlese (1909), are provided each with retractors and protractors that have their origins on the head wall, and with a protractor arising on the tentorial apodeme. Berlese, however, does not mention a muscle continuous between the two mandibles. The muscles present clearly represent the usual tergal muscles, and the hypopharyngeal adductor. NO. 3 INSECT HEAD—-SNODGRASS al In all the apterygote forms thus far described, the mandible has a free attachment to the head, being implanted by most of its length in the ventro-lateral membranous part of the head wall, and articulated to the margin of the chitinous cranium by only a single dorsal point of contact. In the Lepismatidae, a new condition is established in the mandible through the elongation of its dorsal base line forward and ventrally to the anterior end of the lower genal margin of the epicra- nium. The jaw thus becomes hinged to the head on a long basal axis extending from the primitive dorsal articulation, which is now pos- terior, to the angle between the genal margin of the head and the clypeus. At the latter point a secondary, anterior articulation is established between the mandible and the cranium. Borner (1909) describes the articulation of the mandible of Lepisma, but he does not observe that its type of structure is characteristic of the Lepis- matidae only, not of the Apterygota in general. The alteration in the mandibular articulation involves a change in the entire mechanism of the jaw, and initiates the series of modifications that have led to the evolution of the pterygote type of mandibular musculature from that of Machilis, Japyx, and the Collembola. The musculature of the mandible of Lepisma, as described by Borner (1909), is apparently almost the same as that of Machilis. The adductor muscles inserted within the body of the mandible consist of a large dorsal set of fibers (fig. 29 B, KLt) from the tentorium representing the fibers that arise on the hypopharyngeal apodeme of Machilis (figs. 27 D, 29 A, KLt), and of a small ventral set (KLh) arising directly from the hypopharynx. The tergal muscles comprise a pair of abductors (/) inserted on the outer margin of the mandibular base between the two articular points (a, c), and a large dorsal ad- ductor (J) inserted on the inner margin mesad of the posterior artic- ulation. The tergal abductors and adductor, however, are clearly the promotor and the remotor of the mandible of Machilis (fig. 29 A, I, J) and of all other generalized forms, which have assumed a new function by reason of the change in the nature of the mandibular articulation. The structure and musculature of the mandible in nymphs of Ephe- merida is essentially the same as in Lepisma. Borner describes and figures the nymph of Cloéon dipterum, showing the presence of a large tentorial adductor and a very small hypopharyngeal adductor, in addition to the dorsal abductors and adductors; the writer has verified the existence of all these muscles in another ephemerid species. In a dragonfly nymph, Aeschna, a small hypopharyngeal adductor was found, but no tentorial fibers were observed. In the 72 SMITHSONIAN MISCELLANEOUS COLLECTIONS voL. 81 orthopteron, Locusta, Borner shows two small tentorial adductors of the mandible (fig. 29 C, KLt), and a small hypopharyngeal adductor (KLh). The same muscles the writer has found in Microcentrum, the hypopharyngeal fibers being attached medially on the tips of the rudimentary suspensorial arms of the hypopharynx (fig. 20 D, KLh) ; but no trace of either set could be discovered in the acridid, Dissos- teira. Mangan (1908) described in the roach, Periplancta australasiae, both a tentorial adductor and a hypopharyngeal adductor. The first A B Fic. 20.—Three stages in the evolution of the mandibular mechanism in biting insects. A, mandible of Machilis, outer surface, with single dorsal point of articulation (a) with cranium; the jaw moved by tergal promotor (/) and remotor (J), and by ventral adductors (KLk, KLt, see fig. 27 D). B, mandible of Lepisma (from Borner, 1909), articulated with cranium on long basal hinge inclined downward anteriorly from dorsal articulation (a) to anterior articulation (c); the promotors (/) here become abductors, and the remotor (J) becomes a tergal adductor; ventral adductor (KLh, KLt) retained. C, head of Locusta (from Borner, 1909), showing common type of mandibular articulation in pterygote insects, with hinge line inclined downward posteriorly from anterior articulation (c) to posterior articulation (a); tergal abductors and adductors (J, /) highly developed, ventral adductors (KLh, KLt) rudi- mentary. In higher Pterygota the ventral adductors disappear. mention of either of these muscles is by Basch (1865), who found the tentorial adductor. in the mandible of Termes flavipes. The adductor fibers arising directly from the base of the hypo- pharynx are evidently remnants of the primitive sternal adductors that have retained their original connections. Jn the insects, therefore, the primary, sternal adductor muscles (KL) of the mandibles have become differentiated into three groups of fibers, the fibers of one group (KLh) retaining the primitive sternal connection, those of the second (KLt) being carried inward upon the sternal (hy popharyngeal ) apophyses, those of the third (KLk), after having united medially NO. 3 INSECT HEAD—-SNODGRASS 73 with the corresponding set from the opposite mandible, having been detached from all connections except their points of insertion on the mandibles. With the change in the mandibular articulation from a single dorsal suspensory point to a long basal hinge, the primary ad- ductors have lost their importance, and the function of adduction has been secondarily taken over by the primary tergal remotor, while the original tergal promotor becomes the abductor. Remnants of the primary adductors in insects having a hinged mandible persist in the Lepismatidae, Ephemerida, Orthoptera, and Isoptera, but in the higher orders they have disappeared. A still further evolution in the mandibular base has reversed the tilt of the hinge line. Instead of sloping from the posterior articulation downward and forward, as it does in Lepisma and in some ephemerid nymphs, the base of the jaw in all higher insects is inclined from the anterior articulation downward and posteriorly (fig. 29 C). This change in the slope of the axis of the hinge causes the apex of the jaw to swing inward and posteriorly during adduction, instead of in- ward and anteriorly as in the first condition. In the higher decapod crustaceans, and in the amphipods and iso- pods, the mandible has undergone an evolution parallel to that which has taken place in insects. Borner (1909) has described the mandible and mandibular musculature of Gammarus, an amphipod, and has shown the structural similarity with the mandible of Lepisma. In Gammarus locusta (fig. 28 A) the mandible is hinged to the cranium by its long base, which slopes downward and forward from the pos- terior point of articulation. The primitive tergal promotor muscle (7) has then become an abductor, and the remotor (J) has become a dorsal adductor. The primitive ventral adductor (KL) has its origin on a well-developed transverse tentorial bar (B, PT) passing through the back of the head; a hypopharyngeal branch of the adductor is lacking. In the crayfish (Astacus), Schmidt (1915) describes an anterior ventral adductor of the mandible arising on the anterior end of the ventral head apodeme. In the isopods the mandible attains a stage almost exactly comparable with that of the higher pterygote insects (fig. 17 F)—the basal hinge line of the jaw slopes posteriorly and downward, and the only muscles present, so far as the writer could find, are the tergal abductors and adductors. The homologues of the mandibles in Xiphosura and Arachnida, the so-called pedipalps (fig. 17 J, Pdp), scarcely need consideration here. The pedipalps never attain a jaw-like form, but retain always the structure of a jointed limb, though the basal segment may develop a strong gnathal lobe. 74 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 81 THE FIRST MAXILLAE The leading features of the first maxilla have been sufficiently noted in the description of a generalized gnathal appendage (page 60) based on the maxilla of Periplaneta (fig. 25). In none of the other arthropods are the maxillary appendages so highly developed as in the insects, but, in all the arthropods, it appears that the mandible has been evolved from an appendage that was originally very similar Fic. 30—Maxilla of Heterojapyx gallardi. A, left maxilla, posterior (ventral) surface. B, right maxilla and muscles, anterior (dorsal) view. Cd, cardo; e, articulation of cardo with cranium; fga, flexor of galea; flcc, cranial flexor of lacinia; flcs, stipital flexor of lacinia; Ga, galea; HA, hypo- pharyngeal apophysis; HS, rudiment of suspensorial arm of hypopharynx; /, promotor of cardo; KLcd, adductors of cardo; KLst, adductors of stipes; Le, lacinia; OO, muscle of base of palpus; », muscle of terminal segment of palpus; PcR, posterior cranial ridge; Plp, palpus; rplp, first segment of palpus; St, stipes; , line of internal ridge of stipes. to the generalized insect maxilla. In many of the higher insects the maxillae, too, have become specialized, always in adaptation to special modes of feeding, but a description of the modifications involved is beyond the scope of the present paper. The musculature of the organ is essentially the same in all groups of biting insects, except as it suffers a reduction where the appendages become reduced or united with the labium. The maxilla of Japyx (fig. 30) presents a more generalized con- dition in its relation to the head than does the maxilla of the roach, NO. 3 INSECT HEAD—SNODGRASS ai in that the head apophyses (B, HA) upon which the adductor muscles of the appendages arise are still connected with the hypopharynx, whereas in Periplaneta the corresponding endoskeletal arms have lost their primitive sternal connections and have become a part of the tentorium. The adductors of the cardo in Heterojapyx (fig. 30 B, KLcd) are well difterentiated from those of the stipes (KLst), and cross obliquely the inner ends of the latter. The promotor of the cardo (/) arises against a median ridge of the dorsal wall of the cranium. The lacinia (Lc), which is mostly covered dorsally by the galea, has a broad flexor arising within the stipes (fics), and a large cranial muscle (fcc) arising against the dorsal cranial ridge (PcR) on the top of the head, and going dorsal to the other muscles of the appendage to be inserted on a slender apodeme from the inner angle of the lacinial base. The galea (Ga) is provided with a single long flexor (figs. 30 B, 31 D, fga) arising within the stipes, which splits into two bundles of fibers toward its insertion on the ventral wall of the base of the galea. The palpus (P/p) is reduced and otherwise modified as compared with that of the roach (fig. 25), consisting of only three segments, of which the basal one (figs. 30 A, 31 D, rplp) is much elongate and is united with the outer wall of the base of the galea (Ga). There might be some question as to the homology of this basal region of the palpus of Japyx, but the insertion upon its base of the muscle (OQ) from the stipes, evidently representing the usual pair of palpal muscles, and the origin within it of a muscle (p~) going to the distal segment of the palpus identify the part in question as the true basal segment of the palpus. The cardo and the stipes of many insects appear externally to be divided into sub-sclerites, but in most such cases it is found that the so-called “ sutures” are but the external lines of inflections that have formed internal ridges, the ridges being developed either for giving strength to the sclerite, or to furnish special surfaces for muscle attachment. The cardo of Periplaneta, for example (fig. 25 A, Cd), has a “divided” appearance externally, but when examined from within (B) it is seen that the regions apparent on the surface result from the presence of a strong Y-shaped ridge (7) on the inner wall, which extends distally from the base to reinforce with its diverging arms the extremities of the hinge line with the stipes. This structure of the cardo is characteristic of other orthopteroid insects. Crampton (1916) distinguishes the area of the cardo between the arms of the ridge as the “‘ veracardo,” and calls the rest of the sclerite the “ juxta- cardo.” The terms may have a descriptive convenience, but they are misleading if taken to signify a division of the cardo into two parts. 76 SMITHSONIAN MISCELLANEOUS COLLECTIONS voL. 8I The stipes is usually marked by a prominent groove parallel to its inner edge (fig. 25 A, q), setting off a narrow marginal strip. The groove is here likewise but the external line of an internal ridge or plate upon which are inserted the adductor muscles of the stipes | (B, KLst). Crampton designates the area of the stipes external to / the ridge as the ‘‘ verastipes,” and that mesad to it as the “ juxta- ae stipes.” In Heterojapyx the basal part of the stipes is strengthened by an internal ridge (fig. 30 A, w) that forks proximally to the ends of the hinge line with the cardo. ‘ lic. 31—Maxillae of insects and of a chilopod. A, maxilla of Nesomachilis. B, maxilla of Thermobia (Lepismatidae). C, maxilla of larva of Sialis. D, terminal part of maxilla of Heterojapyx. §, maxilla of an adult stonefly (Pteronarcys). F, first maxillae of a chilopod (Lithobius). Base of palpus to be identified by insertions of levator and depressor muscles (O, QO); the palpifer (P/f) has no muscles, and appears as a mere subdivision of stipes; in Sialis larva (C), lobe o is not the galea, but an endite of first segment of palpus (7plp), the latter identified by its muscles (O, QO). The ventral, or distal, end of the stipes bears the lacinia and galea, . and to its lateral surface is attached the palpus. The lacinia and galea are movable lobes, each being provided with muscles having their origin in the stipes, by which they can be flexed posteriorly (or ven- trally if the mouth appendages are horizontal). The lacinia, in ad- dition, has a muscle from the cranial wall inserted on the inner angle of its base, which gives it a mesal flection, or adduction. The base of the galea commonly overlaps anteriorly the base of the lacinia. The maxillary palpus arises from the outer wall of the stipes, usually only a short distance proximal to the base of the galea. The NO. 3 INSECT HEAD SNODGRASS Te area supporting the palpus is frequently differentiated from the rest of the stipes, and is then distinguished as the palpifer (fig. 31 A, Plf). When the delimiting suture of the palpifer region extends to the galea, the palpifer appears to support both the galea and the palpus. That the palpifer is not a segment of the appendage is shown by the fact that muscles neither arise within it nor are inserted upon it. The muscles that move the palpus as a whole have their origins within the main part of the stipes, and always pass through the pal- pifer, if the latter is present, to be inserted on the proximal segment of the palpus (figs. 25 C, 31 A-E, O, Q). The palpus muscles, then, may be taken as identification marks of the true basal segment of the palpus. Since they are typically inserted one dorsally and the other ventrally, relative to the vertical axis of the appendage, they are evidently a levator (O) and a depressor (Q) of the palpus. The number of segments in the maxillary palpus varies much in different insects. Machilis perhaps presents the maximum number of seven (fig. 31 A): the palpus of the roach with five segments is more typical (fig. 25). Evidence will later be given indicating that the palpus is the telopodite of the maxillary appendage, and that its basal articula- tion with the stipes, or palpifer, is the coxo-trochanteral joint of a more generalized limb (fig. 35 A, B, C, ct). A joint near the middle of the palpus (figs. 25 C,35 A, B. ft) often suggests the femero- tibial flexure. THE SECOND MAXILLAE The second maxillae of insects are unquestionably united in the labium. The correspondence in external relations between the parts of each half of a typical labial appendage and those of an entire maxilla is so close that most entomologists have not hesitated to assume an homology of the submentum (figs. 32 A, 40 D, Sim) with the cardines, of the mentum (J/t) with the stipites, of the glossae (G/) with the laciniae, and of the paraglossal (Pg/) with the galeae. Some writers, however, have contended that the submentum, or both the submentum and the mentum represent the sternum of the labial segment. Thus, Crampton in a recent paper (1928) adopts the idea of Holmgren (1909) that the submentum and mentum are derived from the sternum of the labial segment. In an orthopteroid labium (fig. 40 D), the muscles of the palpi (28, 29), and the muscles of the terminal lobes (25) arise in the mentum (//t), and this relation, together with the presence of muscles from the mentum to the tentorium (23, 24), must certainly identify the region of the mentum in the labium with that of the stipes in a 78 SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL. 81 first maxilla (fig. 40 B, St). The wall of the mental region, however, may not be entirely or continuously chitinized (fig. 32 A), and, hence, a distinction must be drawn between the entire region of the mentum, and the area occupied by one or more mental sclerites. The labium may contain muscles not represented in the maxillae, such as the muscles associated with the orifice of the salivary duct in the grass- C= hopper (fig. 40 D, 20, 27), or with the silk press in the caterpillar (fhigs.'53.C, Dps4. AaB: C).7 7.406. 60s The submentum corresponds functionally at least with the cardines of the maxillae, since it serves to attach the labial appendage to the walls of the head. The lateral articulation of its basal angles to the Fic. 32.—Second maxillae. A, typical second maxillae of an insect (Periplaneta) united to form the labium. B, second maxillae of a chilopod (Lithobius) united by inner angles of coxae. ct, coxo-trochanteral joint; Cx, coxa; Gl, glossa; Mt, mentum; O, levator muscle of telopodite (palpus) ; Pgl, paraglossa; Plg, palpiger; Plp, palpus; Q, depressor muscle of telopodite (palpus) ; Smt, submentum. margins of the cranium in orthopteroid insects (figs. 18 B, C, 36 C, f) suggests, moreover, that the points of attachment are the true basal articulations of the second maxillae with the cranium, corre- sponding with the articulations of the cardines (e¢) in the first maxillae. It is possible, of course, that a median part of the labial sternum has been incorporated into the submentum. To accept the proposal, how- ever, that the entire submentum is the sternum of the labial segment. is to assume that the sternum itself has become articulated laterally to the tergum of its segment, and that it alone bears the segmental appendages. Such assumed relations violate the basic principles of segmental morphology, and thus throw suspicion on the evidence given in their support. a ee Fs | NO. 3 INSECT HEAD—SNODGRASS 79 It will be shown in the next section of this paper that the cardines of the maxillae are not true proximal segments of the maxillary appendages, but are secondary subdivisions of the bases of these appendages. It appears probable, therefore, that the submentum represents likewise proximal subdivisions of the bases of the second maxillae, retaining the lateral articulations with the margins of the cranium in generalized insects (fig. 36C,f), and perhaps including between them a median part of the labial sternum. If the insect labium (figs. 32 A, 40 D) is compared with the second maxillae of a chilopod (fig. 32 B), it will be seen that the united basal segments of the latter (Cx), containing the origins of the palpal muscles (O, Q), correspond at least with the mentum of the labium. The large proximal segments of the chilopod maxillae are clearly the bases of a generalized limb, the coxae, according to Heymons (1901), and the limb base, or a subcoxal division of it, bears the primitive dorsal articulation of the appendage with the body. The mentum, and at least the lateral parts of the submentum, therefore, appear to be subdivisions of the primary bases of the second maxillary ap- pendages, corresponding with the stipites and cardines of the first maxillae in insects, and with the similar subdivisions of the bases of the mandibles in the diplopods (fig. 26 A, Cd, St). The median, terminal duct of the labial, or “ salivary,” glands opens anterior to the labium, and, in typical forms, at the base of the mentum (figs. 18 D, 19, SIO). The position of the orifice, anterior to the sub- mentum, however, does not argue that the latter is entirely the sternum of the labial segment, but rather the reverse, for it is likely that the orifice of the duct has not left the sternal region of its segment, and that it has been crowded forward in the latter by the median approach of the labial cardines. The common duct of the labial glands results during embryonic development from the union of the two primary ducts of paired lateral glands of the labial segment. ’ MORPHOLOGY OF THE GNATHAL APPENDAGES It has often been assumed that the segmental appendages of all arthropods are derived from a primitive limb having a biramous type of structure. A two-branched limb, however, occurs actually only in the Crustacea, and there is no certain evidence of a biramous limb structure ever having prevailed in other arthropod groups. In all forms, including the Crustacea, the segmental appendages first appear in the embryo as simple protuberances of the body wall, and some zoologists now believe that the exopodite branch, when present, is 6 SO SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. SI merely a specially developed exite lobe of a single shaft. Borradaile (1917) expresses the opinion that “ probably the primitive crustacean appendage resembled that of the Branchiopoda in being uniramous.”’ Movable lobes individually provided with muscles, however, may be developed along both the outer and the inner margin of the limb, and an excessive development of one of the outer lobes might give rise to Fic. 33.—Parapodium and parapodial musculature of an annelid worm (Nereis virens). A, B, first and third parapodia, left, anterior surfaces. C, cross section of left half of a segment from middle of body, cut anterior to base of parapodium, showing muscles of setae inserted on end of setal pouch (a), and ventral promotor (K) and remotor (L) muscles of parapodium. DMcl, VMcl, dorsal and ventral bands of longitudinal body muscles. D, musculature of third parapodium, right, inner view, showing tergal pro- motor (/) and remotor (J), and sternal promotor (AK) and remotor (L). E, musculature of right side of a segment from middle of body, internal view, lateral oblique muscles and setal muscles removed: b, c, anterior and pos- terior pleuro-sternal muscles; DJ/cl, part of dorsal longitudinal muscles ; /, tergal promotor of parapodium; J, tergal remotor; j, accessory remotor arising anteriorly from intersegmental fold; A, sternal promotor; L, sternal remotor ; Set, bases of setae. a secondary biramous structure of the appendage. Hansen (1925) recognizes the definitive two-branched structure of the typical crus- tacean appendage, but he says it seems “ impossible to deny the possi- bility that the exopod may be analogous with the epipod, and if so the primitive appendage is uniramous.” The segmental appendages, or parapodia, of the polychaete annelids are in some cases simple lobes; in others they are of a two-branched D3 INSECT HEAD—-SNODGRASS SI structure owing to the presence of two groups of setae on each (fig. 33 B,C). In Nereis virens, though most of the parapodia are dis- tinctly cleft, those of the first and second segments do not have the double structure (fig. 33 A). Whatever relations, however, may be traced, or assumed to exist, between the annelids and the arthropods, the relationship must be presumed to have come through a remote worm-like ancestor common to both groups, for none of the highly organized modern annelids can be taken to represent the ancestral form of the arthropods. A comparative study of the legs of mandibulate arthropods will show that in each group there is a maximum of seven limb segments, beyond a subcoxal base, that are individually provided with muscles. The relative size and form of the segments, the character of the articu- lations, and the nature of the musculature present many variations, and it is not to be assumed that segments are to be homologized in all cases by their numerical order beyond the base of the limb. The gnathal appendages undoubtedly constitute a group of organs that are individually homologous in arthropod groups, whether their segments are united with the protocephalon to form a larger head, or with the body segments following. The similarity of the structure of the mandible in all the eugnathate arthropods, and the common plan of its musculature, allowing for modifications of which the evolution can easily be followed, leave no doubt concerning the identity of the jaw in the various groups, or that the jaw attained its basic structure in some very remote common ancestor. The primitive struc- ture of the mandible is not entirely preserved in any arthropod: in the Diplopoda and Chilopoda the movable lacinia is retained, but the palpus has been lost ; in the Crustacea and Hexapoda, the lacinia has lost its independent mobility and has become solidly fused with the base of the appendage, but in many crustaceans a mandibular palpus persists. The first maxilla of the Hexapoda has the structure of a generalized mandible, 7. ¢., it consists of a base supporting a palpus and at least one movable lobe, the lacinia, though generally there is present a second lobe, the galea. The insect labium consists of a pair of ap- pendages that probably once had the structure of the first maxillae. In the Chilopoda the maxillary appendages appear to have under- gone but little modification of structure, and those of the second pair still retain a form similar to that of the body appendages. The corresponding appendages of the Diplopoda are now so highly spe- cialized that it is useless to speculate as to their earlier form. In the Crustacea both pairs of maxillae have been reduced in size and modified i ; i | | } ; 82 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL: OF in structure to serve as organs accessory to the mandibles, but they have not attained the highly specialized form of the corresponding appendages of insects. We may conclude, therefore, that in the common ancestor of the several groups of modern eugnathate arthropods, the mandible alone had attained a gnathal function, and that in form and structure it re- sembled the maxilla of a present day insect, though perhaps lacking a galea, or outer endite lobe of the base. The two maxillae at this period were more or less modified to serve as organs accessory to the mandibles. When the modern groups of arthropods were differentiated, the mandible, in the Diplopoda and Chilopoda, retained the movable lacinia, but lost the palpus ; in the Crustacea and Hexapoda, the lacinia fused with the base of the appendage to form a solid jaw, while the palpus was preserved by the crustaceans, and lost by the insects. The two maxillary appendages retained the leg-like form in the Chilopoda ; in the Diplopoda they became highly specialized in a manner peculiar to the diplopod group; in the Crustacea and Hexapoda they were modified for an accessory gnathal function, but in the insects they ac- quired a form almost identical with that of a primitive mandible. Finally, in the insects, the second maxillae united basally to form the labium. While the insect maxilla appears to be a highly specialized appendage, it will be shown that its basic structure is not far removed from that of a thoracic leg. While the status of the gnathal appendages relative to one another in the various groups of the eugnathate arthropods seems fairly clear, it is a more difficult matter to homologize their parts with the segments of the ambulatory appendages. The structure of the first and second maxillae of a chilopod, or of the first maxilla of an insect, suggests that the gnathal appendages have been derived from an appendage of the ambulatory type—the insect maxilla is certainly more like the leg of an insect, a chilopod, or a decapod than it is like one of the body appendages of Apus (fig. 35 C), or of any other of the lower crustaceans in which the appendages are used for swimming. This condition suggests, therefore, that the ambulatory leg more nearly represents the primitive type of arthropod limb than does an appen- dage, such as that of Apus, clearly modified for purposes of purely aquatic locomotion. If we consider, furthermore, that the appendages of the chelicerate arthropods (Xiphosura and Arachnida) are also of the ambulatory type, the evidence becomes all the more convincing that the primitive arthropod limb was a walking leg and not a swim- ming organ. If this deduction is acceptable, we must conclude that NO. 3 INSECT HEAD—SNODGRASS 83 the Crustacea represent a group of arthropods that have secondarily adopted an aquatic life, and that, while certain forms have become thoroughly adapted to a free life in the water, others have retained, with but little modification, some of the organs that were developed primarily for terrestrial locomotion. This view, perhaps, is contrary to generally accepted ideas concerning the evolution of the arthropods, but it is clearly futile to attempt to derive the appendages of arth- ropods in general from the swimming appendages of Crustacea. If the ambulatory limb be taken as more nearly representative of the basic structure of an arthropod appendage than is the natatory Fic. 34.—Generalized segmentation and musculature of an insect leg, diagram- matic. A, theoretical segmentation and musculature of a primitive arthropod leg, anterior view: the appendage, consisting of a limb base (LB), and a telopodite (Tip) of two segments, moved forward and backward on vertical basal axis et tergal and sternal promotors (/, AK), and tergal and sternal remotors (Cie). D, definitive segmentation of an insect leg by division of limb base (A, LB) into subcoxa (Scx) and coxa (Cx), and by subsegmentation of first part of telopodite into trochanter and femur, and of second part into tibia, tarsus, and praetarsus. a-b, basal axis of limb base; ct, coxo-trochanteral joint; Cx, coxa; F, femur ; ft, femoro-tibial joint; J, tergal promotor; J, tergal remotor; K, sternal pro- motor; L, sternal remotor; O, levator of telopodite; P, tergal depressor of telo- podite (characteristic of insects) ; Ptar, pretarsus; Q, depressor of telopodite ; Scx, subcoxa ; T, depressor of distal segment of telopodite; Tar, tarsus; T), tibia. limb, we have only to inquire as to what was its probable form in the ancestors of terrestrial arthropods. The primitive appendage un- doubtedly turned forward and backward on a vertical axis through its base (fig. 34 A, a, b), as does the parapodium of a modern polychaete annelid (fig. 33 A, B,C). For walking purposes, however, the limb must have acquired joints, and, as Borner (1921) has shown, the simplest practical condition would demand at least two joints with vertical movements (fig. 34 A), one near the union of the leg with the body (ct), dividing the limb into a basal piece (LB) and a telopodite i i | ( 84 SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL. 81 (Tlp), and one (ft) near the middle of the telopodite. These primary joints persist, evidently, as the coxo-trochanteral joint (B, ct) and the femero-tibial joint (ft) of the leg (Hiiftgelenk and Kniegelenk, ac- cording to Borner). The type of leg-segmentation resulting from two joints so placed applies at least to the Chilopoda, Diplopoda, Hexapoda, and Crustacea ; in the Xiphosura and Arachnida, however, it is possible that the mechanism of the leg is given by three primary joints, the second and third setting off a horizontal middle section of the leg (patella). The further segmentation of the limb has been produced by the subdivision of the principal parts of the telopodite. In the mandibu- late arthropods (fig. 34 B), one or two small segments cut off from the basal end of the proximal piece of the telopodite form the tro- chanters (Tr), while the rest of this part becomes the femur (F) ; the distal section beyond the knee joint (ft) subsegments into the tibia (7b), tarsus (Tar), and praetarsus (Ptar). This type of seg- mentation is clearly shown also in the maxillipeds or in any of the anterior body appendages of Apus. In the third maxilliped (fig. 35 C) there are two principal flexures, one (ct) between the limb base (LB) and the telopodite (7/p), the other (ft) beyond the middle of the latter. The part between the two points of flexure is the femur (F) with two indistinctly separated trochanters (77) ; that beyond consists of two shortened segments, and the terminal praetarsus, or dactylopodite. The limb base of Apus is entire, but in some arthropods the basis appears to have become subdivided into a coxa (fig. 34 B, Cx) and a subcoxa (Scx). The coxa may then become the free functional base of the appendage, since the subcoxa usually forms a chitinization in the pleural or the sternal wall of the segment. The primitive musculature of the limb base was such as to swing the appendage forward and backward ; it must have comprised, there- fore, promotor and remotor muscles. Probably there was a tergal promotor (fig. 34 A,/) and a tergal remotor (J), and a sternal pro- motor (K) and a sternal remotor (L). In a thoracic leg of an insect, the base of the telopodite is provided with a depressor muscle (P) having its origin on the tergum of the segment, which greatly increases the lifting power of the appendage, but this muscle is not to be con- sidered as a primitive element of the limb musculature. The usual levator and depressor muscles of the telopodite (O, Q) have their origin within the limb base. The simple type of musculature shown in figure 34 A, and here assumed to be the primitive musculature of an arthropod limb base, is actually present in typical form in the simpler anterior parapodia NO. 3 INSECT HEAD—SNODGRASS 85 of the annelid, Nereis virens (fg. 33 D). Here a dorsal promotor and a remotor (J, J) arise on the tergal wall of the segment, and a ventral promotor and a remotor (K, L) on the midline of the sternal wall. The ventral muscles are repeated regularly in all the segments of the worm (C, E, K, L), but in the more posterior segments the dorsal muscles, though present (FE, J, J), are less symmetrical in ar- rangement, and the primary remotor (J) is subordinated to a large oblique remotor (j) that arises on the anterior margin of the seg- ment. This last muscle is described by Borner (1921) as being the typical dorsal remotor of the parapodium, but by comparison with the simpler anterior appendages (D) it appears to be a secondary acquisi- tion, for the muscle (E, J) dipping beneath it has the same insertion on the parapodial base as that of the tergal promotor of the anterior parapodium (D, J). Borner’s claim, however, that this simple type of limb musculature presented by Nereis must represent the primitive motor mechanism of an appendage turning forward and backward on a vertical axis through its base is scarcely to be questioned. The basal muscles of an appendage do not necessarily retain their original functions, nor their primitive simplicity, for an alteration in the basal articulation of the appendage may change the fundamental movements of the limb, and thereby give quite a different action to the muscles, which, in turn, may shift in position, or become split up into segregated groups of fibers, thus multiplying the number of in- dividual muscles actually present. Returning now to a further consideration of the muscles of the gnathal appendages of the arthropods, it is not difficult to draw a parallel between the musculature of a mandible, or of an insect maxilla, and that of the annelid parapodium (fig. 33 D, EF), or with the hypopthetically primitive musculature of an arthropod limb base as expressed in figure 34 A. In the mandible of Scutigera (fig. 26 B, C), Apus (fig. 27 A), Heterojapyx (fig. 27 C), Nesomachilis (fig. 27 D), a tergal promotor (/) and a remotor (J) have the typical re- lation to the appendage. In some forms the tergal remotor appears to be lacking (Diplopoda, fig. 26 A; Spirontocaris, fig: 27 B). The cranial flexor of the mandibular lacinia in diplopods and chilopods (fig. 26 A, B, C, flcc) is probably derived from the tergal promotor, since it arises on the dorsal wall of the head and goes dorsal (an- terior) to the ventral muscles. The sternal promotor and remotor, which are distinct muscles in the annelid parapodium (fig. 33 C, D, EB; K, L), are united in the gnathal appendages of the arthropods, where they become ventral adductors (KL) asa result of the free movement ~~ ee 86 SMITHSONIAN MISCELLANEOUS COLLECTIONS voL. 8&1 of the base of the appendage on a single dorsal point of articula- tion (a). The adductor fibers of the mandible may all retain their connection with the sternal, or hypopharyngeal, apophyses (Chilopoda, fig. 21 B), or they may become detached from the apophyses and united with the fibers from the opposite jaw to form a transverse dumb-bell muscle (Diplopoda, fig. 26, A, KL; some Crustacea, fig. 27 A, B; most Apterygota, fig. 27 C, D, KLk), though at the same time some of the fibers may retain their connections with the apophyses, or with the tentorium (Apterygota, fig. 27 C, D, KLt; Orthoptera, fig. 29 C, KLt) or with the base of the hypopharynx (Lepisma, fig. 29 B, KLh; Locusta, fig. 29 C, KLh; Microcentrum, fig. 20D, KLh; ephemerid nymph, fig. 20 A, KLh). The evolution of the mandibular muscles in the higher insects has been detailed in an earlier paragraph, wherein it was shown that the ventral adductors are reduced and finally obliterated after the jaw has acquired a double hinge with the edge of the cranium, and that the tergal promotor and remotor muscles then become respec- tively the functional abductors and adductors. _ The basal musculature of the insect maxilla, as already shown, coincides almost exactly with the basal musculature of the mandible of a diplopod or a chilopod, and may be derived from the simple plan of the musculature of the annelid parapodium. The tergal promotor is evidently separated into two groups of fibers inserted on the dorsal and ventral extremities of the anterior rim of the appendage base, the upper set being the muscle of the cardo (figs. 25 C, 30 B, J), the lower set the cranial flexor of the lacinia (fcc). A tergal remotor is lacking in the insect maxilla, but so it appears to be also in the diplopod mandible (fig. 26 A). The sternal promotor and remotor muscles (figs. 33 D, E, 35 B, K, L) are united, as in the mandible, to form a ventral adductor (KL), the fibers of which almost always retain their origin on the hypopharyngeal apophyses, or on the corresponding part of the tentorium, and are distributed to both the cardo and the stipes (figs. 25 C, 30 B, KLced, KLst). In Machilis the maxillary adductors from opposite appendages are united with each other medially, and appear to be detached from the ventral apophyses. The margin of the basal cavity of the maxilla (fig. 35 A) includes the region of the cardo, the stipes, and the lacinia; and the tergal and sternal muscles (/, J, KL) of the appendage are distributed to these three parts. The entire base of the maxilla, therefore, has the fundamental character of a single segment, and there can be no doubt that this segment is the true primitive base of the appendage (fig. 34 A, LB). The base of a leg appendage may be divided into a coxa a3 INSECT HEAD—SNODGRASS 87 and a subcoxa (fig. 34 B, Cv, Scx), and Borner (1909, 1921) would identify the cardo of the maxilla with the subcoxa of a leg. The suture separating the cardo from the stipes, however, terminates at both ends in the marginal rim of the maxillary base (fig. 35 A) instead of running parallel with it; the cardo, therefore, does not have the relation of a true segment to the rest of the appendage. It is perhaps, Fic. 35.—The relation of a maxilla to a generalized limb. A, theoretical generalized structure of a gnathal appendage, consisting of a limb base (LB), bearing a divided basendite (Lc, Ga), and a six-segmented telopodite, or palpus (Pip), with the principal downward flexure at the femoro- tibial joint (ft). The limb base provided with tergal promotor and remotor muscles (J, J), and sternal adductors (KL). B, showing basal musculature of a gnathal appendage analysed into the functional elements of the musculature of an annelid par apodium (fig. 33 D). C, third maxilliped of Apus longicaudata, left, anterior surface, showing division into a limb base (LB) and a telopodite (Tip) ; the base movable on a transverse axis (a-b), the telopodite with a principal flexure (ft) between its third and fourth segments. a-b, basal axis of limb base; Be, basendite; ct, coxo-trochanteral joint; F, femur ; fga, flexor of galea; flcc, cranial flexor of lacinia; flcs, stipital flexor of lacinia; ft, femoro-tibial joint; Ga, galea; J, tergal promotor; J, tergal remotor; K, sternal promotor; KL, ventral adductors (K and L united); L, sternal remotor; LB, limb basis; Lc, lacinia; O, levator of telopodite; Plp, palpus (telopodite) ; QO, depressor of telopodite; T/p, telopodite; Tr, trochanters. though, to be questioned if the subcoxal chitinization, the pleuron, at the base of a thoracic leg is not also a mere subdivision of the basal limb segment similar to the cardo, rather than the remains of a true independent segment. If the cardo does, in any sense, represent the thoracic subcoxa, it is to be noted that the hinge line between it and the stipes has a horizontal position with reference to the axis of the appendage, and this, the writer has argued (1927), must have been the primitive position of the subcoxo-coxal hinge in a thoracic leg. Se. 88 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 81 Though the homology of the cardo must be left in question, the writer would agree with Borner (1909, 1921) that the part of the maxilla bearing the lacinia, galea, and palpus represents the coxal re- gion of the leg base, and that the basal segment of the palpus is a tro- chanter (fig. 35 A, Tv). The lacinia and galea, then, are coxal endites, and, as Borner proposes, the stipes and palpifer are corresponding sec- ondary subdivisions of the coxa, or of the coxal region of the maxillary base. In the maxillae and maxillipeds of the Crustacea, Borner claims, the segments bearing the lobes homologous with the insect lacinia and galea are also subdivisions of the coxa. By this interpre- tation, the palpus (fig. 35 A, Plp) becomes the telopodite of the maxil- lary appendage (B) ; its basal union with the stipes or palpifer is the coxo-trochanteral joint (ct), and its principal distal articulation hav- ing a ventral flexure is the femoro-tibial joint (ft). Other writers have held somewhat divergent views concerning the homologies of the maxillary segments. Goldi (1913) interpreted the cardo as the coxopodite, the stipes as the basipodite, to which he assigned the lacinia and galea as endite lobes, and the basal segment of the palpus as the ischiopodite. Crampton (1922) gave a modifica- tion of this view in that he proposed that the palpifer represents a segment, the ischiopodite, and that the galea is an endite of this seg- ment. Uzel (1897) appears to give confirmation to this view in his description of the development of the maxiila of Campodea; the maxil- lary rudiment, he says, is first divided into an outer and an inner lobe, and then the outer lobe splits into two parts, one of which becomes the palpus, the other the galea. If the maxilla of Campodea resembles that of Japyx (fig. 30 A), however, it is easy to believe that the structure in the embryo might be misinterpreted, in as much as the adult structure is misleading until the muscle relations are taken into consideration (fig. 31 D) ; then it is seen that the basal region of the palpus, which is united with the base of the galea, is the true basal segment of the palp, and not the palpifer—clearly a secondary modi- fication. It has already been pointed out that the entire lack of muscle con- nections in the palpifer is a condition that disavows the segmental nature of the palpifer region. Crampton’s best example among in- sects of a structure corresponding with his idea of the segmentation of a maxillary appendage is the maxilla of the larva of Sialis (fig. 31 C), in which there is a small lobe (0) borne on the apparent first seg- ment of the palpus. This lobe Crampton would identify as the galea, making the supporting segment the palpifer. The muscles (O, Q) in- serted on the base of this segment, however, clearly demonstrate that it NO. 3 INSECT HEAD—SNODGRASS 89 is the true base of the palpus (1plp)—therefore, not the palpifer, which lacks muscles—and that the lobe in question is not the galea, as also the absence of a muscle to it would indicate. The maxilla of the Sialis larva, then, is not a generalized appendage in the sense that Crampton would infer, since it lacks a true galea and is provided with an accessory lobe on the first segment of the palpus. Similar lobes of the palpus segments occur in other insects, particularly in larvae of Coleoptera; they have a suggestion of the endite lobes of the telo- podite in such crustaceans as Apus (fig. 35 C). In the thoracic legs the limb is always flexible at the union between the basis and the telopodite, 7. ¢., at the coxo-trochanteral joint, and in no appendage, where the facts can be clearly demonstrated, is there a union between the coxa and the trochanter. It does not seem reasonable, therefore, to suppose that the proximal segment of the telopodite (the trochanter) should have been incorporated into the limb base in the case of a maxillary appendage. Especially is such a supposition unreasonable in the face of much specific evidence to the contrary. The whole body of evidence bearing on the limb mech- anism points to a primitive uniformity of flexure in all the appen- dages, whereby the limb is divided into a basis and a telopodite. and indicates that the articulation between these two parts is preserved in the entire series of appendages, except, of course, where the telo- podite is lost. The maxillipeds and the anterior body appendages of Apus bear each five endite lobes. The first lobe (fig. 35 C, Be) is a basendite, the second is carried by the proximal division of the trochanter, the third by the femur, the fourth by the tibia, and the fifth by the tarsus. Each endite is independently movable by muscles inserted upon or within its base. The maxillae of Apus are reduced to single lobes, but the first maxilla appears to represent the rudimentary limb base with the large basendite, since it falls exactly in line with the series of basal endites on the following appendages. The basal endites of arthropod limbs in general, including the ‘‘ gnathobases ” of the trilo- bite appendages, the gnathal lobes of the pedipalps in Xiphosura and Arachnida, and the “ styli”’ of the legs of Scolopendrella, are almost certainly analogous lobes in all cases, and they must be represented by the laciniae (at least) of the insect maxillae, by the laciniae of the mandibles of diplopods and chilopods, and by the incisor and molar lobes of the mandibles in all arthropods. It is a question, therefore. whether the galea of the insect maxilla is an accessory lobe of the limb base, or a subdivision of the primary basendite (fig. 35 B, C, Be) The latter seems probable, since, in the more generalized insects, the go SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL. 81 lacinia and galea overlap each other basally, and both are flexed by the muscles inserted upon their bases that have their origin in the stipes. It is impossible at present to arrive at final conclusions on the many problems connected with the morphology of arthropod appendages, and the most that the writer would claim for the present attempt at advancing the subject is that the material here presented gives at least a substantial enlargement to the foundation of known facts from which future work must proceed. There is no question that students of arthropods have given far too little attention to the relationship between skeletal structure and musculature. The more the subject is looked into, the more it will be seen that the characters of the arthropod skeleton are in large part adaptations to the strain of muscle tension, and that they are to be correctly interpreted only through an under- standing of the entire mechanism of which they are a part. The sclerites of the insect cuticula, in particular, have been studied as if they were skeletal elements deftly fitted together in such a manner as to cover the outside of the animal, and we entomologists have played with them, as we might with the sections of a picture puzzle, without looking for their significance in the mechanics of the insect. The arthropod skeleton, it is true, has been formed from a few major centers of increased chitinization, but the minor “ divisions”’ are in almost all cases adaptations to flexion, or the opposite, namely, the strengthening of the skeleton by the development of internal ridges. The scientific study of the comparative anatomy of insects must look for its advance in the future to a wider knowledge of muscles and mechanism. IV. SUMMARY OF IMPORTANT POINTS 1. The arthropods have been derived from creeping animals, not from forms specially modified for swimming; their immediate pro- genitors were annelid-like in structure. .2. The stomodeum marks the anterior end of the blastopore. There are, therefore, no true mesodermal segments anterior to the mouth. The unsegmented preoral part of the animal is the prostomium, and constitutes the most primitive head, or archicephalon, of segmented animals, since it contains the first nerve center, or “ archicerebrum,” and bears the primitive sense organs. 3. The first stage in the development of a composite head in the arthropods, as represented in the embryo, comprises the prostomium and the first two or three postoral segments. The head in this stage may be termed the protocephalon; it is represented by the cephalic lobes of the embryo, which may or may not include the third segment. NO. 3 INSECT HEAD—SNODGRASS gl 4. The protocephalon carried the labrum, the mouth, the eyes, the preantennae, and the antennae, also the postantennae when it included the third body segment. 5. During the protocephalic stage of insects, as shown by the em- bryo, the thorax was differentiated as a locomotor center of the body, and the region between the head and the thorax, consisting of the fourth, fifth, and sixth body segments, became a distinct gnathal region. 6. The gnathal region was eventually added to the protocephalon to form the definitve head, or telocephalon. In the Crustacea, in which there was no thoracic region corresponding with that of the insects, the gnathal region was not definitely limited posteriorly, and the definitive head in this group may include as many as five segments following the protocephalon. In some of the crustaceans the gnathal segments have united with segments following to form a gnatho- thorax, leaving the protocephalon as a separate anterior head piece. In the Arachnida the protocephalon included the prostomium and two postoral metameres, and it has combined with the following six seg- ments to form the cephalothorax. 7. In the definitive insect head, the prostomium, according to some embryologists, contributes the clypeus and frons and the region of the compound eyes; according to others it forms the clypeus and frons only. The labrum is a median preoral lobe of the prostomium. 8. The arthropod brain probably always includes the median pro- stomial ganglion, combined with the ganglia of the preantennal segment to form the protocerebral lobes. It may still be questioned whether the optic lobes are derived from the prostomium or from the prean- tennal segment. The ganglia of the antennal segment form the deuto- cerebrum. The commissures of the protocerebrum and the deutocere- brum are formed above the stomodeum, and unite with the archicere- bral rudiment to form the median part of the brain. The ganglia of the postantennal segment, when united to the preceding ganglia, become the tritocerebral brain lobes, but they, remain separate from the brain in some crustaceans, and their uniting commissure always preserves its sub-stomodeal position. 9. The prostomial region of the adult arthropod is innervated from the postantennal ganglia, but this is probably a secondary condition owing to the loss of the true prostomial nerves. 10. The appendages of the definitive insect and myriapod head are the preantennae, the antennae, the postantennae, the mandibles, the first maxillae, and the second maxillae. Rudimentary, evanescent preantennae have been reported only in the embryo of Scolopendra Q2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 81 (Heymons) and in the embryo of Carausius (Wiesmann). Postan- tennae are commonly present in insect embryos, but their rudiments persist in only one or two doubtful cases in the adult. The postantennal appendages are the second antennae of Crustacea, and probably the chelicerae of Arachnida and Xiphosura. Endites of their bases may have been the functional jaws of the insectan and myriapodan an- cestors in the protocephalic stage. Ir. The gnathal appendages have been derived from organs having the structure of uniramous ambulatory legs. All the primitive ar- thropod appendages were probably uniramous ambulatory limbs. Bira- mous and natatory appendages are characteristic of the Crustacea only, and are probably secondary adaptations to an aquatic life. 12. The mandible is a common inheritance from an early ancestor of the eugnathate group of arthropods. Its primitive structure re- sembled that of the first maxilla of modern insects, and is best pre- served in the Myriapoda. 13. The diplopod mandible consists of a base subdivided into cardo and stipes, bearing a large movable lacinia, but lacking a galea and a palpus. In the typical chilopod mandible, the division between cardo and stipes has been lost, and the lacinia is less free. The musculature of the chilopod mandible is more primitive than that of the diplopod mandible. In the crustaceans and insects the mandibular lacinia is either lost, or is fused with the base to form a solid jaw. The mandib- ular palpus is retained in many Crustacea. The mandible is repre- sented by the pedipalp in Arachnida. 14. The first maxillary appendage is best developed in the insects, and probably here preserves the primitive structure of the mandible. Its musculature is exactly duplicated in the musculature of the man- dibles in the diplopods and chilopods. Neither the first nor the second maxillae of the chilopods gives any evidence of ever having at- tained the special structure of the primitive mandibles and the insect maxillae. 15. A primitive gnathal appendage had the structure of a gener- alized ambulatory appendage, consisting of a limb basis and a telo- podite. The basis represents the coxa and subcoxa of a thoracic leg, but its division into cardo and stipes is not a true segmentation. The galea and lacinia are movable endites of the basis, with the origin of their muscles in the stipital region of the latter. The telopodite becomes the palpus of the gnathal appendage, and its basal articula- tion is the homologue of the coxo-trochanteral joint in the leg. The palpifer is not a segment of the limb, but a subdivision of the stipes NO. 3 INSECT ILEAD—-SNODGRASS 93 bearing the palpus and the galea (as claimed by Borner) ; the muscles of the palpus and the galea pass through the palpifer, but never arise within it. 16. The primitive appendage was implanted in the soft lateral wall of its segment, and turned forward and backward on a vertical axis through its base, as does an annelid parapodium. The first joint set off the telopodite, and gave the latter a mobility in a vertical plane. 17. The primitive muscles inserted on the base of a generalized limb, as on an annelid parapodium, consisted of a dorsal promotor and a re- motor, and of a ventral promotor and a remotor. When tergal plates were developed, the gnathal appendages of the arthropods became attached to their lateral margins, each by single point of articulation. The ventral muscles of the appendages then became sternal adductors. 18. The points of origin of the ventral adductors of the gnathal appendages in myriapods and insects were probably crowded together when the gnathal segments were added to the protocephalon. They have since become supported on a pair of apophyses arising at first from the base of the hypopharynx. In the myriapods and in most of the apterygote insects, the apophyses still maintain their hypopharyn- geal connections ; but in the pterygote insects their bases have migrated laterally to the margins of the cranium, and in all but some of the lower forms have finally moved to a facial position in the epistomal suture. Their posterior ends have united with the transverse ten- torial bar developed in the back part of the head. The hypopharyngeal apophyses of the Myriapoda and Apterygota have thus come to be the anterior arms of the pterygote tentorium. 19. The adductor muscles of the insect maxillae, arising on the tentorium, are the sternal adductors of the appendages, corresponding with the sternal adductors or rotators of the thoracic legs, and are derived from the primitive ventral promotors and remotors of the limb. 20. The ventral adductors of the mandibles in the Chilopoda re- tain their connections with the sternal, or hypopharyngeal, apophyses. In the Diplopoda, Crustacea, and Apterygota, groups of the adductor fibers from the mandibles have lost their sternal connections and have united with each other by a median ligament to form a dumb-bell muscle between the two jaws. Other groups of fibers may retain their connections with the apophyses, or direct with the base of the hypo- pharynx. In the Pterygota, the ventral adductors of the mandibles have been lost, except for a few rudiments in some of the lower orders. 21. The mandible of Lepisma and of pterygote insects is hinged to the head on a long base line with anterior and posterior articulations. | | | " ( Q4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL. 81 The posterior articulation is the primitive one, the anterior a secon- dary one. By this change in the articulation and movement of the jaw, the primitive tergal promotor muscle becomes an abductor, and the primitive tergal remotor becomes an adductor. The base line of the mandible slopes downward and forward in Lepisma and in a few of the lower pterygotes; in all others its slope is reversed, allowing the tip of the jaw to swing inward and posteriorly during adduction. A similar evolution of the mandible has taken place in the Crustacea. 22. The ridge of the base of the insect cranium, on which the pro- thoracic and neck muscles are inserted, is probably a chitinization of the intersegmental fold between the maxillary and labial segments. The posterior tentorial arms arise from its ventro-lateral ends by in- vaginations in the external suture. The neck of the insect, therefore, may be unchitinized parts of both the labial and the prothoracic segments. V. THE HEAD OF A GRASSHOPPER After laying down the general principles worked out in the pre- ceding sections, it will be well to test them with a few specific examples. The head of a grasshopper is a good subject for an elementary study of the structure of the pterygote insect head, because it preserves the generalized orientation in having the face directed forward and the mouth appendages hanging downward. Terms of direction, therefore, do not have to be qualified—ventral is downward, dorsal is upward, and anterior is forward. The descriptions here given are based on the Carolina locust (Dissosteira carolina), a fairly large grasshopper to be obtained in almost any part of the United States. The muscles are designated numerically for convenience of refer- ence only, and the same numbers do not refer to corresponding muscles in the grasshopper and in the caterpillar (Section VII). The myology of insects is as yet too little advanced to furnish a satisfactory general nomenclature for insect muscles, and no attempt is made here to use a set of names for the muscles of the grasshopper that could in all cases be applied to the muscles of other insects. The usual method of naming muscles according to their function, or their supposed func- tion, gives terms fitting for the species described ; but in many cases, by a change in the articulation between the skeletal parts involved, muscles that are clearly homologous have their functions completely altered. Again, it is impossible to name muscles consistently according to their points of origin and insertion, for either end of a muscle may shift and may migrate into a territory quite foreign to its original NO. 3 INSECT HEAD—SNODGRASS 95 connections. A third feature disturbing to a uniform muscle nomen- clature is the fact that any muscle may break up into groups of fibers, or, at least, a single muscle in one species may be represented func- tionally by several muscles in another. Finally, there are muscles that are evidently new acquisitions developed in connection with special mechanisms. The importance of the study of musculature for the understanding of the insect skeleton, however, can not much longer be ignored. STRUCTURE OF THE CRANIUM The walls of the head in the grasshopper are continuously chitinized on the anterior, dorsal, and lateral surfaces (fig. 36 A, B), and the Ant Vx ocs Oc y2 \ \ J. _Poc Fic. 36—Head of a grasshopper, Dissosteira carolina, A, lateral. B, anterior. C, posterior. a, posterior articulation of mandible; Ant, antenna; at, anterior tentorial pit; c, anterior articulation of mandible; Clp, clypeus; cs, coronal suture; cv, cervical sclerites ; E, compound eye; e, articulation of maxilla with cranium; es, epistomal suture; f, articulation of labium with cranium; For, foramen magnum; fs, frontal suture; g, condyle of postocciput for articulation with cervical sclerite ; Ge, gena; h, subocular ridge; 7, frontal carina; j, subantennal suture; , flexible area between lower edge of gena and base of mandible; Lb, labium; Lm, labrum; Md, mandible; Mx, maxilla; O, ocellus; Oc, occiput; ocs, occipital suture; Pge, postgena; Poc, postocciput; PoR, postoccipital ridge; pos, postoccipital suture; PT, posterior arm of tentorium; /f, posterior tentorial pit; sgs, subgenal suture; Vr, vertex. dorsal and lateral walls are reflected upon the posterior surface (C) to form a narrow occipito-postgenal area surrounding the foramen magnum (For). The foramen is closed below by the neck membrane, in which is suspended the base of the labium (Lb). The ventral wall of the head, between the bases of the gnathal appendages, is occupied mostly by the large median hypopharynx, it being otherwise reduced to the narrow membranous areas between the lateral margins of the hypopharynx and the bases of the mandibles and maxillae. Zi go SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 8&1 The facial aspect of the cranium is distinctly separated by the epistomal suture (fig. 36 A, B, es) from the clypeus, but there is no demarked frontal sclerite. The apex of the frons, however, is defined in Dissosteira by two short remnants of the frontal sutures (B, fs) diverging from the end of the coronal suture (cs). The facial area of the head is limited on each side by an impressed line (/) extending from the lower angle of the eye to the anterior articulation of the mandible. The median part of this area forms a broad frontal costa, margined laterally by a pair of sinuous carinae (7) reaching from the top of the head to the lower part of the face. A short, transverse subantennal suture (7) lies on each side of the frontal costa just below the level of the median ocellus. The inner ridges of these subantennal sutures have a close relation to the attachments of the more important muscles of the frons (fig. 38 D, 7). The true frontal area of the grasshopper, therefore, must include the region of these sutures and extend dorsal to them between the bases of the antennae into the angle between the short remnants of the frontal sutures. The lateral areas of the head (fig. 36 A) have no special character- istics. The subgenal suture (sgs) on each side is continuous anteriorly with the epistomal suture (es). The compound eye is surrounded by a distinct suture forming a high ridge internally (fig. 39 A, OR), and setting off a narrow rim, or ocular sclerite, around the base of the eye (fie. 36;-4,,B,C). On the posterior surface of the head (fig. 36C), the occipito- postgenal area (Oc, Pge) is included between the well-marked occip- ital suture (ocs) and the postoccipital suture (pos). In Dissosteira the occiput and postgenae are continuous ; in Melanoplus the occipital arch is separated from the postgenae by a short groove on each side on a level with the lower angles of the compound eyes. Posterior to the postoccipital suture is the postoccipital rim of the head (Poc), widened above and below on each side, to which the membrane of the neck is attached. The postoccipital suture forms internally the ridge on which the muscles of the neck and prothorax that move the head are inserted (fig. 45 A, Pok). Laterally the postoccipital ridge is elevated as a high plate (fig. 36 C, PoR), from the ventral ends of which the posterior arms of the tentorium (PT) proceed inward. The roots of the tentorial arms appear externally as long open slits in the lower ends of the postoccipital suture (Pt, pt). The clypeus and labrum form together a broad free flap (fig. 36 A, B, Clp, Lm) hanging before the mandibles from the lower edge of the frontal region. The fronto-clypeal, or epistomal, suture (es) is a deep groove forming internally a strong epistomal ridge (fig. 39 A, NO. 3 INSECT HEAD—-SNODGRASS 97 B, C, ER), from the lateral parts of which arise the anterior tentorial arms (AT). The roots of these arms appear externally as lateral slits in the epistomal suture (figs. 36 B, 37 A, at, at), just mesad to the anterior articulations of the mandibles (c, c). The clypeus of Dissosteira is partially divided by transverse lateral grooves into anteclypeal and postclypeal areas. The labrum is a broad oval plate, notched at the middle of its ventral margin, freely movable on the lower edge of the clypeus. A median quadrate area on its basal half is limited below by a sinuous transverse groove (fig. 37 A) that forms a low ridge on the inner surface of the anterior wall (B, J.) On the Fic. 37.—Clypeus and labrum of Dissostetra carolina. A, anterior surface. B, posterior surface of anterior wall, showing muscle attachments, and bases of anterior tentorial arms. AT, anterior arm of tentorium; at, anterior tentorial pit; c, anterior articula- tion of mandible; C/p, clypeus; es, epistomal suture; /, ridge of anterior labral wall; Lm, labrum; Tor, torma; 1, labral compressors; 2, anterior retractors of labrum; 3, posterior retractors of labrum; 34, 35, points of origin of anterior dilators of buccal cavity (figs. 41, 44). posterior surface of the clypeo-labral lobe, the area of the clypeus is separated from that of the labrum by two small chitinous bars, the tormae (figs. 37 B, 42 A, Tor). A median Y-shaped thickening (fig. 42 A, m) of the cuticula of the labrum makes a ridge on the inner surface of the posterior labral wall. The clypeus has no muscles for its own movement, but the first two pairs of anterior dilators of the buccal cavity (figs. 41, 44, 33, 3 /) are inserted on the inner surface of its anterior wall (fig. 37 B, 33, 34). The labrum is provided with three sets of muscles, as follows: 1.—Compressors of the labrum (fig. 37 B).—A pair of short mus- cles arising medially on transverse ridge of anterior labral wall (J) ; diverging to arms of Y-shaped ridge in posterior wall (fig. 42 A, m). 98 SMITHSONIAN MISCELLANEOUS COLLECTIONS voL. 81 2.—Anterior retractors of the labrum (fig. 37 B).—A pair of long muscles arising on subantennal ridges of frons (fig. 38 D, 7) ; con- verging downward to insertions on base of anterior wall of labrum (fig. 37 B). 3.—Posterior retractors of the labrum (fig. 37 B).—A pair of long muscles arising on subantennal ridges of frons, each laterad of 2 (fig. 38 D) ; inserted on dorsal processes of tormae at base of posterior wall of labrum (figs. 37 B, 42 A). The articulations of the gnathal appendages occupy typical positions along the lower lateral margins of the cranium. The mandible articu- lates anteriorly with a condyle (fig. 39 A, c) supported at the junction of the epistomal and subgenal ridges (ER, Sg), but projecting on the external surface of the head. Posteriorly the jaw articulates with a facet on the ventral edge of the postgena (figs. 36 C, 39 A, C, a). This articulation as the first is outside the membranous connection of the mandible with the head. The axis of the mandible slopes strongly downward and posteriorly between the articular points. The lateral edge of the mandibular base is separated from the margin of the gena by a narrow, flexible strip of weakly chitinized articular membrane (fig. 36 A, k), at the ventral margin of which arises the abductor apodeme of the mandible (fig. 39 D, § Ap). The maxilla articulates by a single point on the base of the cardo with a shallow facet on the edge of the postgena (figs. 36 C, 40 C, e) almost directly below the posterior tentorial pit (pt). The maxillary articulation is thus crowded unusually far posteriorly in the grass- hopper. In most generalized insects it lies well before the line of the postoccipital suture, as in a roach or a termite, and is often much farther forward. The labium is loosely articulated by the elongate basal angles of the submentum with the posterior margin of the postocciput at points a short distance above the posterior lower angles of the latter (figs. 36 CG, Ao. Ge 7): The tentorium of the grasshopper has the form of an X-shaped brace between the lower angles of the cranial wall (fig. 39 B). The anterior arms (AT) arise from the lateral parts of the epistomal ridge (ER), their broad bases extending from points above the mandibular articulations half way to the median line of the face. In this respect Dissosteira shows an advance over Periplaneta, in which the bases of the anterior tentorial arms arise from the subgenal ridges and extend only a short distance mesad of the mandibular articulations. ‘The posterior tentorial arms of Dissosteira (fig. 39 B, PT) arise from the lower ends of the postoccipital ridge (fig. 45 A, PoR). The median ee INSECT HEAD——-SNODGRASS 99 body of the tentorium is concave below (fig. 39 B, C, Tnt). A thin, flat dorsal arm of the tentorium (fig. 39 C, DT) arises from the base of the inner end of each anterior arm and extends upward and anter- iorly to the wall of the cranium just before the lower angle of the compound eye. The dorsal tentorial arms are attached to the hypo- dermis of the head wall, but make no connection with the cuticula in Dissosteira. THE ANTENNAE Each antenna consists of two larger basal segments, and of a long slender flagellum broken up into about 24 small subsegments. In Fic. 38—Antenna of Dissosteira carolina and of Periplaneta. A, base of left antenna of Dissosteira, ventral surface. B, the same of Periplaneta. C, base of right antenna and antennal muscles of Dissosteira, dorsal view. D, base of right antenna, antennal muscles, anterior tentorial arm, muscles arising on frons, and associated structures of Dissosteira, interior view. 5Ap, apodeme of depressor muscles of antenna; A/X, antennal ridge; At, anterior arm of tentorium; c, anterior articulation of mandible; DT, dorsal arm of tentorium; ER, epistomal ridge; es, epistomal suture; j, subantennal ridge; n, pivot of antenna; OR, ocular ridge; Pdc, pedicel; Scp, scape. Dissosteira the antenna of the male is a little longer than that of the female. Of the two basal segments, the proximal one, or scape (fig. 38 A, Scp), is the larger. It is articulated to the rim of the antennal socket by a small process on the lateral ventral angle of its base that touches upon the margin of the socket (7). The motion of the scape on the head, however, is that of a hinge joint moving in a vertical plane on a transverse axis. The base of the scape is provided 100 SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL. &1 with levator and depressor muscles (C, D). In other insects the antenna is more commonly pivoted on a ventral point of articulation with the rim of the socket, as in Periplaneta (fig. 38 B, n), and thus has greater freedom of movement. As already noted, however, the artic- ular point may be dorsal, as in Japyx, and in the Chilopoda (fig. 23 B, n). The thickened rim of the antennal socket (fig. 38 D, AR) is braced by a short arm against the anterior margin of the heavy cir- cumocular ridge (OR). A crescentic area of the head wall just above and mesad to the antennal socket is depressed externally, and the inflection tilts the place of the antennal socket somewhat dorsally, giving the antenna a more upward play than it otherwise would have. ‘The second basal segment of the antenna, the pedicel (fig. 38 A, B, Pdc), is movable in a horizontal plane on the end of the scape by means of muscles arising within the scape. The other segments of the antenna are flexible but have no muscles. The muscles of the antenna comprise muscles inserted on the base of the scape that move the antenna as a whole, and the muscles of the pedicel that move the pedicel and flagellum. They are as follows: 4.—Levators of the antenna (fig. 38 C, D)—Two muscles arising on tentorium, one (D, 4a) on dorsal arm, the other (4b) on anterior arm; both inserted by a short tendon on a lobe of dorsal side of base or scape (C, D): 5.—Depressors of the antenna (fig. 38 C, D).—Two muscles aris- ing on dorsal arm of tentorium (D, 5a) and on anterior arm (5)) ; both inserted on a long slender tendon arising near ventral margin of scape (A, 54/) in articular membrane of antenna. 6.—Extensor of the flagellum (fig. 38 C).—Arises dorsally and medially in base of scape; inserted medially on base of pedicel. 7.—Flexor of the flagellum (fig. 38 C).—Arises dorsally and later- ally in base of scape; inserted laterally on base of pedicel. THE MANDIBLES The mandible of the grasshopper is a strongly chitinous jaw—a short, hollow appendage with triangular base, thinning down to the cutting margin. The anterior and the posterior angles of the lateral base line carry the articular points with the head, and the apodeme of the adductor muscles arise at the median angle. The distal edge of each mandible presents an incisor and a molar area. The first (fig. 39 D, 0) forms the compressed and toothed apical part of the jaw, the second (p) forms a broad grinding surface on the anterior median face closer to the base of the mandible. The in- cisor and molar areas are not exactly alike on the two jaws, each being NO. 3 INSECT HWEAD—SNODGRASS LOL best developed on the right. The molar area of the right mandible consists of strong, heavy ridges forming a projecting surface; the ridges of the left jaw are low and their area does not project. The two molar surfaces, therefore, fit one upon the other without interfer- ence when the jaws are closed. The incisor lobes of the mandibles close upon the ventral end of the hypopharynx, the molar surfaces over its base, and the anterior contour of the hypopharynx is modeled 9a \ Pe PT a Fic. 39.—Internal structure of the head of Dissosteira carolina, and the mandible and its muscles. A, inner surface of right half of epicranium. B, tentorium and lower margin of epicranium, ventral view. C, inner view of right half of head, with right mandible and its muscles in place. D, right mandible, postero-mesal view. a, posterior articulation of mandible; 8Ap, abductor apodeme of mandible; 9Ap, adductor apodeme of mandible; AR, antennal ridge: AT, anterior tentorial arm; at, anterior tentorial pit; c, anterior articulation of mandible; Clp, clypeus ; cv, cervical sclerite; DT, dorsal tentorial arm; E, compound eye; ER, epistomal ridge; es, epistomal suture; Fr, frons; g, condyle of articulation of cervical sclerite; h, subocular ridge; j, subantennal ridge; Lm, labrum; Md, mandible; O, ocellus; 0, incisor lobe of mandible; », molar area of mandible; Poc, post- occiput; PoR, postoccipital ridge; P7, posterior tentorial arm; pt, posterior tentorial pit; SgR, subgenal ridge; Smt, submentum; 7 nt, body of tentorium. according to the irregularities of the mandibular surfaces. The pos- terior slope of the mandibular hinge lines cause the points of the jaws to turn inward, upward, and posteriorly during adduction. At the base of each molar area of the mandibles a flat brush of hairs (fig. 39 C, D) projects inward, and the two brushes come together anterior to the mouth opening when the mandibles are closed, serving thus evidently to prevent the escape of masticated food material from be- tween the jaws. The anterior surfaces of the mandibles are overlapped by the epipharyngeal surface of the clypeus and labrum, and the 102 SMITHSONIAN MISCELLANEOUS COLLECTIONS voL. &1 asymmetry of the mandibular surfaces and contours is reflected in that of the epipharyngeal surface (fig. 42 A). The mandibles of Dissosteira are moved, so far as the writer could discover, only by tergal abductor and adductor muscles, which, as al- ready explained, are the primitive tergal promotors and remotors transformed in function by the change from a monocondylic to a dicondylic articulation in the mandible (fig. 29 A, B, C). Small ventral adductors of the mandibles arising on the hypopharynx and on the tentorium persist in some of the Tettigontidae (figs. 20 D, KLh, 29 C, KLh, KLt), but these muscles appear to be lost in the Acrididae, as they are in all higher pterygote insects. The fibers of the functional abductors and adductors arise on the walls of the cranium and are inserted on flat apodemal plates of the jaws. The abductor apodeme is a small plate (fig. 39 D, SAP) arising from the articular membrane close to the outer margin of the mandibular base and near the posterior articulation (a). The adductor apodeme (QAP) con- sists of two large thin plates borne upon a common stalk, which arises from the articular membrane at the inner angle of the mandib- ular base, and lies in the lateral angle between the anterior and pos- terior arms of the tentorium (C). One plate extends dorsally in a longitudinal plane, the other, which is smaller, lies in a transverse plane. Each mandibular apodeme is a chitinous invagination from the articular membrane close to the base of the jaw. The muscles of the mandible correspond with the apodemes. They are as follows: 8.—Abductor of the mandible—A small fan of fibers, arising on ventral part of postgena and on extreme posterior part of ventral half of gena; inserted on abductor apodeme of the mandible. 9.—Adductors of the mandible (fig. 39 C).—Two sets of fibers corresponding with the two divisions of the adductor apodeme. The fibers of one set (Qa) arise on dorsal wall of cranium, from a point between compound eyes to occiput, with one bundle attached on post- occiput (Poc) ; inserted on both sides and on posterior margin of the median apodemal plate. Those of the other set (9D), inserted on the transverse plate of the apodeme, arise on lateral walls of cranium from subocular ridge (h) to postgena, and some of the posterior fibers encroach upon outer end of posterior tentorial arm. THE MAXILLAE The maxilla of the grasshopper (fig. 40 A) is so similar to that of the roach (fig. 25 A), already described, that its major features will need no special description. It consists of a triangular cardo (fig. 40 A, Cd), a quadrate stipes (St), with a well-developed palpifer INSECT HEAD—SNODGRASS 103 # 50 92 5S Sl fy Y Yi St Fic. 40.—Maxilla and labium of Dissosteira carolina. A, right maxilla, posterior surface. B, left maxilla, anterior view, exposing muscles of cardo and stipes. C, posterior region of cranium, with cervical sclerites and maxilla, left side. D, labium and its muscles, posterior view. E; stipes and palpifer with bases of palpus, galea and lacinia, lacinial muscles re- moved, anterior view. a, posterior articulation of mandible; 10Ap, apodeme of promotor of cardo: Cd, cardo; cv, cervical sclerite; e, articulation of maxilla with cranium; f, articulation of labium with cranium; Ga, galea; Gl, glossa; Le, lacinia; Mt mentum; Mx, maxilla; Oc, occiput; ocs, occipital suture; Pge, postgena ; elf palpifer; Plg, palpiger; Plp, palpus ; Poc, postocciput ; pos, postoccipital suture ; pt, posterior tentorial pit; g, suture and internal ridge near inner margin of stipes; r, internal ridge of cardo; s, apophysis of cardo for muscle insertion ; SID, salivary duct; Smt, submentum; Sf, stipes; t, suture and internal ridge separating palpifer from stipes; Tt, body of tentorium: uw, inner ridge at base of posterior wall of galea; v, keel of salivary cup. + l : i i i | { ——E ee ee eee ee ee EE Oe ——— 1O4 SMITHSONIAN MISCELLANEOUS COLLECTIONS voL. 81 (PIf), and two terminal lobes, lacinia (Lc) and galea (Ga), and a five-segmented palpus (P/p). The cardo presents an irregular topography on its external surface, and is marked into several areas by the lines of a strong branching ridge on its internal surface (fig. 40 B,r). Crampton (1916) calls the part proximal and posterior to the ridge the juxtacardo and the rest of the sclerite the veracardo, but the inference that these areas , are “ divisions ” of the cardo is scarcely warranted, since the ridge is clearly a mere strengthening device. The articular point (e¢) of the cardo with the cranial margin is a knob on the posterior angle of its base, anterior to which is a long arm to which is attached the apodeme (10Ap) of the promotor muscle (C, ro). A pit in the distal part of the external surface of the cardo (A, s) marks the site of an internal process on which one of the adductor muscles is inserted (B, z7a). The distal margin of the cardo is articulated by a long, flexible hinge line with the base of the stipes, but there are no muscles extending between the cardo and stipes. The quadrate stipes (fig. 40 A, St) has a strong plate-like ridge on its internal surface near the inner margin (q), on which is in- serted one of the adductor muscles (E, 12). Crampton distinguishes the body of the stipes as the verastipes, and the flange mesad of the muscle-bearing ridge as the juxtastipes. The region of the palpifer (A, PIf) is well separated from that of the stipes by an internal ridge (E, t), but the muscles of the palpus (17, 1S), as well as the muscle of the galea (76), have their origin in the stipes, suggesting that the palpifer is a subdivision of the stipes, and not a basal segment of the palpus. The lacinia (fig. 40 A, B, Lc) is borne by the distal end of the stipes, and is capable of flexion anteriorly and posteriorly on an oblique axis with the latter. Distally it tapers and ends in two claws turned in- ward. The lacinia is flexed by a pair of strong muscles arising within the stipes (B, 75a, 15b), and by a slender muscle (774) having its origin on the wall of the cranium. The galea (fig. 40 A, Ga) is carried by a distal subdivision of the palpifer, which Crampton (1916) calls the basigalea. In form, the galea (A, B, C, Ga) is an oval, flattened lobe ; its walls are but weakly chitinized. Its inner margin lies against the lacinia, and its outer surface is modeled to fit the outer part of the posterior surface of the mandible, against which it can be tightly closed. The base of the galea is marked on the posterior wall by an internal ridge (EF, «), upon which is inserted its single flexor muscle (70). NO. 3 INSECT HEAD—SNODGRASS 105 The maxillary palpus consists of five segments (fig. 40 A, B, C, Pip). The basal segment is provided with levator and depressor mus- cles (B, FE, 77, 18) arising within the stipes; each of the other seg- ments has a single muscle arising in the first or second segment proxi- mal to it. The muscles of a maxilla are as follows: 10.—Promotor of the cardo (fig. 40C)—A small fan of fibers arising on lower posterior part of postgena, external and anterior to the mandibular abductor ; inserted on slender apodeme of basal arm of cardo. 11.—Adductors of the cardo (fig. 40 B).—Two muscles arising on posterior end of anterior arm of tentorium, extending ventrally, pos- teriorly, and outward; one (za) inserted on process (s) of inner face of cardo, the other (zzb) mesad to distal end of ridge (r) of cardo. 12.—Proximal adductor of the stipes (fig. 40 E).—Arising on ex- treme posterior end of anterior arm of tentorium; inserted on ridge of inner margin of stipes. 13.—Distal adductors of the stipes (fig. 40 E).—\ Two muscles arising on tentorium, the first (73a) a slender muscle arising, along with rra, r1b, and 12, on posterior end of anterior arm of tentorium, the second (13) a large, thick, digastric muscle arising laterally on concave ventral surface of body of tentorium; both muscles inserted on a slender apodeme attached to inner distal angle of stipes. Muscles rz, 12, and 13 correspond with the adductors of the cardo and stipes that in Apterygota arise on the hypopharyngeal apodemes (fig. 30 B, KLcd, KL st), representing the sternal adductors, or ster- nal promotor and remotor, of a primitive appendage (fig. 35 B, K, L). 14.—Cranial flexor of the lacinia (fig. 40 B).—Arises on gena just before upper end of promotor of cardo (C, 10) ; inserted on inner angle of base of lacinia. This muscle is the homologue of the cranial flexor of the maxillary lacinia in Apterygota (fig. 30 B, fcc), and of the corresponding flexor of the mandibular lacinia in Myriapoda (fig. BoA, B; C, fcc). 15.—Stipital flexor of the lacinia (fig. 40 B)—A large two- branched muscle arising in base of stipes, one branch (15a) medially, the other (75b) in outer basal angle ; both inserted on anterior margin of base of lacinia. These muscles flex the lacinia forward. Berlese (1909) describes the posterior branch of this muscle in Acridium as attached to the posterior wall of the lacinia, and as being an antagonist to the anterior branch, but in no insect has the writer observed an antagonist to the lacinial flexor. : e@ 106 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 8I 16.—Flexor of the galea (fig. 40 B, -).—A large muscle arising mesally in base of stipes, external to lacinial muscles and depressor of palpus ; inserted posteriorly on ridge (EF, ) at base of galea. This muscle probably flexes the galea forward and inward, the point of flex- ion being at the base of the subgalea. 17.—Levator of the maxillary palpus (fig. 40 B, E).—Origin in median basal part of stipes ; insertion on dorsal margin of basal seg- ment of palpus. 18.—Depressor of the maxiilary palpus (fig. 40 B, E).—Origin on inner edge of stipes; crosses anterior to muscle of galea (16) to in- sertion on ventral margin of basal segment of palpus. If the basal segment of the palpus (fig. 35 A) corresponds with the trochanter of the leg (fig. 34 B, Tr), then muscles 77 and 18 represent the levator and depressor of the telopodite (fig. 34 A, O, OQ) arising in the coxal region of the leg base (LB). 19, 20, 21, 22—Muscles of the maxillary palpus (fig. 40 B).—A single muscle for each segment, the first (79) a levator of second seg- ment, the second (20) a productor of third segment, the third (27) a depressor (adductor) of fourth segment, the fourth (22) a reductor of terminal segment. The joint between the third and fourth segments of the palpus apparently represents the femero-tibial flexure of a leg (figs. 34, 35 A, ft), the two small basal segments of the palpus being trochanters. THE LABIUM The labium of the grasshopper (fig. 40 D) is simple in construction, and typical of the labium of biting insects, except in the reduction of the gloss. It consists of a large;submentum (Smt) with the elongate basal angles loosely attached to the posterior margin of the cranium behind the roots of the posterior tentorial arms (C, f). The mentum (D, Mt) is broad, with imperfectly differentiated palpus-bearing lobes, or palpigers (P/g), at the sides of its base. On its ventral margin the mentum bears a pair of large flat lobes, the paraglossae (Pgl), with a pair of rudimentary glossae (G/) between them. Each palpus is three-jointed. At the base of the anterior surface of the mentum, where the wall of the mentum is reflected into that of the hypopharynx (fig. 41), there is a small, median, oval, cup-shaped depression into which opens the duct from the salivary glands (S7D). A small prominence on the base of the hypopharynx fits into the salivary cup and apparently closes the latter when the labium is pressed against the hypopharynx. NO. 3 INSECT HEAD—SNODGRASS 107 The walls of the salivary cup are chitinous, and its posterior inner surface bears a strong chitinous keel (figs. 40D, 41 v) projecting into the interior of the labium in the base of the mentum. Two pairs of muscles (figs. 40 D, 26, 27) are attached upon the keel and the walls of the salivary cup. _ The musculature of the labium is in general similar to that of the maxillae. It includes the following muscles : 23.—Proximal retractors of the mentum (fig. 40D)—A pair of muscles arising on ventral surfaces of posterior tentorial arms; in- serted on lateral basal angles of mentum. 24.—Distal retractors of the mentum (fig. 40 D).—A pair of mus- cles arising on posterior surfaces of posterior tentorial arms ; extend- ing through submentum and mentum to be inserted on anterior wall of labium at inner basal angles of the glossae. The distal parts of these muscles are not seen in figure 40 D, being covered posteriorly by muscles 23 and 25. The labial muscles 23 and 24 evidently cor- respond with the tentorial adductors of the maxillae (E, 12, 13). 25.—Flexors of the paraglossae (fig. 40 D).—A pair of large mus- cles arising in lateral basal angles of mentum; inserted on bases of paraglossae, to posterior walls, near inner ends. Each of these muscles corresponds with the flexor of the galea in the maxilla (E, 16). The small labial glossae of Dissosteira have no muscles. 26, 27.—Muscles of the salivary cup (fig. 40 D).—Two pairs of muscles: one pair (26) arising on basal angles of mentum, converg- ing to insertions on keel of salivary cup; the other pair (27) arising on posterior wall of mentum near bases of palpi, converging proxi- mally to insertions on sides of salivary cup. These muscles apparently have no homologues in the maxillae; perhaps they are special labial muscles having something to do with the regulation of the flow of saliva from the salivary duct. 28.—Levator of the labial palpus (fig. 40 D).—Origin in lateral basal angle of mentum; insertion on dorsal rim of base of palpus. 20.—Depressor of the labial palpus (fig. 40 D).—Origin in distal median angle of mentum; insertion on ventral rim of base of palpus. 30, 31—Muscles in the labial palpus (fig. 40 D).—The first (30) a levator of second segment; second (37) a depressor (adductor) of third segment. THE PREORAL CAVITY AND THE HYPOPHARYNX The intergnathal space, or preoral cavity, of the grasshopper (fig. 41, PrC) is of large size, but it is mostly filled by the thick, tongue- like hypopharynx suspended from its roof (Hphy). Its anterior wall 108 SMITHSONIAN MISCELLANEOUS COLLECTIONS vou. SI is the posterior surface of the clypeus and labrum (Clp, Lin), which in the grasshopper is not produced into a specially developed lobe, or epipharynx. The lateral walls are the inner faces of the mandibles and maxillae ; the posterior wall is the anterior surface of the labium (Lb). The dorsal wall of the cavity represents the true sternal region of the head, sloping downward and posteriorly from the mouth open- ing to the base of the labium. It is mostly produced into the large, / litran Hphy Fic. 41—Stomodeum, and its dilator muscles in right half of head of Dissosteira carolina. Clp, clypeus; Cr, crop; cs, epistomal suture, Fr, frons; Hphy, hypopharynx, Lb, labium; Lm, labrum; Mth, mouth; Phy, pharynx; Prec, preoral cavity; SID, salivary duct; Tut, tentorium; v, salivary cup. median hypopharynx (Hphy), leaving otherwise only a narrow mem- branous area on each side between the base of the hypopharynx and the bases of the mandible and maxilla. The anterior or epipharyngeal wall of the preoral cavity presents a number of features of special interest (fig. 42 A). The lateral parts of the labral region of this wall are concave and fit closely over the smooth, rounded, anterior surfaces of the mandibles. The bands of hairs directed inward on the labral surface guard the exits from NO. 3 INSECT HEAD—SNODGRASS 10g between the mandibles, and the asymmetrical forms of the hair- covered areas here correspond with the different shapes of the two mandibles. Minute sense organs are scattered over this labral surface, especially on the bare lateral regions. A special group of similar but somewhat larger sense organs lies at each side of the notch in the ventral border of the labrum. The median area of the basal half of the labral surface forms a low elevation, the sides of which are thickly covered with long spine-like hairs curved inward and upward. This elevation projects between the inner edges of the closed mandibles, and its irregular contours fit with the lines of the opposing jaws. Its median surface is depressed and embraces the region of the internal Y-shaped ridge (7). The elevation is continued upward on the clyp- eal region, above the spreading arms of the Y-shaped ridge, and be- tween the inner recurved ends of the tormae (7Jor), and then into the mouth (Mth) and upon the anterior wall of the buccal cavity. A sinuous groove begins upon the elevation ventrally between the tormae, which extends dorsally and enlarges into a deep, median channel con- tinued into the anterior wall of the mouth and pharynx. At the sides of the lower end of the channel, between the slender arms of the tormae, are four asymmetrically placed, oval groups of small peg-like sense organs with large circular bases, partly covered from the sides by fringes of long recumbent hairs. The hypopharynx is a large median lobe suspended, as already noted, from the ventral wall of the head between the mouth and the base of the labium (fig. 41, Hphy). Its posterior end is closely covered by the paraglossal lobes, and its sides are concealed by the mandibles and maxillae. In form, as seen from below (fig. 42 C), the hypo- pharynx is somewhat ovate, with the smaller end anterior, but its pos- terior end is set off as a narrowed lobe by lateral constrictions. The lateral surfaces of the anterior division fit into the posterior concavi- ties of the mandibles, those of the posterior lobe are embraced by the concave inner faces of the laciniae. The posterior, basal extremity of the hypopharynx projects as a small median process into the salivary cup on the base of the labium (fig. 41). The lateral line of the hypo- pharyngeal base is marked by a slender, sinuous, chitinous bar on each side (w). The arrangement of the hairs clothing the hypopharynx is sufficiently shown in the figures (figs. 41, 42 C). On its sides and at the posterior end near the salivary cup are a few small sense organs similar to those of the labrum. Dorsal to the anterior end of the hypopharynx is an area that leads directly upward into the floor of the mouth. It possibly represents the sternal region of the protocephalic segments of the head. On its LO SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL. &t median surface (fig. 42 B,C) is a ridge, bordered by long hairs di- rected inward and upward, that continues dorsally from the narrowed end of the hypopharynx, and which is excavated by a median channel where it enters the mouth. At each side of this channel is an oval group of sense organs. Flanking the ridge are two chitinous bars (HS), the ventral ends of which articulate with the anterior extrem- ities of the lateral basal rods of the hypopharynx (w). Dorsally each bar forks into two arms, of which one (1) goes posteriorly to Fic. 42—The epipharyngeal surface, and the hypopharynx of Dissosteira carolina. A, epipharyngeal surface of clypeus and labrum, and ventral extremity of pharynx. B, lateral view of mouth opening, and of suspensorial apparatus of hypopharynx. C, antero-ventral view of hypopharynx and its suspensory rods. gAp, apodeme of adductor muscle of mandible; Clp, clypeus; Hphy, hypo- pharynx; HS, suspensorial bar of hypopharynx ; Lm, labrum; m, Y-shaped ridge in epipharyngeal wall of labrum; Mth, mouth; Phy, pharynx; Put, small lobe behind angle of mouth, possibly rudiment of tritocerebral appendage; Tor, torma; w, lateral basal bar of hypopharynx; +, mandibular branch of sus- pensorial bar (7.S) of hypopharynx; y, oral branch of same. the base of the adductor apodeme of the mandible (9APp), and the other (y) goes anteriorly, laterally, and dorsally into the angle of the mouth, where it forms a support for the insertion of the retractor muscle of the mouth angle (A, B, 38). The two lateral bars (fig. 42 B, HS) in the space between the hypo- pharynx and the mouth, with their posterior dorsal arms (1%) braced against the bases of the mandibular apodemes, and their ventral ends articulated with the basal rods (zw) of the hypopharynx, consti- tute a movable suspensorial apparatus of the hypopharynx. It is evident that a contraction of the mouth angle muscles (38) inserted NO. 3 INSECT HEAD—SNODGRASS RY on the anterior dorsal arms (y) of the bars must effect a movement of the hypopharynx, and that the latter would be lifted and swung forward beneath the mouth opening. The pull of the mouth muscles, however, also retracts the mouth angles, and there is probably thus accomplished a closing of the mouth upon the food mass accumulated in the preoral space above the anterior end of the hypopharynx. In the grasshopper, the mouth is closed also by the opening of the jaws, but, so far as can be observed in a dead specimen, the closing of the mouth in this case results mechanically from the transverse stretch- ing of the oral aperture between the separating bases of the adductor apodemes of the mandibles. Posteriorly the hypopharynx is fixed to the base of the labium, where its wall is reflected into that of the latter (fig. 41). The hypopharynx, therefore, can swing forward only in unison with the labium, but other- wise it is free to move to the extent permitted by the membranous areas laterad of its base. The only muscles properly belonging to the hypopharynx are the following: 32.—Retractors of the hypopharynx (figs. 40D, 41)—A pair of muscles arising posteriorly on extreme lateral ends of anterior arms of tentorium (fig. 40 D) ; inserted on posterior parts of basal rods of hypopharynx (fig. 41). The contraction of these muscles probably retracts the hypopharynx, and pulls the hypopharynx and labium posteriorly. The mouth aper- ture is opened by the contraction of the dilator muscles inserted on its anterior and posterior walls (figs. 41, 44, 33, 34, 41). The rods (HS) of the suspensory apparatus of the hypopharynx in the grasshopper are evidently remnants of the much larger sus- pensory plates of the hypopharynx in Apterygota and Myriapoda (fig. 21 A,B,C,E, HS). In Microcentrum, as already shown (fig. 20D), a small hypopharyngeal adductor muscle of the mandible is attached to the end of each rod. In the roach (Periplaneta) the chi- tinous parts of the hypopharyngeal suspensorium are more strongly developed than in the grasshopper, and their action can be more clearly demonstrated. In the bees, though the hypopharynx itself may be lacking, the oral arms of the suspensory bars are prolonged as slender rods into the lateral walls of the pharynx, and their basal ends are bridged by a wide plate on the pharyngeal floor. In Dissosteira there is at each side of the mouth, in the angle between the dorsal arms of the suspensorial bar of the hypopharynx, a very small but distinct membranous lobe of a definite form (fig. 42 B, Put), but having no apparent function, and bearing neither hairs nor sense 8 II2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOR. ROL # organs. The position of these lobes between the mouth and the ad- ductor apodemes of the mandibles strongly suggests that they are rudiments of the postantennal appendages of the tritocerebral seg- ment, which have otherwise not been observed in the adult of any pterygote insect. THE STOMODEUM At the upper end of the preoral cavity (fig. 41, PrC), anterior to the base of the hypopharynx, and immediately behind the base of the clypeus is the true mouth (Mth), or external opening of the stomo- deum. The mouth of Dissosteira is a transverse aperture having acute lateral angles, but without definite “ lips,” for the epipharyngeal \\ 2/ aC x ee Gz S = Pvent FrGny’ A ie —< SC MS J =r << Say PL iene Fic. 43.—Pharynx, crop, anterior gastric caeca, and associated organs of Dissosteira carolina. Ao, aorta; CA, corpus allatum (7); Cr, crop; FrGng, frontal ganglion; GC, gastric caecum; LNv, lateral stomodeal nerve; OeGng, posterior median or oesophageal ganglion; Phy, pharynx; PLGng, posterior lateral stomodeal gan- glion; Pvent, proventriculus. and the supra-hypopharyngeal walls are directly continued into the anterior and posterior walls of the buccal cavity and pharynx. The stomodeum of the grasshopper extends from the mouth upward in the anterior part of the head (fig. 41), then turns posteriorly above the tentorium, and continues rearward through the head and thorax into the base of the abdomen (fig. 43). By differences in its diameter and in the character of its walls, the stomodeum is differentiated into several parts, but only three parts are well defined in the grasshopper ; these are the pharynx, the crop, and the proventriculus. The pharynx, or first division of the stomodeum, is a narrow, mus- cular-walled tube bent downward to the mouth between the anterior arms of the tentorium (figs. 41, 43, 44, Phy). The region of the mouth, including the upper end of the preoral cavity (fig. 41, PrC) and the part of the stomodeum just within the oral aperture, may be distinguished as the buccal cavity because the muscles inserted on it NO. 3 INSECT HEAD—SNODGRASS IT3 (figs. 41, 44, 33, 34, 38, 41) function in connection with the mouth. The dorsal dilators (33, 34) arise upon the clypeus (fig. 41, Clp). The true pharyngeal region of the stomodeum of the grasshopper is differentiated into an anterior pharynx and a posterior pharynx, the two parts being thus named by Eidmann (1925) in the roach. The principal differences between the two parts of the pharynx, how- ever, are in the conformations of the cuticular lining, though the posterior end of the anterior pharynx is marked externally by a slight bulging of the lateral walls. The circumoesophageal connectives (fig. 44, CoeCon) lie approximately between the two pharyngeal sections. The crop (fig. 43, Cr) is a large, rather stiff-walled sack, represerit- ing probably both oesophagus and crop in insects with a long oesoph- ageal tube, though the posterior section of the pharynx in the grass- hopper appears to be the oesophageal region in the caterpillar (fig. 55). The anterior end of the crop in Dissosteira lies in the back of the head where it rests upon the bridge of the tentorium(fig. 41) ; the ventral surface of the thoracic part of the organ is supported by the spreading apophyses of the thoracic sterna. The anterior third of the crop (fig. 43) is somewhat set off from the rest by a slight nar- rowing of the walls; the posterior part tapers between the large anterior caecal pouches of the ventriculus, and ends in the proventric- ulus (Pvent). The proventriculus is a small, cup-shaped enlarge- ment of the posterior end of the stomodeum, mostly concealed between the bases of the ventricular pouches (GC). The frontal ganglion of the stomodeal (stomatogastric) nervous system (fig. 43, FrGng) rests against the anterior wall of the pharynx, and the posterior median oesophageal ganglion (OeGng) lies over the posterior end between the spreading bases of the last pair of dorsal dilator muscles of the pharynx (37). From this second median gan- glion a long lateral nerve (Nv) goes posteriorly on each side of the crop, ending on the rear part of the latter in a posterior lateral gan- glion (PLGng). A pair of short anterior lateral nerves from the oesophageal ganglion go laterally to a pair of globular bodies, possibly the corpora allata (CA), lying at the sides of the posterior pharynx. The anterior dilated end of the aorta (Ao) rests upon the oesophageal ganglion, and its open, trough-like lower lip is extended forward be- neath the brain. The Inner Wall of the Stomodeum—The surface of the intima, or cuticular lining, of the pharynx, crop, and proventriculus is diversified by various folds and ridges, most of which are clothed with hairs or are armed with small chitinous teeth. IIl4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL. 81 The channels on the walls of the preoral cavity that lead into the mouth are continued upon the inner walls of the anterior pharynx. The median epipharyngeal groove proceeds upward on the anterior pharyngeal wall between two converging ridges, but it soon ends in a thick median fold which follows the midline of the roof of the posterior pharynx to the end of the latter. Likewise, the median channel leading upward from the base of the hypopharynx is con- tinued on the rear wall of the anterior pharynx, between two con- verging ridges, and ends in a median ventral fold on the floor of the posterior pharynx. From the lateral angles of the mouth, wide chan- nels go dorsally in the side walls of the anterior pharynx, but these again end each in a lateral fold of the posterior pharynx. Thus the relative positions of the principal ridges and grooves in the walls of the two parts of the pharynx are reversed. In the posterior pharynx there is a slenderer intermediate fold between each two of the major dorsal, lateral, and ventral folds. These eight folds of the posterior pharynx end at the entrance of the crop, giving the aperture a stellate appearance when seen from the lumen of the crop. All the pharyngeal folds, except the midventral fold of the posterior pharynx, are clothed with hairs directed backward. In the crop, a wide dorsal channel proceeds from the pharyngeal opening posteriorly on the anterior third of the upper wall between converging folds of the intima. A narrower ventral channel follows the midline of the floor between a pair of folds that diverge posteriorly and are lost beyond the middle of the organ. The lateral walls of the anterior half of the crop are closely corrugated by obliquely trans- verse ridges, which bear rows of small, slightly curved, sharp-pointed, chitinous teeth projecting backward. The anterior three or four transverse ridges on each side are particularly conspicuous by reason of their greater width, and because they are thickly beset with similar but slightly larger teeth than those of the other ridges. In the posterior, narrowed part of the crop the transverse ridges are replaced by fine, parallel, lengthwise folds, following the lines of the longitudinal muscle fibers. Numerous teeth are present here also, but they are smaller and blunter than those of the anterior region, and are mostly arranged in small groups, usually two or three together, on elevations of the intima along the folds. The interior characters of the crop are better developed and the teeth are more numerous in the larger organ of the female grasshopper than in that of the male. They can be studied best on pieces of the intima stripped from the tough muscular sheath of the crop. NO. 3 INSECT HEAD—-SNODGRASS 15 The walls of the short proventriculus are produced into six flat, triangular elevations having their bases contingent anteriorly, and their apices directed backward, where they all end on the rim of the wide, round orifice into the ventriculus. The proventricular ridges are not mere folds of the intima, for each is formed by a thick mass of the underlying epithelial cells. The surface of the intima in the pro- ventriculus is smooth, except for a few very small teeth on the edges of the triangular ridges, and areas of minute granulations on the distal halves of the latter. The posterior margin of the proventricular wall is reflected outward upon itself to form a short circular fold project- ing into the anterior end of the ventriculus, reaching just past the openings of gastric caeca. The intima covers the outer surface of the fold, but terminates at the base of this surface. The line of the latter, therefore, marks the end of the stomodeal or anterior ectodermal section of the alimentary canal. The Muscular Sheath of the Stomodeum.—The stomodeal walls are everywhere covered with flat bands of muscles, which in general take a transverse and a longitudinal direction, the transverse bands being external and the longitudinal internal; but the distribution of the two sets is not such as to form a regular net-pattern on all parts of the stomodeum. On the posterior two-thirds of the crop, the ex- ternal transverse fibers have the form of continuous rings encircling the organ, and the longitudinals run with its length. On the anterior third, however, the ring muscles are interrupted laterally and dorsally, and their layer is continued only on the ventral surface as a series of ventral arcs; but the fibers of a Jatero-ventral tract of the posterior longitudinal muscles on each side curve upward on the lateral wall of the crop where the circular bands are interrupted, and are continuous with those from the opposite side over the dorsal surface as an external layer of obliquely transverse fibers reaching to the base of the pharynx. On the pharyngeal tube the muscles again take the pattern of regularly arranged external circular and internal longitudinal fibers. The cir- cular fibers of the pharynx may belong to the interrupted set of circular fibers of the crop, but the longitudinal fibers are continued irregularly into the walls of the crop on the inner surface of the anterior circular fibers of the latter, and they do not, therefore, belong to the same layer as the posterior longitudinal crop muscles. A close study of the stomodeal musculature of the grasshopper would show some com- plexity of detail in the arrangement and relationship of the muscle fibers, but nothing approaching the intricacy of the fiber connections in the muscular layers on the pharynx and crop of the caterpillar, to be described later. 116 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 81 The Dilator Muscles of the Stomodeum.—Thirteen paired sets of muscle fibers and one median unpaired muscle arising on the skeletal parts of the head or thorax are inserted on the stomodeal walls in Dissosteira (figs. 41, 43, 44). These muscles may be classed as dor- sal, lateral, and ventral according to their insertions, though because of the downward flexure of the pharynx, the first “ dorsal” and “ ventral’? muscles are anterior and posterior. The dilator muscles of the stomodeum, sometimes called also suspensory muscles, enumer- ated from 33 to 46 inclusive, are as follows: Fic. 44.—Dilator muscles of the buccal region, pharynx, and crop of Dissosteira carolina. CoeCon, circumoesophageal connective ; Cr, crop ; Mth, mouth; Tnt, tentorium. 33.—First anterior dilators of the buccal cavity (figs. 41, 44).— A pair of fan-shaped muscles arising on inner wall of clypeus (fig. 37 B), the fibers spreading to their insertion on anterior wall of buccal cavity (fig. 44) mostly distal to oral aperture. 34.—Second anterior dilators of the buccal cavity (figs. 41, 44).— A pair of fan-shaped muscles similar to 33, arising on clypeus near epistomal ridge (fig. 37 B) ; inserted laterad of 33 and mostly proxi- mal to oral aperture. 35.—First dorsal dilators of the pharynx (figs. 41, 44).—A pair of slender muscles arising on frontal area of head wall, each attached between labral retractors of same side (fig. 38 D) ; inserted on anterior wall of pharynx. NO. 3 INSECT HEAD—SNODGRASS ey 36.—Second dorsal dilators of the pharynx (figs. 41, 44) —Each arises by a slender stalk on subantennal ridge of frons (fig. 41) ; in- serted by spreading base on upper end of anterior pharynx. 37-—Third dorsal dilators of the pharynx (figs. 41, 43, 44). —Each arises by slender stalk on vertex near inner rim of compound eye just anterior to first dorsal fibers of mandibular adductor ; inserted by widely spreading base on dorsal wall of posterior pharynx. 38.—Retractors of the mouth angles (figs. 41, 44)—These, the largest muscles of the stomodeum, and the first of the lateral series, arise on the subantennal ridges of the frons (fig. 38 D), and extend downward and posteriorly to their insertions on the oral arms of the hypopharyngeal suspensorial rods (figs. 42 A, B, 44) in the lateral angles of the mouth. 39.—F*irst lateral dilators of the pharynx (fig. 44) —A pair of slender muscles arising laterally on frontal region ; inserted on sides of anterior pharynx. 40.—Second lateral dilators of the pharynx (figs. 41, 44).—A slen- der muscle on each side, arising on posterior face of distal end of dor- sal arm of tentorium (fig. 41) ; inserted by spreading base on upper end of anterior pharynx (fig. 44). 41.—V entral dilator of the buccal cavity (figs. 41, 44) —A median, unpaired, strap-like muscle arising on ventral face of body of tentor- ium ; inserted on median groove of posterior wall of mouth. 42.—First ventral dilators of the pharynx (figs. 41, 44).—A pair of fibers arising on ventral surface of tentorium ; going anteriorly to insertions medially on lower end of posterior wall of anterior pharynx. 43.—Second ventral dilators of the pharynx (figs. 41, 44)—A group of diverging fibers on each side, arising on anterior edge of tentorium ; inserted latero-ventrally on anterior pharynx. 44.—Third ventral dilators of the pharynx (figs. 41, 43, 44) —A large fan of fibers on each side, arising on dorsal edge of posterior arm of tentorium; the spreading fibers inserted ventro-laterally along entire length of posterior pharynx. 45.—Anterior dilators or protractors of the crop (figs. 41, 43, 44). —A large group of fibers arising on each posterior tentorial arm. behind origin of 44; spreading posteriorly to insertions ventro-later- ally along anterior third of crop. These are the last of the stomodeal muscles that have their origin in the head. 46.—Posterior protractors of the crop and gastric caeca (fig. 43).— A pair of long, branched muscles, each arising by a slender stalk on inner surface of prothoracic tergum, just anterior to base of troch- antinal muscle ; branching downward and posteriorly, one branch in- 118 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 81 serted on lateral wall of crop just above posterior lateral stomodeal ganglion (PLGngq), the others on tips of the gastric caeca (GC) of same side. THE MECHANISM FOR MOVING THE HEAD The head of the grasshopper is freely attached to the prothorax by a membranous neck, but its movements are somewhat limited by the overlapping anterior edgés of the protergum, and by the pair of cervical sclerites on each side (fig. 45 B, rev, 2cv) which link the head with the concealed episternal plate of the prothorax (Eps;). The cervical sclerites, however, constitute an important part of the mechanism for moving the head The two plates of each pair are articulated end to end, and ordinarily they are bent downward at an angle to each other (fig. 36 A, cv). The first is articulated anteriorly to the posterior margin of the postoccipital rim of the head (fig. 45, g), the second posteriorly to the anterior edge of the prothoracic episternum (Eps,). The neck plates thus constitute a fulcrum on each side between the head and the thorax, giving a leverage to the dorsal and ventral muscles extending from the postoccipital ridge and ten- torium to the prothorax and the first thoracic phragma. Moreover, upon each plate are inserted strong levator muscles (fig. 45 B) arising on the back of the head and on the prothoracic tergum, and the con- traction of these muscles, with the consequent straightening of the angle between the two plates of each pair, must cause the protraction of the head. From each anterior plate a horizontal muscle extends to the prosternal apophysis of the opposite side (fig. 45 A, B, 54). Be- sides the muscles that connect the skeletal parts of the head, neck, and prothorax, there are two muscles on each side inserted directly upon the neck membrane (A, 56, 57). It is difficult to give names signifying function to the neck muscles, for it is evident that the function will depend on whether the two muscles of any pair act in unison, or as antagonists. The neck muscles of Dissosteira are as follows, on each side: 47.—First protergal muscle of the head (fig. 45 A).—A slender muscle arising dorsally on prothoracic tergum ; inserted dorso-later- ally on postoccipital ridge of head (Po). 48.—Second protergal muscle of the head (fig. 45 A).—A larger muscle arising on principal ridge of protergum (e) ; inserted with 47 on postoccipital ridge of head. 49.—Longitudinal dorsal muscle of the prothorax. (fig. 45 A).— Extends from first thoracic phragma (7Ph) to postoccipital ridge (PoR) just below 48. NOR 1S INSECT HEAD—SNODGRASS 11g 50, 51.—Cephalic muscles of the cervical plates (fig. 45 A, B).— Origin on postoccipital ridge below 49; both extend ventrally and posteriorly, the first (50) inserted on first cervical plate, the second (52) on second plate. 52, 53:-—Protergal muscles of the cervical plates (fig. 45 B).— Origin dorso-laterally on prothoracic tergum; both extend ventrally and anteriorly, crossing internal to 50 and 57, to be inserted on first PoR A Fic. 45.—Muscles of the neck of Dissosteira carolina, right side, internal view. A, muscles extending between head and prothorax, omitting 52, 53 and 54, inserted on cervical sclerites (B). B, head and prothoracic muscles of cervical sclerites. Bs,, basisternum of prothorax; c, first ridge of protergum; cv, first cervical plate; 2cv, second cervical plate; d, second ridge of protergum; e, third ridge of protergum; Eps, episternum of prothorax; Eps, episternum of mesothorax ; g, process of head articulating with first cervical sclerite; H, head; 1Ph, first thoracic phragma; PoR, postoccipital ridge; PT, base of posterior arm of tentorium; Rd, posterior fold of protergum; SA, apophysis of prothoracic sternum; Spn, spina; 71, tergum of prothorax. cervical plate, the first (52b) with a branch (52a) to articular process (g) of postoccipital ridge. 54.—Prosternal muscle of the first cervical plate (fig. 45 A, B).—A diagonal, horizontal muscle arising on apophysis of prothoracic ster- num (A, SA), crossing its fellow to insertion on inner edge of first cervical plate of opposite side (B). 55.—Longitudinal ventral muscle of the prothorax (fig. 45 A).—A broad, flat muscle from prosternal apophysis (54) to base of poste- rior arm of tentorium (PT). I20 SMITHSONIAN MISCELLANEOUS COLLECTIONS voL. 81 56.—Dorsal lateral neck muscle (fig. 45 A).—A band of slender fibers from first phragma (7P/), inserted on base of neck membrane. 57—Ventral lateral neck muscle (fig. 45 A, B).—A short, flat muscle from anterior edge of prothoracic episternum (/ps,), inserted on base of neck membrane. VI. SPECIAL MODIFICATIONS IN THE STRUCTURE OF THE, HEAD. The important structural variations in the head of biting insects affect principally the fronto-clypeal area, and the posterior lateral and ventral regions. Modifications of the facial plates are often to be correlated with variations in the relative size of the buccal and pharyn- geal parts of the stomodeum, or with a special development of the mouth cavity. Modifications in the posterior ventral parts of the head are correlated with a flattening and elongation of the cranial capsule, usually resulting from an upward tilting of the head on the neck by which the mouth parts become directed forward, and, in certain orders, are accompanied by an elongation of the submentum an- teriorly, with a differentiation of this plate into a posterior gular sclerite and a secondary anterior submental sclerite. MODIFICATIONS IN THE FRONTO-CLY PEAL REGION The prostomial part of the insect head includes the frons, the clyp- eus, and the labrum. Whether or not it comprises also the region of the compound eyes may be regarded as an open question, and one for the embryologists to settle. If the compound eyes belong to the first true segment of the head, it is probable that the frontal sutures define the posterior limit of the prostomium; otherwise the sutures must be secondary formations within the area of the prostomium. The frontal sutures do not always mark the lines of cleavage in the head cuticula at the time of a molt. In an odonate nymph, for example (fig. 46 1), the facial clefts (t) of the molting cuticula extend from the coronal suture outward and downward on each side between the eyes and the bases of the antennae, far outside the possible limits of the frons (Fr). , The part of the postembryonic head that may be defined as the frons is the area included between the frontal sutures, where these sutures are fully developed (fig. 46 B, Fr). The frontal sutures (fs) extend typically from the coronal suture (cs) to the neighborhood of the anterior articulations of the mandibles (c, c). The true frontal region, therefore, can not include the bases of the antennae, which NO. 3 INSECT ITEAD—-SNODGRASS 121 organs belong to the second head segment behind the prostomium, and acquire their facial positions secondarily by a forward and upward migration. Ventrally the frons is limited, and separated from the Fic. 46—Modifications in the facial structure of the insect head. A, Forficula auricularia. B, Popillia japonica, larva. C, Pteronidea ribesi, larva, inner surface of front of head. D, espa maculata, well-chitinized larva. E; Pteronidea ribesi, adult. F, Apis mellifica. G, Psocus venosus. H, Magicicada Eepiendecim. I, molted skin of an Aeschna larva. Aclp, anteclypeus ; Ant, antenna; AT, anterior arm of tentorium; at, anterior tentorial pit; c, anterior ‘articulation of mandible; Clp, clypeus ; dt, attachment of dorsal tentorial arm to head wall; es, epistomal suture; Fr, frons; fr, “adfrontal”; Lin, labrum; O, ocellus; s, suture of Forficula diverging from end of coronal suture; SgR, subgenal ridge; t, molting split in Aeschna larva diverging from end of coronal suture, but is not frontal suture. clypeus, by the epistomal suture (fig. 46 B, es), except when this suture is lacking. If a median ocellus is present, it is situated in the upper angle of the frons (figs. 46 E, 47B). The muscles of the labrum, some of the dilator muscles of the pharynx, and the retractors of the mouth angles, when present, have their origins on the frons. By I22 SMITHSONIAN MISCELLANEOUS COLLECTIONS- voL. 81 these characters, especially the position of the median ocellus and the origin of the labral muscles, the true frontal region is to be identified when the frontal sutures are imperfect or obsolete (fig. 46 FE, F, Fr). As was shown in the study of the grasshopper (fig. 36 B), the frontal region of the face may present a number of secondary lines formed by ridges of the inner surface. In the Dermaptera two sutures (fig. 46 A, s) diverge widely from the end of the coronal suture (cs) and extend outward to the compound eyes. It appears doubtful that these are the frontal sutures, for the true frontal region should be the smaller triangular area indistinctly defined on the median part of the face. The clypeus (fig. 46B, Clip) is a distinct area of the prostomial region, and is to be identified by the origin of the dilator muscles of the mouth and buccal cavity on its inner wall. It is almost always in biting insects separated from the labrum by a flexible suture, and it is demarked from the frons whenever the epistomal suture is present. The clypeus is sometimes divided into an anteclypeus and a postclyp- eus by a partial or complete transverse suture; but often the term “anteclypeus’”’ is given to a more or less membranous area between the clypeus and the labrum (fig. 46G, Aclp), and it is likely that regions named “ anteclypeus ” are not equivalent in all cases. The labrum (fig. 46 B, Lim) hangs as a free flap before the mouth. It is a preoral lobe of the prostomium characteristic of insects, myria- pods, and crustaceans. The insect labrum is usually movable, and is provided with one or two pairs of muscles (though both may be ab- sent), which, as above noted, have their origin on the frons. The labral muscles, therefore, are strictly muscles of the prostomium. The principal departure from the typical structure in the pro- stomial sclerites arises from variations in the development or in the position of the epistomal suture, and from a partial or complete sup- pression of the frontal sutures. The epistomal suture is the external groove formed incidentally to the development of an internal transverse ridge across the prostomial area. Since this ridge in generalized insects lies approximately be- tween the anterior articulations of the mandibles, its primitive position suggests that it was developed to strengthen the lower edge of the face between the mandibular bases. The epistomal ridge itself is a con- tinuation of the subgenal ridges, and the epistomal suture is, there- fore, continuous with the subgenal sutures. In the Ephemerida and Odonata, as we have seen, the anterior arms of the tentorium arise in the subgenal sutures laterad of the bases of the mandibles. In some of the Orthoptera, as in the roach, and in larvae of Coleoptera, NOH S INSECT HEAD—SNODGRASS 123 the tentorial arms have moved forward to a position above the mandib- ular articulations, and their external openings, the anterior ten- torial pits, appear in these positions (fig. 46 B, at, at). In some of the more generalized insects, the epistomal ridge and its suture are lacking, as in the roach, and there is then present only a single fronto-clypeal sclerite (fig. 47 A, Fr-Clp). In such cases, the tentorial pits (at) lie in the anterior extremities of the subgenal sutures (sgs), above the anterior articulations (c) of the mandibles. Where an epistomal ridge unites the subgenal ridges across the face, separating the clypeus from the frons, the tentorial pits may retain Fic. 47.—Diagrams showing variations in the position of the epistomal suture (es), and the relations of the frons and the clypeus. Aclp, anteclypeus; at, anterior tentorial pit; c, anterior articulation of mandible ; Clp, clypeus; es, epistomal suture; Fr, frons; fr, “adfrontal” ; Fr-Clp, fronto-clypeus; fs, frontal suture; /, line of secondary ridge across lower part of clypeus; Lm, labrum; LmMcls, labral muscles, with origin always on frons; O, median ocellus. their positions above the mandibular articulations (fig. 46 A, B, at, at); but more commonly they move into the epistomal suture (fig. 47 B). In any case, the tentorial pits identify the epistomal suture, when this suture is present. The mandibular articulations (c, ¢) are carried by the ventral margin of the epicranium and are not true land- marks of the epistomal suture, as has been pointed out by Yuasa (1920), and by Crampton (1925). As long as the epistomal suture maintains its direct course across the face, no complications arise; but the suture is frequently arched upward, and this shift in the position of the suture extends the clyp- eus into the facial region above the bases of the mandibles, and re- duces the area of the frons (fig. 47C). A modification of this kind I24 SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL. 81 has taken place in the Hymenoptera. In the larval head of Vespa (fig. 46 D) the clypeus has clearly encroached upon the area of the frons by a dorsal arching of the epistomal suture (cs). In an adult tenthredinid (FE), the same condition is observed, but the lower parts of the frontal sutures (fs) are lost, and the bases of the antennae have approached each other mesally, and have constricted the frontal area between them. In the adult of Apis (F) the condition is more exaggerated—the epistomal suture (es), identified by the tentorial pits (at, at), is arched upward almost to the bases of the antennae, and the frontal sutures are obsolete. The frontal area (Fr), however, is to be identified by the position of the median ocellus, and the points of origin of the labral muscles between and just above the antennal bases. The head of a larval tenthredinid (fig. 46C) presents a specialized condition, for the single large facial plate is here clearly a fronto-clypeus, as shown by the origin of the labral muscles on its upper parts, and by the origin of the tentorial arms (AT) from the ridges at its sides. Evidently, the median part of the epistomal ridge and its suture has been suppressed. A similar condition is to be observed in some trichopteran larvae. A still greater degree in the upward extension of the clypeus is shown on the face of a psocid (fig. 46 G). Here the epistomal suture (es) 1s arched high above the tentorial pits (at, at), and the clypeus (Clp) becomes the large, prominent, shield-shaped plate of the face between the bases of the antennae. The frontal sutures are lacking, but the frontal area (Fr) is that between the bifid end of the coronal suture and the clypeus, on which is located the median ocellus. A weakly chitinized area below the clypeus is sometimes called the anteclypeus (Aclp), but it appears to be only a chitinization of the connecting membrane between the clypeus and the labrum. The clypeus, finally, attains its greatest development at the expense of the frons in the Homoptera (fig. 47 D). In the cicada (fig. 46 H), the clypeus is the great bulging, striated plate of the face upon which arise the dilator muscles of the mouth pump. The dorsal arch of the epistomal suture (es) lies on a level with the antennal bases, and the anterior tentorial pits (at, at) are in its upper lateral parts, just above the dorsal extremities (c, c) of the mandibular plates (Md). The frons is a small, indistinctly defined triangular area (Fr) bearing the median ocellus in the adult. It is more strongly marked in the nymph, and is cut out by the opening of the frontal sutures at the time of the molt. The plate below the principal clypeal sclerite is probably an anteclypeus (Aclp), because in some Hemiptera it is not distinctly separated from the area above it, but it is questionable if NO. 3 INSECT HEAD—-SNODGRASS 1 on it is homologous with the preclypeal area of the psocid (fig. 46 G, Aclp). The terminal piece in the cicada (H, Lm) that closes the groove in the upper part of the labium would appear to be the labrum by comparison with Heteroptera. The “ mandibular plates” (Md) on the sides of the head must be the true bases of the mandibles. Their upper ends (c, c) have the same relations to the surrounding parts that the anterior mandibular articulations have in biting insects. The mandibular bristles are chitinous outgrowths from the ventral pos- terior angles of the plates, and the protractor apparatus of each bristle in the adult is differentiated from the posterior margin of the mandib- ular plate, as the writer has elsewhere shown (1927). In the larvae of Lepidoptera, a somewhat different type of modi- fication has produced an unusual distortion in the relation between the frons and the clypeus. The caterpillar head shows no essential varia- tion within the order, but the homologies of the facial structures are clear if interpreted by the characters which serve as identification marks in the other orders. The triangular facial plate (fig. 50 A) thus becomes the clypeus, because the suture (es) bounding it is identi- fied as the epistomal suture by the origin of the anterior tentorial arms from its lateral parts (fig. 50 I, AT). Upon this plate arise the muscles of the buccal region of the stomodeum. The median part of the frons is invaginated and forms the thick internal ridge (Fr) dorsal to the apex of the clypeus, which is to be identified as the frons by the origin of the labral muscles upon it. The so-called “ adfrontals ” (A, fr) are probably lateral remnants of the frons at the sides of the clypeus, and the ‘“‘adfrontal” sutures are the true frontal sutures (fs). That the relations of the plates of the caterpillar’s head, as thus established, are identical with those in other insects is made clear in the diagram given at E of figure 47. The clypeus (C/p) has simply extended into the area of the frons, and the median part of the latter plate (Fr), bearing the origins of the labral muscles, has been in- flected, while its distal parts, the so-called “ adfrontals ” (fr), maintain the original lateral ground of the primitive frontal area. The lower part of the clypeus is sometimes strengthened between the bases of the jaws by a secondary thickening forming a submarginal ridge (/1) on its inner surface. MODIFICATIONS IN THE POSTERIOR VENTRAL REGION OF THE HEAD The structural changes in the posterior parts of the head described here are associated with an elongation of the postgenal regions, re- sulting in the production of a long interval between the foramen 126 SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL. &I magnum and the posterior articulations of the mandibles. Two dif- ferent types of structure follow from this style of modification, one shown in adult Hymenoptera and in the larvae of Lepidoptera, the other in those orders in which a gular plate is developed. The morphology of the posterior surface of the hymenopteran head is comparatively easy to understand, for, in the larval stages, the rear aspect of the head presents the same structure as does that of an adult orthopteron (fig. 36C). In the head of the larva of Vespa, for example (fig. 48 A), the details of the structure are exactly as in the grasshopper. There is a distinct postoccipital suture (pos) ending below in the invaginations of the posterior arms of the ten- torium (pt, pt). The postocciput (Poc) is very narrow, but it forms the marginal lip of the head capsule behind the postoccipital suture. Fic. 48.—Development of the posterior head region in Hymenoptera. A, posterior surface of head of larva of Vespa maculata. C, same of the adult. D, corresponding view of head of adult Apis mellifica. Cd, cardo; Lh, labium; Oc, occiput; Pge, postgena; Poc, postocciput; pos, postoccipital suture; pt, posterior tentorial pit; St, stipes. The labium (Lb) is suspended from the ventral neck membrane, and the cardines of the maxillae (Cd) are articulated to the ventral cranial margins just anterior to the tentorial pits. In the adult wasp (fig. 48 B) the back of the head presents a quite different appearance from that of the larva. The foramen magnum is greatly contracted and is reduced to a small aperture in the center of a broad occipito-postgenal field. It is surrounded by a wide post- occipital collar (Poc) set off by the postoccipital suture (pos), in which suture are located the posterior tentorial pits (pt, pt). The labium (Lb) is detached from the neck and displaced anteriorly (ventrally), and the space between its base and the neck is closed by mesal extensions of the inner angles of the postgenae (Pge, Pge). » The articulations of the cardines (Cd) are also far removed from the tentorial pits (pt, pt), and are separated from them by the interven- ing bridge of the postgenae. In the wasp the postgenal bridge pre- 3 INSECT HEAD—-SNODGRASS 127 serves a median suture, but in the honeybee (C) the line of union between the postgenal lobes is obliterated, and the bridge presents a continuous surface in the space between the foramen magnum and the fossa containing the bases of the labium and maxillae. In an adult tenthredinid (Pteronidea), on the other hand, the foramen magnum, though greatly reduced in size by the development of a wide occipito-postgenal area, is still ““ open ’’ below, that is, it is closed by a narrow remnant of the neck membrane between the approximated angles of the postgenae. The labium, however, is displaced ventrally and united with the bases of the maxillae. In the Hymenoptera, then, there can be little question as to the line of evolution that has produced the structure of the back of the head in the higher forms. The resulting condition has been correctly observed by Stickney (1923), who says: “In many Hymenoptera the mesal margins of the postgenae are fused between the occipital foramen and the articulation of the labium.’’ A very similar modi- fication of the head has taken place in the caterpillars, as will be shown later, in which the parts constituting the “hypostoma”’ (fig. 51 A, Hst) correspond with the postgenal bridge of adult Hymenoptera. In either case, an unusual thing has happened in that the labium, after being moved forward to unite with the maxillae, has been separated from its own segment by the intervention of parts of the first maxil- lary segment. If the postgenae are lateral tergal elements of the head wall, their ventral union finds a parallel in the prothorax of the honeybee, which is completely encircled behind the bases of the legs by the prothoracic tergum. The modifications in the posterior ventral parts of the head in those orders in which a “ gula ” is developed are difficult to explain if studied only in the higher phases of their evolution, but they can be understood if traced from forms that show the simpler earlier stages of departure from the normal. In the Blattidae, the cranium is much flattened, but the essential head structure has not been altered, its posterior parts retaining the same form as in the less movable head of the grasshoppers. In many insects, especially in the Neuroptera and Coleoptera, however, the flattened head is not only turned upward on the neck, causing the true anterior surface to become dorsal and the mouth parts to be directed forward, but the ventral surface of the head has been elongated to preserve the vertical plane of the foramen magnum. In such insects the bases of the mouth parts become separated from the foramen magnum by a wide space, and in this space there appears a median 128 SMITHSONIAN MISCELLANEOUS COLLECTIONS voL. 81 plate called the “gula.” The nature of the gula has long been a puzzle to entomologists, but Crampton (1921, 1928) has given reasons for believing that it is a differentiation of the base of the labium, and a few examples taken from the Coleoptera will amply substanti- ate this view. In a scolytid or scarabaeid beetle larva the structure of the head does not differ essentially from that of the grasshopper. The face is directed forward, the mouth parts hang downward, and the under surface of the head is short. In the scarabaeid larva (fig. 49 A) the occipital and postgenal regions terminate in a postoccipital suture (pos), in the ventral ends of which are situated the large invagina- tions (pt, pt) of the posterior arms of the tentorium. Beyond the suture is a narrow postoccipital rim of the cranium (Poc), best de- veloped ventrally, where the lateral cervical sclerites (cz) are articu- lated to it. The postoccipital ridge is developed on each side of the foramen magnum into a broad apodemal plate (Pok), the two plates constricting the foramen laterally, and uniting ventrally in the broad tentorial bridge, which is concealed in the figure by the ventral part of the neck membrane (NV/b). The labium, the maxillae, and the mandibles of the scarabaeid larva are suspended from the ventral edges of the cranium exactly as in the grasshopper (fig. 36 C), but the two forms differ by the elongation in the beetle (fig. 49 A) of the postgenal margins of the head between the articulations of the car- dines (e) and the posterior articulations of the mandibles (a). The basal part of the submental region of the labium in the scara- baeid larva, Popillia japonica (fig. 49 A), is chitinized to form a triangular plate (Smt). This plate is attached to the mesal points of the postgenae (Pge), and has its extreme basal angles prolonged be- hind the tentorial pits to points (f, f) corresponding with the basal articulations of the submentum with the postocciput in an orthopteron (fig. 36 C, f). There can be no doubt that this) plate in the beetle head is the submentum, or a chitinized basal part of the submentum. It is marked by a transverse groove between the tentorial pits (pt, pt). In a silphid larva (fig. 49 B) the general structure of the head is similar to that in the scarabaeid larva, but the ventral postgenal mar- gins between the articulations of the cardines (e, ¢) and the mandibles (a) are much longer, and the posterior tentorial pits (pt, pt) are approximated in the mesally prolonged basal angles of the postgenae. The submentum (Smt) is large; its base is narrowly constricted be- tween the tentorial pits, which here almost cut off a small but distinct proximal area (Gu). The lateral angles of this extreme basal area of the submentum are prolonged behind the tentorial pits and become con- NO. INSECT HEAD—SNODGRASS 12 9 tinuous with the postocciptal rim of the cranium (Poc), which is set off by the postoccipital suture (pos) ending ventrally in the tentorial pits (pt, pt). “ce Fic. 49.—Evolution of the gula” in Coleoptera. A, Postero-ventral view of the head of a scarabaeid larva, Popillia japonica. B, same of a silphid larva, Silpha obscura. C, ventral surface of an adult meloid, Epicauta pennsylvanica. D, same of a carabid larva, Scarites. a, posterior articulation of mandible; Cd, cardo; cv, cervical sclerite; e, an- terior articulation of mandible; Gu, gula; Page, postgena; Poc, postocciput ; PoR, postoccipital ridge; pos, postoccipital suture; pt, posterior tentorial pit; I Smt, submentum. The characteristic structure of an adult coleopteran head is well illustrated in the head of a meloid beetle (fig. 49 C). The form of the . cranial capsule here differs principally from that of the scarabaeid or silphid larva in the lengthening of the postgenal regions between the foramen magnum and the articulations of the cardines (¢, e). | & T30 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 8I The extension of the ventral surface of the cranial wall accommodates the head to its horizontal position, and has involved a great elongation in that part of the submentum which lies between the posterior ten- torial pits (pt, pt) and extends forward to the articulations of the cardines (e, e). This region of the submentum is known as the gula. In Epicauta (fig. 49 C) the tentorial pits lie at about the middle of the lateral margins of the gula, and the ventral ends of the postoccipital suture (pos) are, consequently, turned anteriorly and lengthened in the same direction behind the pits. The ventral parts of the post- occipital suture, terminating in the tentorial pits, now become the so- called “ gular sutures.” It is evident that the large gular region in the adult meloid head (fig. 49 C) lying posterior to the tentorial pits and continuous basally with the postoccipital rim of the cranium (Poc) is produced from the small but corresponding area in the larval silphid head (B, Gu), and that this area, in turn, is merely the basal strip of the submentum in the scarabaeid larva (A, Smt), attached to the postocciput by its lateral extremities (f, f). In adult Coleoptera the distal end of the gula may be differentiated as a “ pregula” or “ gular bar” (C, Pgu). It supports the terminal part of the original submental plate (Smt), which lies between the bases of the maxillae, and which, in a restricted sense, is usually called “the submentum” by coleopterists. The pregular region may fuse laterally with the ‘“‘ hypostomal” regions of the postgenae, and in other ways the more primitive structure may become so obscured that the relations of the parts are difficult to determine except by studying them in a gradient series of simpler forms. The comparative studies made by Crampton (1921, 1928) on the gula in various orders show fully its numerous variations, and demonstrate its origin from the proximal part of the primitive submental plate. Stickney (1923) also has well illustrated the structure of the gula and associated parts in a large number of coleopteran forms. Stickney fails to recognize, however, that the “ gular sutures” are direct continuations of the ventral ends of the postoccipital suture, and that, therefore, the gular plate between them must be the basal part of the submentum. He would explain the gular bridge in the Coleoptera as a product of the ventral fusion of the edges of the postgenae, and the gular sclerite as a plate cut out of this newly-formed region by the anterior exten- sion of the “ gular sutures.” As we have seen, the ventral bridge of the cranial walls is formed in this manner in the Hymenoptera (fig. 48), as Stickney has pointed out, but in the Hymenoptera the ten- torial pits have remained at the sides of the foramen magnum, and the labium has lost its original connection with the postoccipital region. ce a coe INSECT HEAD—SNODGRASS 131 The facts are quite otherwise in the Coleoptera, for here the labium retains its postoccipital connections, and its base has been drawn out between the lengthened postgenal margins to form the gula. In certain Coleoptera the postgenal margins do become closely ap- proximated (fig. 49 D), but, in such cases, the gula is compressed be- tween the postgenae, and sometimes almost obliterated. The gular sutures may then be partially or wholly united into a median gular suture, with which are closely associated the two tentorial pits (pt, pt). Intermediate stages of this condition are well shown in some of the Rhyncophora, in which the head is drawn out into a “ snout.” In the Neuroptera, both larvae and adults, and in larval Trichoptera, a gular plate is developed showing essentially the same structure and variations. of form as in the Coleoptera. The gular structure has been described in various members of these orders and others in addition to the Coleoptera by Crampton (1921, 1928). In the Termitidae, Crampton shows, the gular region of the submentum may be very much elongated, and in the soldier of Termopsis its margins become united with the lengthened edges of the postgenae to form a typical gular plate. The question of the derivation of the gula, the answer to which is, that the gula is a part of the submental region of the labium, is not to be confused with the question as to the origin of the submentum itself. The various views concerning the nature of the submentum have been already discussed in an earlier section of this paper (page 77), and the writer will reiterate here only his own personal opinion that, since the submentum in generalized insects is attached laterally to the postoccipital tergal region of the head, it comprises the basal parts of the second maxillary appendages, to which, however, there may be added a median field of the sternum of the corresponding seg- ment. If the submentum is regarded as entirely the labial sternum, then the sternum becomes suspended directly from the tergum of its segment, and bears the appendages—a condition so at variance with ordinary morphological relations as to discredit the premises from which it is deduced. VII. THE HEAD OF A CATERPILLAR The caterpillars are remarkable for their standardization of struc- ture. In none of the other larger groups of insects is there such unt- formity in fundamental organization as in the larvae of the Lepidop- tera. Some species are superficially specialized, but apparently there is no “generalized ” caterpillar. Ontogenetically, the caterpillars prob- ably represent a stage below that of the larvae of Neuroptera, and of 132 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 81 the larvae of the more generalized adult Coleoptera (Adephaga), since the young of these insects are closer in form to that of a typical adult insect. The caterpillars show primitive conditions in the origin of the antennal muscles on the walls of the cranium, in the musculature of the thoracic legs, in the monocondylic leg joints, in the dactylopo- dite-like end segments of the legs, and in the retention of the abdominal “legs,” if these organs are remnants of true abdominal appendages, as they appear to be. The general form of the alimentary canal, of the tracheal system, and of the nervous system are fairly generalized, though the brain is specialized by an extreme condensation of its ganglia. On the other hand, the head, the maxillary appendages, the muscle sheath of the alimentary canal, and the body musculature are all highly specialized. While the form of the caterpillar’s body is worm-like, it is not to be supposed that it represents a worm stage or even a primitive stage in the insect ancestry, for the structure of the head shows that the caterpillar belongs to the highly evolved stage of the pterygote insects. The caterpillar’s form is merely one that adapts the insect to a wide feeding environment. The extremely com- plicated body musculature must be regarded as acquired through an excessive multiplication of the segmental muscles to give unlimited mobility to a soft-bodied animal. The fly maggot likewise has an intricate body musculature, but of quite a different pattern from that of the caterpillar. STRUCTURE OF THE HEAD CAPSULE The caterpillar head is an example of the type of head structure in which the lower genal and postgenal regions of the cranium (fig. 51 E) are lengthened to give a long ventro-lateral area on each side between the foramen magnum and the posterior articulation of the mandible. The facial aspect of the head (fig. 50 A) is characterized by the ex- tension of the clypeus into the area of the frons, and by the invagina- tion of the median part of the frons dorsal to the clypeus. The prominent triangular plate so characteristic of the facial aspect of a caterpillar’s head is unquestionably the clypeus (fig. 50 A, B, C, F, H, Clp), though it has usually been called the “ frons.” Its margins are defined internally by a strong V-shaped ridge (FE, I, ER), the in- verted apex of which is continued into a thick median ridge of the dorsal wall of the cranium. From the arms of the V-ridge arise the anterior tentorial apophyses (AT), and the latter identify the V-ridge as the epistomal ridge (ER). The space between the diverging arms, therefore, is the true clypeus (C/p). It has already been shown that the clypeus in other orders of insects may be extended into the facial region dorsal to the mandibular articulations (figs. 46 D, F, G, 47 C). 543 INSECT HEAD—SNODGRASS S65 Further evidence that the area thus designated the clypeus in the lepidopteran larva is the true clypeal area, and not the frons, is given Fic. 50—Head structure of caterpillars: anterior cranial wall, labrum, antenna, and mandibular muscles. A, anterior surface of head of Lycophotia (Peridroma) margaritosa. B, same of Thrydopteryx ephemeraeformis. C, same of Sibene stimulea, showing areas of origin of mandibular muscles (4, 5a). D, antenna of Malacosoma americana, left, anterior view. E, interior view of anterior wall of head of Prionoxystus robiniae (Cossidae), with labral muscles and adductor of left mandible in place. F, anterior surface of head of Mnemonica aurocyanea. G, labrum of Lycophotia margaritosa, anterior view, showing muscle insertions. H, fronto-clypeal area of same. I, inner view of same. AT, anterior arm of tentorium; c, anterior articulation of mandible; C/p. clypeus; ER, epistomal ridge; es, epistomal suture; [’r, frons; fr, “ adfrontal ve fs, frontal suture; fh, submarginal thickening of clypens; Lm, labrum; Md, mandible; Nv, antennal nerve; 7a, antennal trachea. by the origin of the clypeal dilator muscles of the stomodeum upon it (fig. 55, 20, 21). Finally, it is to be observed, the muscles of the la- brum, which, in all cases where the identity of the facial plates is clear, SS pS Te RF LC ag TS RR ee ee 134 SMITHSONIAN MISCELLANEOUS COLLECTIONS voL. 81 arise on the frons, are never attached to the triangular plate of the caterpillar face, but take their origin from the median ridge dorsal to it (fig. 50 B, E, 7). In many caterpillars the lower part of the clyp- eus is strengthened by an internal submarginal thickening (E, I, h) forming a bracing ridge between the articulations of the mandibles (cee): The frontal area of the head, as has been shown, is to be identified by the origin of the labral retractor muscles upon its inner surface (fig. 47 B,C). In the caterpillar the labral muscles arise either upon the median internal ridge of the cranium that extends between the apex of the posterior emargination of the vertex and the apex of the clyp- eus, or upon the dorsal bifurcations of this ridge that are continued into the margins of the vertical emargination (fig. 50 B, EF, 53 FE, 7). This ridge, then, is at least a part of the frons. It is formed by a deep inflection of the median line of the cranium dorsal to the apex of the clypeus, which appears externally as a median suture (fig. 50 A, B, C, H, Fr). Ina softened specimen this frontal invagination can often be widely opened, when it is seen that its inflected surfaces are continuous with the so-called “ adfrontal ” strips lying laterad of the clypeus and extending ventrally to the bases of the mandibles. The sutures, o1 membranous lines, along the outer margins of the “ adfrontals ” thus become the true frontal sutures (fig. 50 A, H, I, fs). The frontal region of the caterpillar, therefore, includes the invag- inated frontal groove (fig. 50 A, E, Fr), the “adfrontals ” (fr), and perhaps the apical margins of the vertical emargination. When the mature caterpillar sheds its skin at the pupal molt, the head cuticula splits along two lines, which, beginning at the notch of the vertex, follow the external lips of the median frontal invagination and then diverge along the “adfrontal ” sutures to the bases of the mandibles. An elongate piece is thus cut out which includes the median frontal inflection, the “‘ adfrontals ’’ and the clypeus. In some caterpillars the molting cleft follows only one of the adfrontal sutures, the other re- maining closed. The median part of the vertex in the caterpillar’s head is obliterated ‘by the dorsal emargination, and the angle of the emargination usually extends into the frontal invagination (fig. 501); in some cases the notch is so deep that the latter is reduced to a very small area dorsal to the apex of the clypeus (F). The labrum of the caterpillar (fig. 50 A, B, Lim) is commonly sep- arated from the lower edge of the clypeus by a wide, flexible membran- ous area. Some writers, having mistakenly identified the true clypeus as the frons, have regarded this membranous area as the clypeus, NO. 3 INSECT HEAD—SNODGRASS 135 but the error of this interpretation is shown by the fact that none of the stomodeal muscles arise upon the membrane, the clypeal dila- tors having their origin on the triangular plate above. The caterpillar labrum has but a single pair of muscles: 1.—Retractor muscles of the labrum (figs. 50 E, G, 53 E) —A pair of long slender muscles arising on the inflected frons (figs. 50 E, 53 E, Fr) ; inserted by long tendons on bases of tormae (figs. 50 G, 53 E) The ventral surface of a caterpillar’s head presents a number of secondary modifications that, at first sight, somewhat obscure the basic structure ; but, when the general head “ landmarks” are once recog- nized, it is not difficult to see that the fundamental structure is no different from that in an orthopteroid head. As we have noted, the caterpillar head is characterized by an elon- gation of the postgenal regions between the foramen magnum, or the end of the neck membrane (fig. 51 E, NMb), and the posterior articu- lations of the mandibles (a). On each side, a posterior median part of the postgena (A, E, Hst) is separated from the more lateral post- genal region (Pge) by a suture (7). The median area thus set off is called the hypostoma (Hst), and the inner angles of the two hypostomal areas are approximated and sometimes united on the median line behind the base of the labium, which is thus separated from its usual basal connection with the neck membrane, or with the postoccipital rim of the cranium. In this manner a condition has been evolved which is almost a replica of that in the head of adult Hymenoptera (fig. 48 B, C), except that in the latter the hypostomal areas are not separated from the rest of the postgenal regions. In some caterpillars a well-developed subgenal ridge (fig. 51 D, Sg) follows the outer margin of the membranous area of the an- tennal base from the anterior articulation of the mandible (c) to the posterior (a), and is then continued along the anterior mesal margin of the hypostoma (Hst). Some entomologists distinguish the part of the subgenal ridge that skirts the mandibular area as the “ pleuro- stomal ridge,’ or “ pleurostoma,” and that part which follows the hypostomal margin as the “ hypostomal ridge.’ The external suture that defines the hypostomal area on each side (E, 7) forms internally a strong ridge (D, 7) extending from the subgenal ridge at the pos- terior mandibular articulation (a) to the postoccipital ridge (Pok). The subgenal ridge, especially its hypostomal part, is lacking or but weakly developed in some caterpillars (C), but the ridge of the hypostomal suture (j) is always well developed, and apparently serves to brace the genal area between the mandible and the posterior rim 136 SMITHSONIAN MISCELLANEOUS COLLECTIONS voL. 81 of the head. The maxillae are suspended in the usual manner by the articulations of the cardines against the margins of the hypostomal areas of the postgenae (C, Cd, E, e). Fic. 51.—Structure of the posterior and ventral parts of the head of a caterpillar. A, postero-ventral view of head of a noctuid (Lycaphotia margaritosa). B, dorsal view of.same. C, interior view of postgenal and hypostomal regions, showing posterior arm of tentorium (P77), and articulation of cardo (Cd). D, inner face of same region in Malacosoma americana. E, ventral view of right half of cranium, with mandible and antenna, of Estigmene acraea. a, posterior articulation of mandible; Ant, antenna; 44p/, base of adductor apodeme of mandible; AT, anterior arm of tentorium; c, anterior articulation of mandible; Cd, cardo; dap, dorsal apodemal plate of postoccipital ridge; e, articulation of cardo to cranium; Fr, frontal ridge; Hst, hypostoma; 7, line of base of neck membrane; 7, hypostomal suture, hypostomal ridge; Lb, labrum; Md, mandible; Mx, maxilla; NMb, neck membrane; Pge, postgena; Polk, post- occipital ridge; PT, posterior arm of tentorium; 7nt, transverse bar of ten- torium; vap, ventral apodemal plate of postoccipital ridge. The foramen magnum is extraordinarily large in the caterpillar, being almost as wide as the cranium, and is extended forward dorsally in the median notch of the vertex (fig. 51 A). The postoccipital ridge (PoR) is inflected from the rear margin of the cranial walls, there NO. 3 INSECT HEAD—SNODGRASS £37 being no perceptible chitinization beyond it to form a postoccipital rim in the neck region. The postoccipital ridge gives origin to plate- like apodemes that constrict the actual opening of the head cavity into that of the neck. Usually there is a pair of dorsal apodemes (A, B, dap) in the notch of the vertex, and a pair of larger ventral apodemes (A, D, E, vap) arising from the postgenal and hypostomal parts of the postoccipital ridge. The apodemes vary much in size and shape in different species, but those of the ventral pair are usually the larger and the more constantly developed. The apodemes furnish surfaces of attachment for the anterior ends of prothoracic muscles inserted on the back of the head (fig. 57 A, C). In the caterpillars the foramen magnum is crossed laterally by oblique foraminal muscles, which are the following: 2.—Muscles of the foramen magnum (figs. 51 E, 57 A).—Attached below on each side to ventral postoccipital apodeme (fig. 51 E, vap) laterad of posterior root of tentorium ; spreading dorsally and laterally, sometimes as a broad fan (fig. 57 A), to the dorso-lateral parts of postoccipital ridge. The foraminal muscles are of the nature of the transverse muscles of the intersegmental folds in the body of the caterpillar. From their position it would appear that they must pro- duce a tension on the hypostomal regions of the head wall. Foraminal muscles are not present in insects generally. The tentorium of the caterpillar is a simple structure consisting of two slender longitudinal bars, and of a delicate transverse posterior bridge. The longitudinal bars, which represent the anterior arms of the tentorium (fig. 53 D, E, AT), arise from the lateral parts of the epistomal ridge at the sides of the clypeus (fig. 50 E, I, AT). They - extend horizontally through the head (fig. 53 E), and are united posteriorly with the ends of the posterior bridge (figs. 51 A, C, E, 53 D, Tnt). The bridge represents the united median parts of the posterior tentorial arms (fig. 51 A, C, PT), the origins of which (EF, pt) are at the posterior angles of the hypostomal plates in the deep inflections that form the inner ends of the ventral postoccipital apo- demes (vap). The positions of all the tentorial roots in the caterpillar, thus, are identical with those of the tentorial roots in an orthopteroid head, notwithstanding the considerable alterations which the surround- ing parts have suffered. THE ANTENNAE The antennae are much reduced in all caterpillars, being so small by comparison with the adult organs that the latter are forced to de- velop by recession, and during the propupal stage their tips only lie within the antennae of the larva. The antennae of the caterpillar are 138 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. OF situated on membranous areas just laterad of the bases of the man- dibles, while the antennae of the adult arise from the facial region above the compound eyes. The ventro-lateral position of the larval antennae, therefore, appears to be a primitive character in the cater- pillars. Each antenna of the caterpillar consists of three segments, of which the middle one is usually the largest, the proximal segment being often reduced to a mere basal ring (fig. 51 E, Ant), and the terminal one appearing as a minute apical papilla of the second. The mem- brane of the antennal base may form a large mound with the antenna retractile into it, or sometimes a long cylindrical projection simulat- ing a basal segment (fig. 50 C). A hypodermal fold projects inward from the base of the antenna (fig. 50 D) which receives the antennal nerve and trachea. Each antenna is moved by a single set of muscle fibers, which are: 3.—The retractor muscles of the antenna (fig. 50 B—F ).—A group of slender fibers arising on the parietal walls of the cranium laterad of adfrontal area ; inserted on anterior inner angle of base of proximal antennal segment. Extension of the antennae is probably effected by blood pressure from within the head. THE MANDIBLES The mandibles of the caterpillar are typical insect jaws suspended from the lower margins of the cranium by a hinge line sloping down- ward posteriorly, with well-developed anterior and posterior articula- tions. The anterior articulation of each mandible consists of a condyle on the cranial margin placed just laterad of the clypeus (fig. 52 A, c), received into a socket on the base of the jaw ; the posterior articulation (a) is the reverse, consisting of a socket on the cranial margin receiv- ing a condyle of the mandible. As in all insects, the articular points of the jaw lie outside the membrane that connects the base of the mandible with the head. A line between the two articulations divides the base of the jaw unequally (fig. 52 B), the larger part being mesad to the axis. The muscles of the mandibles are inserted on large but weakly chitinized apodemal inflections arising at the outer and inner margins of each jaw. The muscles take their origin on the walls of the cranium and on the ventral apodemes of the postoccipital ridge. Their fibers occupy most of the cavity of the head, and the cranial hemispheres appear to model their form on that of the bases of the great adductor muscles of the jaws. NO. 3 INSECT HEAD—SNODGRASS 139 4.—The abductor muscles of the mandible (figs. 50 C, 52 B)—A group of fibers, small by comparison with the adductor group, arising on lower lateral and posterior walls of cranium, and on ventral apo- deme of postoccipital ridge laterad of posterior root of tentorium; fibers converging ventrally, anteriorly, and mesally to insertion on abductor apodeme of mandible. 5.—The adductor muscles of the mandible (figs. 50 C, E, 52 B, 53 E).—An enormous mass of fibers disposed in two sets (figs. 52 B, 53 E, 5a, 5b). The fibers of one group arise from almost entire dorsal, anterior, lateral, and posterior walls of corresponding half of epi- cranium above the ocelli (figs. 50 C, E, 53 E, 5a) ; they converge down- ward upon both surfaces of the broad, adductor apodeme of mandible. The fibers of the other group (figs. 52 B, 53 E, 5b) arise on ventral apodeme of postoccipital ridge (fig. 53 FE, vap) mesad of bases of Fic. 52—Mandibles of a caterpillar. A, mandibles and antennae of Estigmene acraea, ventral view. B, left mandible of a noctuid, with bases of muscles, dorsal view. a, posterior articulation of mandible; Ant, antenna; c, anterior articulation of mandible ; C/p, edge of clypeus ; Mds, mandibles ; 4, abductor muscle of mandible ; 5a, fibers of adductor arising on wall of cranium; 5), adductor muscles arising on ventral apodeme of postoccipital ridge (see fig. 53 E). abductor fibers, and extend horizontally to posterior edge of adductor apodeme of mandible. The obliquity of the mandibular axes causes the points of the jaws to turn upward and somewhat posteriorly during adduction. When the mandibies are closed, the teeth on the cutting edges of the two jaws are opposed to each other (fig. 52 A), not interlocked ; but usually one mandible closes first and its toothed edge passes inside that of the other. Live caterpillars examined by the writer always closed the right mandible over the left, and species of several families preserved in alcohol were found to have the jaws in the same position. THE MAXILLAE AND LABIUM The basal parts of the maxillae and labium are united, and their chitinous areas are reduced or variously broken up into small plates (figs. 51 A, 53 A), which may differ much in different species. With Te I40 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 81 the anterior wall of the labium, apparently, is united also the hypo- pharynx (fig. 54D, Hphy), and the duct of the silk gland opens through a hollow spine, the spinneret, at the tip of the labium. Each maxilla includes a cardinal area (fig. 53 A, Cd), a stipital area (St), both united with the basal part of the labium, and a free terminal lobe (Lc), which appears to be the lacinia. A maxillary palpus is lacking. The area of the cardo includes one principal sclerite (fig. 53 A, B, E, F, Cd), and generally one or two accessory plates (A, E, F,k,k). The principal sclerite is always articulated to the hypostomal margin at a point (e) corresponding with the articulation of the cardo to the cranium in orthopteroid insects. The area of the stipes (St) is variously chitinized, or unchitinized, but it always preserves the ridge (q) of its inner margin, upon which are attached all the stipital muscles. The homology of the terminal lobe of the maxilla is difficult to determine. The musculature of the maxilla of a caterpillar comprises muscles pertaining to its three parts, most of which are comparable to the maxillary muscles of the grasshopper or other generalized insects, though there is little similarity in the general appearance of the struc- ture in the two cases. The cardo, in the caterpillar, is provided with two or three muscles (fig. 53 B, FE, F, 6, 7, 8), all of which arise on the anterior arm of the tentorium (D, E), and, therefore, represent the tentorial adductors of the cardo in orthopteroid insects. The usual cranial muscle of the cardo (fig. 25, J, fig. 40 C, ro) is lacking in the caterpillar. The stipes is provided likewise with tentorial adductors (fig. 53 B, D, E, F, 9, ro, 11) inserted on its mesal chitinous ridge (q). The terminal maxillary lobe is moved by muscles that arise within the stipes (B, F, 12, 73), and also by a long muscle (B, 14) having its origin in the posterior angle of the hypostomal plate (Hst) of the epicranium. These three muscles are inserted upon a basal sclerite in the ventral wall of the maxillary lobe (A, B, /). The first two suggest the ordinary stipital muscles of the lacinia, but the third (14) appears to have no homologue in more generalized insects, since the usual cranial flexor of the lacinia (fig. 30 B, fcc) is inserted on the median angle of the latter. The insertion of the three muscles on a single sclerite in the base of the maxillary lobe leaves no evidence to indicate the presence of a galea, and suggests that the lobe is the lacinia alone, complicated in form by the development of large sensory papillae. Certainly, the musculature of the lobe shows that none of the papillae can be a palpal rudiment. Bois; INSECT HEAD—SNODGRASS I4I Fic. 53—Maxilla, labium, and silk press of a caterpillar. A, Estigmene acraea, maxillae and labium, with hypostomal plates of head, posterior (ventral) view. B, internal view of left maxilla and hypostomal region of same, showing muscles. C, Malacosoma americana, distal part of labium and hypopharynx, lateral view, showing silk press and muscles. D, Lycophotia margaritosa, muscles of maxillae, labium, and hypopharynx, internal (dorsal) view. E, the same, right side of head, internal view, showing muscles of labrum, mandible, maxilla, and labium. F, cardo and lateral parts of stipes of a) maxilla, showing bases of muscles, dorsal (anterior) view. (Compare wit ; a, anterior articulation of mandible; AT, anterior arm of tentorium; Cd, cardo; Clp, clypeus; dap, dorsal apodeme of postoccipital ridge; ¢, articulation of cardo with cranium; Fr, frons; h, submarginal ridge ot clypeus: Hp/ly, hypopharynx; Hst, hypostoma; 7, hypostomal suture; k, accessory plates of cardo; /, basal sclerite of lacinia; Lc, lacinia; Lm, labrum; Md, mandible; Mt, mentum; NMb, neck membrane; PT7, posterior tentorial arm; Pf, posterior tentorial pit; g, ridge on inner edge of stipes; r, articular nodule between end of stipital ridge (q) and mentum; S/D, silk gland ducts; Sim?, submentum; Spt, spinneret; St, stipes; Jt, transverse bar of tentorium; vap, ventral apodeme of postoccipital ridge. 142 SMITHSONIAN MISCELLANEOUS COLLECTIONS voL. 81 The muscles of the maxilla may be enumerated as follows, and they will probably be found to differ but little in different species of caterpillars : 6.—First adductor of the cardo (fig. 53 B, D, E, F).—Origin on posterior end of anterior arm of tentorium (AT); goes ventrally to insertion on base of cardo. 7—Second adductor of the cardo (fig. 53 B, D, E, F).—Origin anteriorly on tentorial arm (D, E) ; insertion on distal end of cardo. 8.—Third adductor of the cardo (fig. 53 D, E, F).—This muscle found in noctuid larvae, perhaps a subdivision of 7. Origin anterior to 7 on tentorial arm (D, E) ; insertion on accessory plate (E, F, k) mesad to the articulating sclerite of cardo (Cd). 9.—First adductor of the stipes (fig. 53 B, D, E, F).—Arises near anterior end of anterior tentorial arm (D, E) ; goes obliquely ventrally and posteriorly to insertion on marginal ridge (B, D, E, F, q) of stipes. to.—Second adductor of the stipes (fig. 53 B, D, E, F).—Origin at anterior end of tentorial arm, just before 9 (D, E) ; insertion on stipital ridge (D, FE, F, q) anterior to 9. 11.—Third adductor of the stipes (fig. 53 B, D, E, F).—Arises pos- teriorly on anterior tentorial arm, just before first adductor of cardo (6) ; goes obliquely ventrally and anteriorly (D, E), internal to 7, 8, 9, and ro, to insertion on anterior end of stipital ridge (B, D, E, F, q) 12—External retractor of the lobe (fig. 53 B, F).—Origin on base of stipital ridge (q); insertion laterally on basal plate (A, B, J) of terminal lobe of maxilla. 13.—Internal retractor of the lobe (fig. 53 B, F).—Origin on base of stipital ridge (q) ; insertion mesally on basal plate (A, B, 7) of terminal lobe of maxilla. 14.—Cranial abductor of the lobe (fig. 53 B).—Origin in basal angle of hypostomal plate of epicranium (Hs?) ; insertion on outer end ‘of basal plate (7) of terminal lobe of maxilla. A corresponding muscle is not present in orthopteroid insects. The labium of the caterpillar (fig. 53 A) lies between the maxillae. The broad membranous surface of its large submental region is united on each side with the marginal ridges (q) of the stipites, and its basal part is continuous laterally with the membrane of the cardinal areas. Proximally the labium may be continuous with the neck mem- brane (NMb) between the approximated ends of the hypostomal plates (Hst), but, when the latter are united, the labium becomes NO. 3 INSECT HEAD—SNODGRASS 143 separated from the neck. A large submental plate occupies the median basal part of the submental region in some species (A, Smt). The distal, free lobe of the labium probably represents the mentum and ligula of other biting insects, combined with the hypopharynx, which forms its anterior surface (fig. 54. A). Evidence of this in- terpretation is found in the fact that the labial and hypopharyngeal Fic. 54.—Distal part of labium, hypopharynx, and silk press of a noctuid caterpillar. A, mentum and hypopharynx, with silk press partly exposed, lateral view. B, the same, dorsal view. C, the same, posterior view, showing support on arms of stipites (q, q). D, lateral view, showing muscle attachments. Hphy, hypopharynx; Mt, mentum; Pr, silk press; q, g, ridges of stipes; 7, r, articular nodules between stipital arms and mentum; SI/D, silk duct; Spt, spinneret. muscles are inserted on the base of the lobe (figs. 53 C, D, 54 C, D, 15, 16), and in the position of the spinneret (fig. 54 A, D, Spt), which contains the opening of the silk duct (salivary duct), the latter being normally situated between the labium and the hypopharynx (fig. 18 DB, SIO). The mental region of the mento-hypopharyngeal lobe appears to be that occupied by the large proximal plate (fig. 53 A, Mt) that em- Io a. 144 SMITHSONIAN MISCELLANEOUS COLLECTIONS voL. 81 braces the base of the lobe ventrally and laterally, but which is not continued across the hypopharyngeal surface (figs. 53 C, 54 A, C, D, Mt). This plate is supported upon the distal ends of the ridges of the stipites (fig. 54 C, D, g, q), which are turned forward and artic- ulated with the dorsal arms of the mentum (V/t) by small, chitinous nodules (7, 7). By this mechanism, the mentum-hypopharynx, which carries the spinning apparatus, is freely movable on a transverse axis between the ends of the supporting stipital ridges. The motion in a vertical plane is the only movement that can be given to the spinning apparatus, except by the action of the entire head; but the head of the caterpillar is highly mobile by reason of the great number of mus- cles inserted upon its posterior margin (fig. 57). The musculature of the mentum-hypopharynx, or spinning organ, is as simple as its mechanism, consisting of two pairs of muscles, as follows: 15.—Reductors of the spinning organ (figs. 53 C, D, E, 54 C, D)— A pair of double muscles arising at posterior ends of tentorial arms (fig. 53 D, E) ; converging ventrally and anteriorly to insertions on ventral edge of mentum (figs. 53 C, E, 54 C, D, Mt). These muscles probably represent the mento-tentorial muscles of orthopteroid insects (fig. 40 D, 23), which are primitive adductors of the second maxillae. 16.—Productors of the spinning organ (figs. 53 C, D, 54 C, D).— A pair of broad muscles arising medially on transverse bridge of ten- torium (fig. 53 D, Tt), diverging ventrally and anteriorly to base of hypopharynx (figs. 53 C, D, 54 C, D, Hphy). These muscles are prob- ably the retractors of the hypopharynx in orthopteroid insects (fig. Al, 32): The silk press of the caterpillar is a special development of the common duct of the labial glands (here, the silk glands). The deeply invaginated dorsal wall of the organ exerts a pressure on the silk ma- terial, which is regulated by two sets of opposing muscles that, prob- ably acting together, effect a dilation of the lumen of the press by elevating the invaginated roof. The muscles of the press arise within the mentum, and the two sets may be distinguished as follows: 17, 18.—Dorsal muscles of the silk press (fig. 54 A, B, C)—Two lateral series of muscles, the number on each side varying in different species of caterpillars, arising on dorsal arms of mentum; converg- ing to insertions on chitinous raphe in dorsal (anterior) wall of press. 19.—Ventral muscles of the silk press (fig. 54 A, B, C).—Origin in ventrolateral parts of mentum; insertion on dorso-lateral edges of silk press. These muscles are antagonists to the dorsal muscles, since the fibers of the two sets oppose each other in the crossed lines of an X NO. 3 INSECT HEAD—SNODGRASS 145 (fig. 54 C) ; but in function the ventral muscles are probably accessory to the dorsals by counteracting the pull of the latter on the press. It is difficult to discover a parallelism between the muscles of the silk press in the caterpillar and muscles of the labium in other insects. However, it may be possible that the two sets of muscles in the labium of the grasshopper (fig. 40 D, 26, 27) inserted on the salivary cup (v) are the prototypes of the silk press muscles, though their insertion points are ventral instead of dorsal. THE STOMODEUM The stomodeum of the caterpillar (fig. 55) is differentiated into four parts. The first part is a bucco-pharyngeal region (BuC, Phy) ; Fic. 55.—Anterior part of the stomodeum of a noctuid caterpillar, showing muscles of the stomodeal wall, and the dilator muscles arising in the head. a-m, muscles of stomodeal wall; BuC, buccal cavity; Cr, crop; OE, oesoph- agus; Phy, pharynx; 20-23, muscles of buccal region, arising on clypeus; 24-27, dorsal dilators of anterior pharyngeal region; 28-30, dorsal dilators of oesophagus (posterior pharyngeal region) ; 31-36, ventral dilators. the second, a cylindrical tube with strong transverse muscle rings, constitutes an oesophagus (OE) in the caterpillars, but it evidently corresponds with the posterior section of the pharynx in Orthoptera ; the third part is the large sack-like crop (Cr) ; the fourth is the con- stricted posterior region of the stomodeum (fig. 56 F, Pvent), which may be termed the proventriculus, though it has no special develop- ment of the lining intima, such as usually distinguishes the proven- tricular region in other insects. The muscular sheath of the entire alimentary canal of the caterpillar is strongly developed, and in some parts becomes highly complicated in structure. The alimentary muscles are particularly strong in the noctuids, and the following descriptions are based mostly on Lyco- photia margaritosa: 146 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 81 The lateral walls of the bucco-pharyngeal region are marked on each side by an oblique ridge (fig. 55), formed by a specially chitinized groove of the intima, which gives a firm line of insertion for the ex- ternal muscles. The latter consist of thick, broad bands of strongly fibrillated muscle tissue, for the most part lying in one plane, though varying in position from transverse to longitudinal. The anterior- most muscles consist of two dorsal ares (a, b), and of a corresponding wide ventral are (d), their ends inserted laterally on the oblique ridges. This part of the stomodeum may be defined as the buccal region because its dilator muscles (20-23) have their origins on the clypeus. The anterior end of the pharyngeal region following is cov- ered dorsally by a broad transverse muscle (c) attached laterally on the oblique ridges. The frontal ganglion lies over the posterior border of this muscle. Each side of the pharynx presents two muscle plaques (e, f) attached to the ventral margins of the upper half of the oblique ridge, but extending posteriorly to the oesophagus. The posterior dorsal wall of the pharynx is covered with several longitudinal mus- cles, the most prominent of which is a wide, median, external band of fibrils (g) deflected from the posterior part of the broad anterior transverse muscle (c). Concealed by this muscle are two longitudinals of a deeper set, arising anteriorly on the buccal region beneath the first transverse muscle (a) and extending posteriorly to the anterior end of the oesophagus. Several superficial longitudinal fibers lie more laterally. The buccal region of the stomodeum is thus distinguished by its strong circular musculature, which evidently gives it a powerful constrictor action. The pharynx is provided principally with longi- tudinal muscles, and its action, except for that produced by the an- terior dorsal transverse muscle, must be one of lengthwise cuntrac- tion. The entire length of the oesophageal tube is sheathed in a close series of strong circular fibers (7) which are complete rings, except a few of the most posterior interrupted dorsally at the anterior end of the crop. The inner walls of the pharynx and oesophagus form four longt- tudinal folds—one dorsal, one ventral, and two lateral. The dorsal fold is broad, flat, and straight-edged. It arises at the base of the labrum, where its margins begin at the tormae, and continues to the posterior end of the oesophagus, where it is lost with the sudden widening of the stomodeal tube in the crop. Between the pharynx and the oesophagus, the continuity of the dorsal fold is interrupted by a transverse fold. The ventral and lateral folds are less definite, rounded NO. 3 INSECT HEAD—SNODGRASS 147 inflections of the stomodeal wall, continuous from the pharynx into the oesophagus. In Lycophotia margaritosa each of these folds ends at the opening of the crop in a prominent fleshy papilla covered with small chitinous points. Between the folds are four deep channels ex- tending from the mouth to the crop, two dorso-lateral, and two latero- ventral. Possibly it is through these channels that the alimentary liquid, which caterpillars frequently eject from the mouth when irritated, is conveyed forward from the crop. The muscles of the crop (fig. 55, Cr) are arranged longitudinally and circularly. The circular muscles (J), except for a few closely placed anterior bands (#), are widely spaced, external circular fibers. They all completely surround the crop like the hoops of a barrel. At the junction of the crop with the oesophagus, there are several short transverse fibers (7) confined to the dorsal surface. All the muscles of the crop are strongly fibrillated (fig. 56 A, B, C, D). The circular bands have distinct nuclei, but nuclei were not observed in the longi- tudinal muscles of noctuid species examined. The longitudinal muscles of the crop (fig. 55, m) have their origin in single fibrillae (fig. 56 A) or small bundles of fibrillae (B) given off from the posterior margins of the circular fibers. They are, there- fore, of the nature of branches of the circular fibers, and this fact may account for their lack of nuclei. Moreover, the longitudinal muscles are not continuous, individual bands, but are everywhere branched and intimately united by intercrossing bundles of fibrillae in such a manner that the entire layer becomes a plexus of muscle tissue (fig. 56C). Most of the fibrillae of this layer spring from the anterior circular fibers, but probably all the circular fibers contribute at least a few elements to the longitudinal plexus. On the anterior end of the crop, the longitudinal fibrillae appear as simple connectives between the transverse fibers (fig. 55, 7). On the posterior end of the crop (fig. 56 F), the longitudinal muscles again break up into smaller fibril bundles, and at last into fine strands that reunite with the external circular fibers of the crop or the proventriculus. The proventricular region (fig. 56 F, Pvent) resembles the oesoph- agus in being surrounded by a close series of strong circular muscle fibers (n). There is no distinct inner muscular sheath here, but the circular fibers are all connected by small bundles of fibrillae going from one to another (G), some to the first neighboring fibers, others to the second, third, or fourth removed in either direction. The proven- triculus has a special feature in the presence of an external layer of fine, widely-spaced, longitudinal muscles, stretched freely between its two ends (fig. 56F, 0). These threadlike strands arise anteriorly es el 148 SMITHSONIAN MISCELLANEOUS COLLECTIONS vot. 81 from branches that spring from the posterior ends of the longitudinal crop muscles, and from the anterior circular fibers of the proventric- ulus. Posteriorly they again break up into branches that are lost in a plexus of fibers at the junction of the proventriculus with the ven- triculus (Vent). A study of the stomodeal muscle sheath of the caterpillar thus shows that the usual brief statement that the insect stomodeum is surrounded by an external layer of circular fibers and an internal layer of longitudinal fibers must be considerably modified and amplified to fit conditions in the caterpillar. The proctodeal muscles of the cater- pillar are even more complicated than are those of the stomodeum. The high degree of development in the alimentary musculature of the caterpillars accords with the general specialization of the caterpillar as an animal most efficient in feeding, and the extreme development of the somatic musculature is only another adaptation to the same end. The dilator muscles of the stomodeum are inserted dorsally and ventrally on the stomodeal walls. The dorsal muscles are grouped into three sets corresponding with the buccal, pharyngeal, and oesophageal regions of the stomodeum. The dilator muscles of the dorsal and central series, enumerated according to the order of their insertions, are as follows: 20.—First dorsal dilators of the buccal cavity (fig. 55).—A pair of slender muscles arising on submarginal ridge of clypeus (fig. 50 I, h) ; extending posteriorly to insertions laterally on roof of mouth cavity just before first band of circular stomodeal muscles. 21.—Second dorsal dilators of the buccal cavity (fig. 55 ).—Origins on clypeus, above middle and close to lateral margins; insertions medially on dorsal wall of mouth cavity between insertions of 20. 22, 23.—Third and fourth dorsal dilators of the buccal cavity (fig. 55).—Two pairs of slender muscles: those of each side arising to- gether in ventral angles of clypeal triangle just above ends of sub- marginal ridge; inserted dorso-laterally on buccal region, 22 before second band of transverse muscles (b), 23 behind it. A wide space occupied by the third transverse muscle band (c) intervenes between the dilators of the buccal region and those of the true pharyngeal region. 24.—First dorsal dilators of the pharynx (fig. 55).—Origin on upper part of clypeus just internal to epistomal ridge; insertion medially on dorsal wall of pharynx laterad of frontal ganglion. These are clearly true pharyngeal muscles; their points of origin have evi- dently crossed the epistomal ridges to the clypeus. INSECT HEAD—-SNODGRASS 149 LAD I TI TAI 4 2 iy LU} pe a il HN) ill - au nk MW Fic. 56.—Muscles of the stomodeum of a noctuid caterpillar. A, B, origin of longitudinal muscles (mm), of crop (see fig. 55) from fibrils deflected from the anterior circular muscles (j, k). C, plexus of longitudinal muscles, anterior part of crop. D, piece of circular fiber from anterior part _ of crop. E, a connecting fiber between circular and longitudinal muscles. F, posterior end of crop (Cr), proventriculus (Pvent), and anterior end of ven- triculus (Vent): 1, m, circular and longitudinal muscles of crop; n, circular muscles of proventriculus; 0, external longitudinal fibers of proventriculus; p, first suspensory muscles of ventriculus. G, parts of seven consecutive circular fibers of proventriculus, showing bundles of uniting fibrils, external. 150 SMITHSONIAN MISCELLANEOUS COLLECTIONS voL. 81 25.—Second dorsal dilators of the pharynx (fig. 55)—Origins on epistomal ridges near union with frontal ridge; insertions dorso- laterally on pharynx. 26.—Third dorsal dilators of the pharynx (fig. 55).—Each arises on cranial wall laterad of origins of antennal muscles ; extends medially, posteriorly, and downward to insertion on pharynx just laterad of 25. The insertions of muscles 24, 25, and 26 all lie posterior to the frontal ganglion connective. 27.—Fourth dorsal dilators of the pharynx (fig. 55).—A group of fibers on each side, arising on outer surface of lower end of frontal ridge; converging to one or two stalks inserted on dorsal wall of pharynx just before brain. The following dorsal muscles are inserted behind the brain and on the region of the stomodeum that may be distinguished in the cater- pillar as the oesophagus, but which is the so-called posterior pharynx in Orthoptera. 28, 29, 30.—Dorsal dilators of the oesophagus (fig. 55)—Three fans of muscles arising on posterior margin of cranial walls on each side of vertical emargination; the spreading fibers inserted dorso- laterally on oesophagus from brain to crop. 31.—First ventral dilators of the pharynx (fig. 55).—A pair of long slender muscles arising on transverse bar of tentorium (fig. 53 D, Tnt), converging to ventral wall of pharynx where inserted just behind first ventral transverse muscle (d). 32, 33.—Second and third ventral dilators of the pharynx (fig. 55). —aA pair of small muscles on each side arising on extreme outer ends of transverse tentorial bar ; fibers spreading at insertion ventro-later- ally on pharynx just before anterior circular muscles of oesophagus. 34, 35, 36.—Ventral dilators of the oesophagus (fig. 55).—Three large fans of fibers arising on postoccipital apodemes on each side laterad of posterior roots of tentorium; the spreading fibers inserted ventro-laterally on oesophagus from circum-oesophageal nerve con- nective to crop. THE MUSCULATURE OF BACK OF HEAD, AND NATURE OF THE INSECT NECK The head of the caterpillar is remarkably mobile. It is provided with a wonderful system of muscles, the fibers of which arise mostly in the prothorax and are distributed at their insertions upon the post- occipital ridge of the head in such a manner as to enable the caterpillar to make all possible head movements of which it conceivably might have need (fie. 57 A, B,C): NO. 3 INSECT HEAD—SNODGRASS ISI The muscles of the prothorax of the American tent caterpillar, Malacosoma americana, are illustrated in figure 57. At A are shown the lateral and ventral muscles as seen from a posterior dorsal view, with the head turned somewhat downward on the neck: B shows the ’ dorsal muscles as seen from below; C presents an inner view of all Fic. 57—Muscles of the prothorax of a caterpillar, Malacosoma americana. A, prothoracic muscles inserted on lateral and ventral parts of back of head, and ventral muscles of mesothorax, posterior view. B, dorsal muscles of pro- thorax and mesothorax inserted on dorsal half of back of head, seen. from below. C, innermost layers of muscles of right half of prothorax, internal view. D, external muscles of right half of prothorax. Isg, intersegmental fold; Zi, base of prothoracic leg; Pok, postoccipital ridge of head; s, edge of tergal plate of prothorax; 1S), first spiracle; Tut, Beverse bar of tentorium; vap, ventral apodemal plate of postoccipital ridge. the muscles in the right half of the prothorax inserted on the head ; and D gives the muscles of the same side that lie external to those shown in C, except the single fiber arising just dorsal to the spiracle, which is shown in both figures. iN I a a es ee Be 152 SMITHSONIAN MISCELLANEOUS COLLECTIONS voL. 81 The various fibers of the head muscles are mostly arranged in groups, and it is easiest to trace them from their points of insertion on the back of the head. Inserted in the median notch of the vertex there is a dorsalmost group of long fibers that diverge posteriorly to the dorsal wall of the prothorax (B, C), the middle fibers of each group going to the posterior margin of the segment. External to these muscles, a group of short fibers, inserted serially on each side, extends posteriorly and dorsally to the tergal plate of the prothorax. Laterally there are inserted on the postoccipital ridge several fibers that spread to their origins on the tergal plate, and a group of four long fibers going dorsally and medially to the intersegmental fold (Isg), with the two median fibers crossing the latter to the dorsum of the mesothorax. Three lateral groups of fibers (A, C) go ventrally and posteriorly from their head insertions, one to the sternal interseg- mental fold, another to the region just before the base of the protho- racic leg, and the third to the median longitudinal fold between the legs. Ventrally there are inserted on the ventral apodeme of the hypostomal region (C, vap) the anterior ends of the ventral longitudi- nal muscles of the prothorax (A, C), and a group of four long fibers on each side that arise on the region above the spiracle. It is of particular interest to observe that, in the caterpillar, the ventral longitudinal muscles of the prothorax are not inserted on the tentorium (fig. 57 A, C) as they are in orthopteroid insects, and fur- thermore, that all the principal longitudinal ventral muscles of the thorax have their origin on the imtersegmental folds, and not on in- trasegmental apophyses. The primitive anterior insertion of these muscles in the prothorax, therefore, should be on a ventral interseg- mental fold between the prothorax and the last head segment. We have already seen that there is evidence of the loss of the true labio- prothoracie intersegmental fold, since the postoccipital ridge, which bears the anterior attachments of the prothoracic muscles in all known insects, appears to be the fold between the maxillary and the labial segments. If so, the original attachments have been lost and the muscles now extend through the length of two primary segments. Furthermore, the attachment of the ventral muscles of the cater- pillar on the hypostomal regions of the head must signify a migration of the muscles from their primitive sternal insertions, for the hypo- stomal lobes clearly belong to the postgenae, and are, therefore, ventral extensions of the tergal area of the head wall. In any case, an attach- ment of the ventral muscles on the bridge of the tentorium certainly represents a farther displacement of the muscle insertions by a final NO. 3 INSECT HEAD—SNODGRASS 153 migration from the tergal postoccipital ridge to the posterior ten- torial apophyses. The question of the morphology of the cervical region of the insect must yet remain a puzzle ; but the musculature gives no evidence of the existence of a neck segment. On the other hand, the fold in the integument of the caterpillar between the neck (fig. 57D, Cv) and the prothoracic tergum (T,) is suggestive of being the true interseg- mental line between the labial segment and the prothoracic segment, and several muscles of the prothorax have their anterior attachments upon it (D). If the primitive insect is conceived as a continuously segmented, vermiform animal, the neck, or any other secondary inter- segmental area, must be a part of a primary segmental region. From the evidence at hand it seems more probable that the region of the insect neck belongs to the labial segment, than to an anterior part of the prothoracic segment. ABBREVIATIONS USED ON THE FIGURES Ab, abdomen. abplp, abductor of palpus. adplp, adductor of palpus. Am, amnion. AMR, anterior mesenteron rudiment. An, anus. Ant, antenna. AntNv, antennal nerve. Ao, aorta. AP, apical plate. AR, antennal ridge. Arc, archicerebrum. as, antennal suture. AT, anterior arm of tentorium. at, anterior tentorial pit. BC, body cavity. Bdy, body. Blc, blastocoele. Bld, blastoderm. Bp, blastopore. 1Br, protocerebrum. 2Br, deutocerebrum. 3Br, tritocerebrum. Bs, basisternum. BuC, buccal cavity. CA, corpus allatum. Cd, cardo. Cer, cercus. Ch, chelicera. Cho, chorion. Clp, clypeus. CoeCon, circumoesophageal connective. Com, commissure. 3Com, commissure of tritocerebral lobes. Con, connective. cs, coronal suture. ct, coxo-trochanteral joint. Cth, cephalothorax. Cv, neck, cervix. cv, cervical sclerite. C%, coxa. dap, dorsal apodemal plate of postoc- cipital ridge. DMcl, dorsal longitudinal body muscle. DWNvz, dorsal longitudinal nerve. DT, dorsal arm of tentorium. dt, attachment of dorsal tentorial arm to wall of cranium. E, compound eye. Ecd, ectoderm. End, endoderm. Endp, endopodite. Ephy, epipharynx, epipharyngeal sur- face. Eps, episternum. CY > 154 SMITHSONIAN MISCELLANEOUS COLLECTIONS ER, epistomal ridge. es, epistomal suture. Exp, exopodite. F, femur. faa, flexor of galea. Fl, flagellum. fic, flexor of lacinia. ficc, cranial flexor of lacinia. fics, stipital flexor of lacinia. For, foramen magnum, or “ occipital ” foramen. Fr, frons. fr, ‘ adfrontal ”’ FrGng, frontal ganglion. fs, frontal suture. ft, femoro-tibial joint. Ga, galea. GC, gastric caecum. Gc, gastrocoele, archenteron. Gch, gnathochilarium. Ge, gena. GI, glossa. Gn, gnathal segments. Gnec, gnathocephalon. Gng, ganglion. Gu, gula. H, head. Hphy, hypopharynx. Hst, hypostoma. I, tergal promotor muscle of an appen- dage. I-VI, segments of the head. Isg, intersegmental fold. J, tergal remotor muscle of an appen- dage. K, sternal promotor muscle of an ap- pendage. KL, ventral adductor muscles. KLh, ventral adductors arising on hy- popharynx. KLk, ventral adductors united by liga- ment (k) forming ‘ dumb-bell muscle.” VoL. 81 KLt, ventral adductors arising on ten- torium, or hypopharyngeal apo- demes. L, leg. rL, first leg. 1, prothoracic leg. sternal remotor muscle of an appen- dage. LB, primitive limb base (coxa and subcoxa). Lb, labium. LbNv, labial nerve. Ibmcl, labial muscles. Le, lacinia. Lm, labrum. LNuvz, lateral stomodeal nerve. Md, mabdible. MdC, mandible cavity. MdNv, mandibular nerve. Ment, mesenteron. Mps, mouth parts. Msb, primary mesoblast. Msc, mesenchyme. Msd, mesoderm. Mst, metastomium. Mt, mentum. Mth, mouth. Mx, maxilla. IM x, first maxilla. 2M x, second maxilla. M-xC, maxilla cavity. MaNv, maxillary nerve. NC, nerve cord. NMb, neck membrane. N ph, nephridium. O, ocellus. levator muscle of palpus, or of tro- chanter. Oc, occiput. OcR, occipital ridge. ocs, occipital suture. OE, oesophagus. OeGng, oesophageal, or posterior me- dian stomodeal ganglion. OpL, optic lobe. OR, ocular ridge. os, ocular suture. NO. 3 INSECT HEAD—SNODGRASS T55 P, thoracic depressor muscle of tro- chanter. PcR, posterior cranial ridge. Pdc, pedicel. Pdp, pedipalp. Pge, postgena. Pgl, paraglossa. Ph, phragma. Phy, pharynx. PLGng, posterior lateral stomodeal ganglion. Pip, palpus Pnt, postantennal appendage. Poc, postocciput, postoccipital rim of foramen magnum. PoR, postoccipital ridge. pos, postoccipital suture. Pp, “pleuropodium,” specialized ap- pendage of first abdominal seg- ment. Ppd, parapodium. Ppt, periproct. PrC, preoral cavity. Pre, protocephalon. Prnt, preantennal appendage. Proc, proctodeum. Prst, peristomium. Prtp, protopodite. Pst, prostomium. PT, posterior arm of tentorium. pt, posterior tentorial pit. Ptar, praetarsus. Q, depressor muscle of palpus, or of trochanter. Rd, posterior fold of tergum. rh, retractor of hypopharynx. RNv, recurrent nerve. SA, sternal apophysis. Scp, scape. Scx, subcoxa. Ser, serosa. Set, seta, setae. SgR, subgenal ridge. sgs, subgenal suture. SID, salivary duct, silk gland duct. STO, opening of salivary duct. Smt, submentum. SeoGng, suboesophageal ganglion. Sp, spiracle, rSp, first thoracic spir- acle. Spn, spina. Spt, spinneret. St, stipes. Stom, stomodeum. T, tergum. depressor muscle of tibia. Tar, tarsus. Tb, tibia. Th, thorax. TI, tentacle. Tlp, telopodite. Tnt, tentorium. Tor, torma. Tr, trochanter. V’, fifth head segment. vap, ventral apodemal plate of postoc- cipital ridge. VI, sixth head segment. V Mcl, ventral longitudinal body mus- cle. V NC, ventral nerve cord. V Nv, ventral longitudinal nerve. Va, vertex. REFERENCES Arttems, C. (1926). Myriapoda. Krumbach’s Handbuch der Zoologie, 4. Ber- lin, Leipzig. ; Batrour, F. M. (1880). Notes on the development of the Araneina. Quart. Journ. Micr. Sci., n. s., 20: 1-23, pls. 19-21. (1883). The anatomy and development of Peripatus capensis. Quart. Journ. Micr. Sci., n. s., 23: 213-259, pls. 13-20. Bascu, S. (1865). Untersuchungen itber das Skelet und die Muskeln des Kopfes von Termes flavipes (Kollar). Zeit. wiss. Zool., 15: 56-75, pl. 5. 156 SMITHSONIAN MISCELLANEOUS COLLECTIONS voL. 81 BERLESE, A. (1909). Gli Insetti, 1. Milan. (1910). 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Zur Kenntnis der Anatomie und Entwicklungsge- schichte der Stabheuschrecke Carausius morosus Br. III. Entwicklung und Organogenese der Célomblasen, pp. 123-328, 86 figs. Published by Zool.- vergl. anat. Inst. Univ. Zurich. Yuasa, H. (1920). The anatomy of the head and mouth-parts of Orthoptera and Euplexoptera. Journ. Morph., 33: 251-200, 9 pls. ZocraF, N. (1883). Contributions to a knowledge of the embryological de- velopment of Geophilus ferrugineus L. K. and Geophilus proximus L. K. (in Russian). Studies Lab. Zool. Mus. Moscow Univ., 2, pt. 1, 77 pp. 108 figs. ie NIAN MISCELLANEOUS COLLECTIONS VOLUME 81, NUMBER 4 UES LEMOYNE DE MORGUES OF ATURIOUA, A TIMUCUA CHIEF (Wirn ONE PLATE) _ DAVID I. BUSHNELL, Jr Ccececese?= __GITY OF WASHINGTON JED BY THE SMITHSONIAN INSTITUTION AUGUST 23, 1928 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLUME 81, NUMBER 4 DRAWING BY JACQUES LEMOYNE DE MORGUES OF SATURIOUA, A TIMUCUA CHIEF IN FLORIDA, 1064 (WiTH ONE PLATE) BY DAVID I. BUSHNELL, Jr (PUBLICATION 2972) CITY OF WASHINGTON PUBLISHED BY THE SMITHSONIAN INSTITUTION AUGUST 23, 1928 The Lord Battimore Press BALTIMORE, MD., U. 8. As DRAWING BY JACQUES LEMOYNE DE MORGUES OF SATURIOUA, A TIMUCUA CHIEF IN FLORIDA, 1564 BYEDAVID Ss: BUSENE EE Re (WitH ONE PLATE) When it became known in Europe that a new continent had been discovered beyond the sea, that the lands reached by Columbus and his companions did not form part of Asia but were a new and distinct region, wonder was aroused as to the sort of people who were to be found in the strange and unknown country. So great was the interest thus manifested that many narratives of early voyages contain accounts of the natives encountered along the coasts and some refer, all too briefly, to the manners and customs of the Indians, a term erroneously applied to the inhabitants of the New World. Many records are preserved of natives having been taken to Europe by the explorers. It is written that when the Cabots—first to reach the con- tinent of North America—returned to England in the year 1497, they carried three of the strange people from the newly discovered lands, and that four years later Cortereal compelled others to return with him to Europe. Likewise when Jacques Cartier reached France in 1535, after exploring the great River St. Lawrence, he had on board his small vessel a native chief taken in the wilderness. This tends to prove that many were eager to learn about the people who lived in the mysterious region far to the westward, beyond the sea. With this evidence of interest in the people of the New World it is difficult to believe that pictures were not made of them; sketches or paintings to portray their peculiar customs, strange ornaments and dress, and frail habitations. But no drawings are known to have been made dur- ing the voyages of the Cabots, of Ponce de Leon, Varrazano, Narvaez, de Soto, or Cartier. No proof that any pictures of Indians of North America were made during the first half of the sixteenth century has been discovered. And although the account of the voyages of Cartier, as presented by Ramusio, is accompanied by several crude illustra- tions, there is no evidence to indicate that the drawings were made by a person who had visited Canada. Thus it would appear that not until the year 1564, when the French expedition led by Laudonniere set SMITHSONIAN MISCELLANEOUS COLLECTIONS, VoL. 81, No. 4 2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL. 81 sail from Havre de Grace for the Land of Florida, did an artist accompany an expedition for the definite purpose of making drawings to be taken back to Europe. Consequently Jacques Lemoyne de Morgues, artist, who accompanied Laudonniere, made the earliest known pictures of Indians of North America. Many sketches were undoubtedly made by the artist during the eventful year he remained in Florida but only one original example of his work can now be traced, this being a drawing of the great chief Saturioua who claimed the land on which the French erected Fort Carolina. JACQUES LEMOYNE DE MORGUES Very little is known of the life and career of the artist who ac- companied Laudonnieére to Florida. He appears to have been a man of culture and learning. He was a Huguenot and seems to have been known personally by Charles IX. He prepared a brief Narrative of events in Florida which was printed by Theodoro de Bry, in the year 1591, as the second part of Grand Voyages. Together with this text were the engraved reproductions of 42 drawings made by Lemoyne revealing scenes in Florida, the natives, their habitations, and events of interest." To quote from the English translation of Lemoyne: “ Charles IX, King of France, having been notified by the Admiral de Chatillon that there was too much delay in sending forward the re-enforcéments needed by the small body of French whom Jean Ribaud had left to maintain the French dominion in Florida, gave orders to the admiral to fit out such a fleet as was required for the purpose. The admiral, in the mean while, recommended to the king a nobleman of the name of Renaud de Laudonniere; a person well known at court, and of varied abilities, though experienced not so much in military as in naval affairs. The king accordingly appointed him his own lieutenant, and appropriated for the expedition the sum of a hundred thousand frances.”’ The Narrative continues: ‘I also received orders to join the expedition, and to report to M. de Laudon- niere . . . . I asked for some positive statements of his own views, and of the particular object which the king desired to obtain in com- *Two works have been quoted in preparing these notes: a. History of the First Attempt of the French to Colonize the Newly Discovered Country of Florida. By Rene Laudonniére. In His- torical Collections of Louisiana and Florida. By B. F. French. New York, I&60. b. Narrative of Le Moyne, an Artist who accompanied the French Expedition to Florida under Laudonniére, 1564. Translated from the Latin of De Bry. Boston, 1875. NO. 4 JACQUES LEMOYNE DE MORGUES BUSH NELL 3 manding my services. Upon this he promised that no services except honorable ones should be required of me; and he informed me that my special duty, when we should reach the Indies, would be to map the seacoast, and lay down the position of towns, the depth and course of rivers, and the harbors; and to represent also the dwellings of the natives, and whatever in the province might seem worthy of observa- tion: all of which I performed to the best of my ability, as I showed his majesty, when, after having escaped from the remarkable perfidies and atrocious cruelties of the Spaniards, I returned to France.” The three vessels of the expedition sailed from Havre de Grace April 20, 1564. Their first stop was at the Canaries, thence they sailed to the West Indies. At one island, “called Dominica, we watered. Making sail again, we reached the coast of Florida, or New France as it is called, on Thursday, June 22.” They had arrived off the mouth of the River of May, the present St. Johns. Soon ascending the stream a few miles they selected a site where Fort Carolina was erected. Lemoyne was in Fort Carolina September 20, 1565, when it was attacked and taken by the Spaniards. He fled and wandered through the swamps several days before meeting Laudonniére and some fifteen others who had escaped the massacre. Later they reached the mouth of the river, boarded one of the small ships and made sail for France, “all manned and ill provisioned. But God, however, gave us so fortunate a voyage, although attended with a good deal of suffering, that we made the land in that arm of the sea bordering on England which is called St. George’s Channel.” Now to quote from Laudonniere’s record. The first of the three ships to return to France departed from Florida July 28, 1564. About November 10, 1564 “ Captain Bourdet determined to leave me, and to return to France.” During the summer of 1565 the French were visited by the English Admiral Hawkins. Laudonniére, with his small party including the artist Lemoyne, sailed from Florida September 25, 1565. “‘ About the 25th of October, in the morning, at the break of day, we described the /sle of Flores, and one of the Azores, where, immediately upon our approaching to the land, we had a mighty gust of wind, which came from the north- east, which caused us to bear against it four days; afterwards, the wind came south and south-east, and was always variable. In all the time of our passage, we had none other food saving biscuit and water.” About November 10, 1565, they reached the coast of Wales and landed, having been carried out of their course and thus failed to reach France. They had landed at Swansea. Laudonniere then wrote: 4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 81 “ For mine own part I purposed, with my men, to pass by land; and, after I had taken leave of my mariners, I departed from Swansea, and came, that night, with my company, to a place called Morgan, where the lord of the place, understanding what I was, staid me with him for the space of six or seven days; and, at my departure, moved with pity to see me go on foot, especially being so weak as I was, gave me a little hackney. “Thus I passed on my journey—first to Bristol, and then to London, where I went to do my duty to M. de Foix, which, for the present, was the King’s ambassador, and helped me with money in my necessity. From thence I passed to Calais, afterward to Paris, where I was informed that the king was gone to Moulins, to sojourn there ; incontinently, and with all the haste I could possibly make, I got me thither, with part of my company.” Lemoyne was probably one of the company, and it may have been at this time that he revealed to the king the work he had done in Florida. How long Lemoyne continued to live in France is not known but later he crossed the channel and resided in London. He was a Hugue- not and for that reason may have sought safety in flight. During 1587 Lemoyne was in London, in the service of Sir Walter Raleigh, when he was visited by De Bry in the endeavor to purchase his papers relat- ing to the expedition to Florida, but as has been written: “ Lemoyne resisted all persuasions to part with his papers. After Lemoyne’s death De Bry bought them of his widow (1588), and published them in 1591. What became of Lemoyne’s drawings is not known. Possibly those secured by De Bry were taken to Frankfort and there copied by the engravers, later to be lost or scattered. No example of the artist’s work is in the British Museum, London; the Louvre, Paris; or the Galleria degli Uffizi, Florence. It may be suggested that Lemoyne’s connection with Sir Walter Raleigh influenced the latter in sending the English artist John White to Virginia, in 1585. White’s instructions were quite similar to those received by Lemoyne some twenty years before. Their work was of the same nature. SATURIOUA RE DELLA FLORIDA Saturioua was a Timucua chief whose tribe claimed and occupied territory on both sides of the St. John River, from its mouth inland for some distance as well as up and down the coast. NO. 4 JACQUES LEMOYNE DE MORGUES—BUSH NELL 5 During the summer of 1564, while Fort Carolina was being con- structed by Laudonniere, “ several chiefs visited our commander, and signified to him that they were under the authority of a certain king named Saturioua, within the limit of whose dominions we were, whose dwelling was near us, and who could muster a force of some thou- sands of men.” Saturioua soon desired to see the work being done by the French and visited the site chosen for the fort. “ He sent forward, however, some two hours in advance of his own appearance, an officer with a company of a hundred and twenty able-bodied men, armed with bows, arrows, clubs, and darts, and adorned, after the Indian manner, with their riches; such as feathers of different kinds, necklaces of a select sort of shells, bracelets of fishes’ teeth, girdles of silver-colored balls, some round and some oblong; and having many pearls fastened on their legs. Many of them had also hanging to their legs round flat plates of gold, silver, or brass, so that in walking they tinkled like little bells. This officer, having made his announcement, proceeded to cause shelter to be erected on a small height near by of branches of palms, laurels, mastics, and other odoriferous trees, for the accom- modation of the king.” And soon the great chief arrived, “ accom- panied by seven or eight hundred men, handsome, strong, well-made, and active fellows, the best-trained and swiftest of his force, all under arms as 1f ona military expedition.” The meeting proved one of great interest to both French and Indian. Laudonniére made known to Saturioua that he had been “sent by a most powerful king, called the King of France, to offer a treaty by which he should become a friend to the king here, and to his allies, and an enemy to their enemies ; an announcement which the chief received with much plea- sure. Gifts were then exchanged in pledge of perpetual friendship and alliance.’ The Indians soon departed, but the French hastened with greater energy the completion of the fort. Some days passed and the time arrived when Saturioua desired to test the sincerity of the French. “The chief sent messengers to M. de Laudonnieére, not only to confirm the league which had been made, but also to procure the performance of its conditions, namely, that the latter was to be the friend of the king’s friends, and the enemy of his enemies; as he was now organizing an expedition against them.” A vague, ambiguous reply was received by the messengers and by them carried to Saturioua. The great chief then visited the fort, accompanied by a large number of men. He attempted to have the French go with him on his expedition against his enemies farther up the river but they declined. “ Failing, however, to obtain what he 6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 81 wished, he set out on his expedition with his own men. While these affairs were in progress, M. de Laudonniére sent his second ship, com- manded by Pierre Capitaine, to France.” Saturioua, surrounded by his chiefs and warriors, preparing to start on the expedition, was the subject of a drawing by Lemoyne, one engraved by De Bry, but the description of the picture as given by the artist is really more complete than the reference just quoted. The description of the engraving, given by De Bry, was evidently prepared by Lemoyne himself. The English translation is now quoted: Frc. 1.—Ceremonies performed by Saturioua before going on an expedition against the enemy. From De Bry, 1501. “ It is mentioned in the account of the second voyage that the French made a treaty of friendship with a powerful chief of the vicinity, named Saturioua, with agreement that they were to erect a fort in his territory, and were to be friends to his friends, and enemies to his enemies ; and, further, that on occasion they should furnish him some arquebusiers. About three months afterwards, he sent messengers to Laudonniere to ask for the arquebusiers according to the treaty, as he was about to make war upon his enemies. Laudonniére, however, sent to him Capt. La Caille with some men, to inform him courteously that he could not just then supply any soldiers, for the reason that he hoped NO. 4 JACQUES LEMOYNE DE MORGUES—BUSH NELL 7 to be able to make peace between the parties. But the chief was in- dignant at this reply, as he could not now put off his expedition, having got his provisions ready, and summoned the neighboring chiefs to his aid; and he therefore prepared to set out at once. He assembled his men, decorated, after the Indian manner, with feathers and other things, in a level place, the soldiers of Laudonniére being present ; and the force sat down ina circle, the chief being in the middle. A fire was then lighted on his left, and two great vessels full of water set on his right. Then the chief, after rolling his eyes as if excited by anger, uttering some sounds deep down in his throat, and making various ges- tures, all at once raised a horrid yell; and all his soldiers repeated this yell, striking their hips, and rattling their weapons. Then the chief, taking a wooden platter of water, turned toward the sun, and wor- shipped it; praying to it for a victory over the enemy, and that, as he should now scatter the water that he had dipped up in the wooden platter, so might their blood be poured out. Then he flung the water with a great cast up into the air; and, as it fell down upon his men, he added, ‘ As I have done with this water, so I pray that you may do with the blood of your enemies.’ Then he poured the water in the other vase upon the fire, and said, “So may you be able to extinguish your enemies, and bring back their scalps.’ Then they all arose, and set off by land up the river, upon their expedition.” Laudonniere wrote regarding these happenings: ‘‘ About two months after our arrival in Florida, the Paracoussy Saturioua sent certain Indians unto me to know whether I would stand to my promise, which I had made him at my first arrival in that country: which was, that I would show myself friend to his friends, and enemy unto his enemies ; and, also, to accompany him with a good number of harque- buses, when he should see it expedient, and should find a fit occasion to go to war.”’ Laudonniére declined to join his forces with those of Saturioua and the latter departed on the war-like expedition without the promised aid of the French. Laudonniére then continued his nar- rative: “ The ceremony which this savage used, before he embarked his army, deserveth not to be forgotten; for, when he was sitting down by the river’s side, being compassed about with ten other paracoussies, he commanded water to be brought him speedily. This done, looking up into heaven, he fell to discourse of divers things, with gestures that showed him to be in exceeding great choler, which made him one while shake his head hither and thither ; and, by and by, with, I wot not what fury, to turn his face towards the country of his enemies, and to threaten to kill them. He oftentimes looked upon the sun, praying him to grant him a glorious victory of his enemies; which, when he 8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 81 had done, by the space of half an hour, he sprinkled, with his hand, a little of the water, which he held in a vessel, upon the heads of the paracoussies, and cast the rest, as it were, in a rage and despite, into a fire, which was there prepared for the purpose. This done, he cried out, thrice, He Thimogoa! and was followed with five hundred In- dians, at the least, which were there assembled, which cried, all with one voice, He Thimogoa!” These events transpired during the latter part of August, 1564. SATURIOUA—DRAWING BY LEMOYNE The original drawing now reproduced for the first time, is in crayon—black and sanguine. It bears a legend in Italian which reads: Saturioua Re della Florida nell’ America Settertionale in atto di andare alla Guerra. Translated it is: ‘* Saturioua King of Florida in North America in the act of going to war.” This evidently shows the chief immediately after the completion of the ceremony mentioned on preceding pages. He has grasped his spear but continues to hold the wooden bowl containing water. Details are revealed in the drawing with great clearness. Several of these may be explained by quoting from Lemoyne’s notes attached to various sketches reproduced by De Bry. Describing the peculiar ear ornament represented as being worn by Saturioua, Lemoyne wrote : “ All the men and women have the ends of their ears pierced, and pass through them small oblong fish-bladders, which when inflated shine like pearls, and which, being dyed red, look like a light-colored car- buncle.”’ Tattooing was practiced extensively and “all these chiefs and their wives ornament their skins with punctures arranged so as to make certain designs. . . . . Doing this sometimes makes them sick for seven or eight days. They rub the punctured places with a certain herb, which leaves an indelible color.” But the strangest of their customs, “ For the sake of further ornament and magnificence, they let the nails of their fingers and toes grow, scraping them down at the sides with a certain shell, so that they are leit very sharp. They are also in the habit of painting the skin around their mouths of a blue color.” Elsewhere Lemoyne wrote: “ They let their nails grow long both on fingers and toes, cutting the former away, however, at the sides, so as to leave them very sharp, the men especially; and, when they take one of the enemy, they sink their nails deep in his forehead, and tear down the skin, so as to wound and blind him.” Such were some of the strange and curious customs of the people of Florida more than three and one-half centuries ago. SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 81, NO. 4, PL. 1 #2 dtiacued ped Ss ES aaa ne a WE os Some ee er cae eee Size 10 by 7 inches SATURIOUA NO. 4 JACQUES LEMOYNE DE MORGUES—BUSH NELL 9 Unfortunately the history of this very interesting drawing, now in the author’s collection, is not known; however, it is possible to reach certain conclusions regarding its origin. The legend is in Italian and this offers a clue as to the time the picture was actually made. The youthful Charles IX was King of France in 1564, the year of the French expedition under Laudonniére to Florida, but all were dominated by the Queen-mother, Catherine de’ Medici, surrounded as she was by groups of Italians who had accompanied or followed her to France. Italian was spoken at the French Court. Lemoyne had accompanied the expedition to Florida for the pur- pose of preparing a series of drawings and sketches, these, as he him- self wrote: “ I showed his majesty, when, after having escaped from the remarkable perfidies and atrocious cruelties of the Spaniards, | returned to France.” And it may be assumed that all such work, when exhibited at Court, bore legends written in Italian. The draw- ing of Saturioua Ke della Florida, may have been one of the sketches thus displayed. It is not possible to determine exactly when the drawing was made. Knowing the manner in which the artist escaped from Fort Carolina the night it was taken by the Spaniards there is no reason to believe he was able to save any drawings. All his possessions appear to have been abandoned and lost. The event of Saturioua starting for war, the subject of the drawing now reproduced, occurred late in August, I 564. Little more than two months later, early in November, the second of the French vessels returned to France. Undoubtedly it carried dispatches and various papers relating to the progress of affairs in the Colony, and quite likely sketches and drawings by the artist of the expedition, Lemoyne, were included with the official reports. The drawing of the chief with whom the French were then in contact, and who claimed the region in which they had settled, may have been sent to France at that time. Again it 1s suggested that the picture may have been made after Laudonniére and his small party, including Lemoyne, had returned to Europe but before they had reached Moulins and met the king. The fact that the legend on the drawing is in Italian, not French, would seem to prove beyond doubt that it was so written for the benefit of Charles IX, the Queen-mother, and their Italian followers and associates. SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLUME 81, NUMBER 5 THE RELATIONS BETWEEN THE SMITHSONIAN INSTITUTION AND THE WRIGHT BROTHERS : BY t GHARLES G. ABBOT é Secretary, Smithsonian Institution (PUBLICATION 2977) a GITY OF WASHINGTON | PUBLISHED BY THE SMITHSONIAN INSTITUTION SEPTEMBER 29, 1928 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLUME 81, NUMBER 5 THE RELATIONS BETWEEN THE SMITHSONIAN INSTITUTION AND THE WRIGHT BROTHERS BY GHARLES G. ABBOT Secretary, Smithsonian Institution (PUBLICATION 2977) CITY OF WASHINGTON PUBLISHED BY THE SMITHSONIAN INSTITUTION SEPTEMBER 29, 1928 + ae a 2 xa » a 2s , o = £ oe s es) a eR eo ou 2 a » < oO b : a = a $ 2 & a < a PREFATORY-NOEE This statement represents an attempt on the part of the Smithsonian Institution to clarify an unfortunate con- troversy, and to correct errors where errors have been made, in order to do justice alike to three great pioneers of human flight—Wilbur and Orville Wright, and Samuel Pierpont Langley—as well as to the Smithsonian Insti- tution. VEAP GR EE ATIONS BED WEE NG iit (SMITE SONTAN INST EEULION AND, THE WiRIGHEE BROTHERS By CHARLES: G ABBOm SECRETARY, SMITHSONIAN INSTITUTION For several months past, beginning February 13, 1928, when I first addressed Mr. Orville Wright, a month after my election as Secretary, I have sought to end the so-called Langley-Wright controversy. In a friendly, personal con- ference with Mr. Orville Wright on April 19, he explained to me the points regarding which he feels that the Smith- sonian Institution has dealt unjustly with the Wright brothers, and stated that what he termed a “ correction of history ” by the Smithsonian was essential. So far as lam aware, all men agree that on December 17, 1903, at Kitty Hawk, North Carolina, Orville and Wilbur Wright, alternately piloting their plane, made the first sus- tained human flights in a power propelled heavier-than-air machine. These successful flights by the Wright brothers came as the culmination of: (1) Their extensive laboratory experi- ments to determine the behavior of plane and curved sur- faces in air. (2) Their numerous gliding flights during several years at Kitty Hawk and elsewhere. (3) Their original design and construction of their flying machine and of the engine and propellers. The Smithsonian Institution has recognized these achievements in the following manner: 1. By printing articles by Wilbur and Orville Wright in the Smithsonian Annual Reports. (See Smithsonian An- nual Reports, 1902, pp. 133-148; 1914, pp. 209-216.) SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 81, No. 5 2 SMITHSONIAN MISCELLANEOUS COLLECTIONS voL. 81 2. By printing other articles descriptive of their achieve- ments. (See Smithsonian Annual Reports, 1903, pp. 170- 180; 1908, p. 133; IQIO, pp. 147-151, 160-161.) 3. By making the first award of the Langley gold medal for aeronautics to Wilbur and Orville Wright. This award was made on February 10, 1909, and the medal was for- mally presented on February to, 1910. (See Smithsonian Annual Reports, 1000, pp: 22, 107, 11l; 1O1o;, pp 22-227 104-110.) 4. By formal vote of the Board of Regents, March 15, 1928, as follows: Wuereas, To correct any erroneous impression derived from published statements that the Smithsonian Institution has denied to the Wright brothers due credit for making the first successful human flight in power-propelled heavier-than-air craft ; Resolved, That it 1s the sense of the Board of Regents of the Smithsonian Institution that to the Wrights belongs the credit of making the first successful flight with a power-propelled heavier- than-air machine carrying a man. 5. By requesting the Wright brothers to furnish for ex- hibit in the National Museum the originals or models of any planes made by the Wrights up to 1910, the selection to be at their discretion. (The request specifically included the Kitty Hawk plane. See pages 5 and 6 following, for letters of Secretary Walcott to Wilbur Wright of March 7, 1910, and April 11, TOmos) 6. By exhibiting in the National Museum the plane flown at Fort Myer in 1908 by Orville Wright, which is the first airplane bought for military purposes by any government. 7. By exhibiting since 1922 in the National Museum twelve double-sided frames containing forty-nine photo- graphs showing the circumstances of the Kitty Hawk and Fort Myer flights. Mr. Wright feels, however, that the Smithsonian Insti- tution has appeared to be engaged in propaganda with the NO. 5 SMITHSONIAN INSTITUTION AND WRIGHT BROTHERS 3 object of exalting Langley at the expense of himself and his brother as follows: 1. By predominant mention of the achievements of Langley in the addresses at the time of the first presenta- tion of the Langley medal. 2. By a misleading account of the exercises of Febru- ary 10, 1910, printed in the Smithsonian Annual Report of 1910. 3. By what he regarded as the lack of cordiality in an invitation by Secretary Walcott in April, 1910, to the Wright brothers to deposit the Kitty Hawk or other planes in the U. 5. National Museum. 4. By the contract, in 1914, for experiments with the Langley machine made with Mr. Glenn Curtiss, at that time a defendant in a patent suit brought by the Wright brothers. s, By claims of priority in capacity to fly, for the Langley machine, based on the experiments of 1914, and repeated in Smithsonian publications as well as on labels in the National Museum. 6. By failure to recognize properly the abilities of the Wrights as research men. I propose to take up these points seriatim: 1. Mr. Wright’s feeling that predominant mention of the achievements of Langley was made at the presenta- tion of Langley medals to him and his brother. The main address on February 10, 1910, was by the late Dr. Alexander Graham Bell, a friend of Langley, a close observer of his experiments for a period of ten years, anda Regent of the Smithsonian Institution. The occasion was the first award of a gold medal bearing Langley’s name, which had been established at the suggestion of Dr. Bell to perpetuate Langley’s place in aeronautics. Responding to a feeling then prevalent that Langley, on account of the FE) SUES TA ALA See 1 See Smithsonian Annual Report, 1910, PP. 104-108. 4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 81 ill success of his experiments of 1903, had met with unjust ridicule, and doubtless inspired also by the partiality of a friend, it cannot be denied that Dr. Bell made less promi- nent in comparison with Langley’s achievements the suc- cessful pioneer work of the Wrights than he might well have done appropriately on that occasion. But Dr. Bell was not lacking in appreciation of the Wrights. In the following letter recommending establishment of the Lang- ley medal he suggests the fitness of awarding it to the Wright brothers: Beinn Bhreagh, Near Baddeck, Nova Scotia, December 5, 1908. Hon. C. D. Walcott, Secretary, Smithsonian Institution, Washington, D. C. Dear Secretary Walcott: The Wright brothers are being deservedly honored in Europe. Can not America do anything for them? Why should not the Smithsonian Institution give a Langley medal to encourage avia- tion? Yours, sincerely, ALEXANDER GRAHAM BELL. (See Smithsonian Annual Report, 1909, p. 107.) By refer- ence to the same Report ° it will be seen also how strongly Senator Lodge felt in regard to the merits of the Wright brothers. 2. Mr. Wright’s feeling that the summary of the exercises of February 10, 1910, printed in the Smithsonian Annual Report of 1910 was misleading. I acknowledge with regret that the summary of the pro- ceedings given at an earlier page of the Smithsonian An- nual Report for 1910 (pp. 22-23) is misleading. The sum- mary quotes the following words from Mr. Wilbur Wright: * Smithsonian Annual Report, 1900, p. I1T. NO. 5 SMITHSONIAN INSTITUTION AND WRIGHT BROTHERS 5 “The knowledge that the head of the most prominent sci- entific institution of America believed in the possibility of human flight was one of the influences that led us to under- take the preliminary investigation that preceded our active work. He recommended to us the books which enabled us to form sane ideas at the outset. It was a helping hand at a critical time, and we shall always be grateful.” From the context it would appear that Mr. Wright made this statement at the ceremony. This was not the case. Actually the statement was quoted by Dr. Bell in his speech from an extract of a private letter from the Wright brothers which Dr. Octave Chanute had quoted at the Langley Me- morial meeting, December 3, 1906. The full statement made by Wilbur Wright at the ceremony is given as ap- proved by him at pages 109-110 of the same Smithsonian Annual Report, that for 1910. Mr. Orville Wright assures me that though he and his brother both drew encouragement from the fact that so celebrated a scientific man as Dr. Langley had adventured his reputation in the field of heavier-than-air aviation, the Wrights did not rely on Langley’s experimental data or conclusions, but made laboratory researches of their own, on which their constructions were based exclusively. I fully accept this assurance as a true statement of historical fact. 3. Mr. Wright's feeling that Secretary Walcott’s invita- tions to deposit the Kitty Hawk and other planes in the National Museum lacked cordiality. The letters referred to are as follows: Smithsonian Institution, Washington, U. S. A., March 7, IgIo. My dear Mr. Wright: The National Museum is endeavoring to enlarge its collections illustrating the progress of aviation and, in this connection, it has “See Smithsonian Miscellaneous Collections, Vol. XLIX, Art. IV, Publ. No. 1720, p. 32. SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL. 81 been suggested that you might be willing to deposit one of your machines, or a model thereof, for exhibition purposes. The great public interest manifested in this science and the numerous inquiries from visitors for the Wright machine make it manifest that if one were placed on exhibition here it would form one of the most interesting specimens in the national collections. It is sincerely hoped that you may find it possible to accede to this request. With kindest regards, I am Very truly yours, Cuar_es D. WALcorTT, Mr. WILBUR WRIGHT, Secretary. Dayton, Ohio. Dayton, Ohio, March 26, 1910. Mr. Charles D. Walcott, Washington, D. C. My dear Dr. Walcott: Your letter of the 7th of this month has been received. If you will inform us just what your preference would be in the matter of a flier for the National Museum we will see what would be possible in the way of meeting your wishes. At present nothing is in con- dition for such use. But there are three possibilities. We might construct a small model showing the general construction of the airplane, but with a dummy power plant. Or we can reconstruct the 1903 machine with which the first flights were made at Kitty Hawk. Most of the parts are still in existence. This machine would occupy a space 40 feet by 20 feet by 8 feet. Or a model showing the general design of the latter machine could be constructed. Yours truly, WicLsur WRIGHT. Smithsonian Institution, Washington, U. S. A., April 11, 1910. Dear Mr. Wright: Yours of March 26th came duly to hand, and the matter of the representation of the Wright airplane has been very carefully con- sidered by Mr. George C. Maynard, who has charge of the Division of Technology in the National Museum. I told him to indicate what he would like for the exhibit, in order that the matter might NO. 5 SMITHSONIAN INSTITUTION AND WRIGHT BROTHERS 7 be placed clearly before you and your brother. In his report he says: The following objects illustrating the Wright inventions would make a very valuable addition to the aeronautical exhibits in the Museum: t. A quarter-size model of the airplane used by Orville Wright at Fort Myer, Virginia, in September, 1908. Such a model equipped with a dummy power plant, as suggested by the Wrights, would be quite suitable. 2. If there are any radical differences between the machine referred to and the one used at Kitty Hawk, a second model of the latter machine would be very appropriate. 3. A full-size Wright airplane. Inasmuch as the machine used at Fort Myer has attracted such world-wide interest, that machine, if it can be repaired or reconstructed, would seem most suitable. If, however, the Wright brothers think the Kitty Hawk machine would answer the pur- pose better, their judgment might decide the question. 4. If the Wright brothers have an engine of an early type used by them which could be placed in a floor case for close inspection that will be desirable. The engine of the Langley Aerodrome is now on exhibition in a glass case and the original full-size machine is soon to be hung in one of the large halls. The three Langley quarter-size models are on exhibition. The natural plan would be to install the different Wright machines along with the Langley machines, making the exhibit illustrate two very important steps in the history of the aeronautical art. The request of Mr. Maynard is rather a large one, but we will have to leave it to your discretion as to what you think it is practicable for you to do. Sincerely yours, Cuartes D. WALCOTT, Mr. WILBur WRIGHT, Secretary. 1127 West Third Street, Dayton, Ohio. I cannot but feel that Mr. Wright has erred in ascribing to Dr. Walcott any but a sincere invitation to the Wrights to make their own selection of whatever they thought best suited and most available to deposit in the National Museum for the purpose of illustrating their achievements. It is to be recalled, too, that in 1910 the world was ringing with the triumphant demonstrations of the Wrights at Fort Myer and in France of ability to make long-continued air flights. At that moment the Fort Myer plane was far more cele- 8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 81 brated than the Kitty Hawk plane. Now, of course, all is changed. We have the Fort Myer plane. But it is pro- foundly regretted by patriotic Americans that the Kitty Hawk plane is not in a place of honor in the United States National Museum. 4. Mr. Wright’s feeling that the contract to test the Langley plane in 1914 with Mr. Glenn Curtiss, then a defendant in a suit with the Wrights, was un- friendly to them. I concede to Mr. Wright that it lacked of considera- tion to put the tests of the Langley plane into the hands of his opponent, Mr. Curtiss. As early as 1908 Dr. Walcott had had correspondence with Mr. Manly and with Dr. Chanute on the desirability of further experiments with the Langley Aerodrome under Manly’s direction. Lack of means, from which the Smithsonian then as now suffered, doubtless stood in the way. Without having been familiar myself with all the circumstances at that time, I believe it was owing to the fact that Mr. Curtiss had the available plant and Manly had not, so that the former could make the tests at smaller expense than the latter, that Dr. Walcott determined to place the machine in Curtiss’ hands for trial. The Smithsonian paid Mr. Curtiss $2,000 to make the ex- periments. Yet the fact that the results of these tests might prove valuable to Mr. Curtiss in his defense against Mr. Wright’s suit, and the unfavorable aspect in which that might put the Smithsonian Institution, if foreseen, might well have deterred from the course of action adopted. The appointment of Dr. A. F. Zahm to represent the Smith- sonian as official observer at the Hammondsport tests has been criticized. At that time Dr. Zahm, a recognized aero- nautic authority, was the official recorder of the Langley Aeronautical Laboratory of the Smithsonian Institution, a position he had held since May, 1913, so that his appoint- ment as indicated was natural. NO. 5 SMITHSONIAN INSTITUTION AND WRIGHT BROTHERS 9 As to the propriety of testing Langley’s machine in 1914, some have objected on the ground that it was a precious specimen, taken from the National Museum to be wantonly subjected to destruction. This is not true. The machine, excepting its engine, was never on public exhibition until 1918. In 1904 it was specifically placed by the War Depart- ment * at the disposal of the Smithsonian for further tests. It had been kept continuously in the shops where it was made from the winter of 1903 until it was taken to Ham- mondsport. In 1914 airplane construction had not reached the com- paratively standardized stage of the present day. It was then thought possible that the tandem, dragon-fly type of the Langley Aerodrome had merits which should be devel- oped. There was also the thought that a decisive success might rescue from unmerited ridicule Langley’s fame. These, I submit, were circumstances very properly inviting the making of the tests. But I feel that it was a pity that Manly, Dr. Langley’s colleague, could not have been the man chosen to make them. 5. Mr. Wright’s feeling that claims in priority of capacity to fly for the Langley machine based on 1914 experi- ments were unjustified and prejudicial to the Wright brothers. The claims published by the Smithsonian relating to the 1914 experiments at Hammondsport were sweeping. In the Report of the U. S. National Museum for 1914, page Agee 1Tt is frequently erroneously stated that the Congress appropriated $100,000 to Langley for his experiments. The sum of $50,000 allotted to him by the Board of Ordnance and Fortifications of the War Department was all the public money that he ever had for the purpose. There was no direct Congres- sional appropriation whatever. 2See also Smithsonian Annual Report, 1914, pp. 9-10 and 217-222; also the label of the full-sized Langley machine as first installed in 1918 in the Na- tional Museum, hereafter quoted. IO SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 81 we read: “ Owing to a defect in the launching apparatus, the two attempts to fly the large machine during Dr. Lang- ley’s life proved futile, but in June last, without modifica- tion, successful flights were made at Hammondsport, N. Y.”’ Certainly this was not literally true, but Assistant Sec- retary Rathbun, who wrote the statement given above, I am certain believed this to be true. There were, however, many differences. (I refer only to the first tests when the original Langley-Manly engine was used.) Mr. Wright claims that essential changes tending to improve the chances of success were made on the basis of knowledge gained subsequent to 1903. Some of the differences were favorable, some unfavor- able, to success. Just what effects, favorable or unfavor- able, the sum total of these changes produced can never be precisely known. In the opinion of some experts, the tests demonstrated that Langley’s machine of 1903 could have flown, and in the opinion of others, these tests did not demonstrate it. It must ever be a matter of opinion. In 1918, the Langley plane, reconstructed as nearly as possible as of 1903, using all available original parts, by Mr. R. L. Reed, the foreman who had most to do with it 1n Langley’s time, was exhibited in the U. S. National Museum with this label: THE ORIGINAL, PULL-SIZE. LANGLEY FLYING MACHINE, 1903 Later this label was amplified to read as follows: ORIGINAL LANGLEY FLYING MACHINE, 1903 THE FIRST MAN-CARRYING AEROPLANE IN THE HISTORY OF THE WORLD CAPABLE OF SUSTAINED FREE FLIGHT. INVENTED, BUILT, AND TESTED OVER THE POTOMAC RIVER BY SAMUEL PIERPONT LANGLEY IN 1903. SUCCESSFULLY FLOWN AT HAMMONDSPORT, N. Y., JUNE 2, IQ1T4. DIMENSIONS: 55 FEET LONG, 48 FEET WIDE; SUSTAINING WING SUR- FACE, 1,040 SQUARE FEET. NO. 5 SMITHSONIAN INSTITUTION AND WRIGHT BROTHERS 1 Vigorous criticism of the statements made by the Smith- sonian relative to the test of 1914, and the capability of flight of Langley’s machine having appeared, Dr. Walcott im 1925 asked Dr. J. S. Ames and Admiral David W. Taylor, members and now Chairman and Vice-Chairman, respectively, of the National Advisory Committee for Aero- nautics, to examine the circumstances and report. Their conclusions were summarized in the following letter, sup- ported by several appendices which are printed herein, the whole of which was given to the press by Dr. Walcott on June 9, 1925. Washington, D. C., Juners 1925: Dr. Charles D. Walcott, Secretary, Smithsonian Institution, Washington, D. C. Dear Doctor Walcott : The announcement that Mr. Orville Wright had arranged to have the first Wright airplane deposited in a British museum having aroused considerable controversy as to the accuracy of the label attached to the Langley flying machine now on exhibition in the Smithsonian Institution, you have asked us to examine the Langley machine, look into its history, and advise you whether we consider it desirable to modify the present label. We have made a careful study, not only of the history of the Langley machine itself, but also of all the circumstances connected with its tests. We append to this letter (Appendix I) a suggested modified label, and a statement of our views and conclusions ( Ap- pendix I1), upon which our recommendation is based. There is no question but that the Wrights were the first to navi- gate the air, thus reaching the goal long sought by many, but in our opinion, when Langley’s 1903 machine was wrecked in launching, he too, after years of effort, following a different road, was in sight of the same goal. He was like the prophet of old who, after forty years of wandering in the wilderness, was permitted to view the promised land upon which he never set his foot. Langley’s accom- plishments in aeronautics were notable, and he is entitled to full credit for them. We believe that the Langley machine of 1903 was capable of sustained flight had it been successfully launched, and it is naturally fitting that the Smithsonian Institution should SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 81 perpetuate with pride, by exhibiting his models and flying machine, suitably labeled, the aeronautical achievements of its distinguished secretary. It is unfortunate that in the past the situation has been beclouded by patent litigation, in which the Smithsonian Institution had no part, involving temptation for one side to exaggerate and distort favorably Langley’s work, and for the other side to belittle and deny it. While bitterness thus engendered survives, it cannot be expected that any label can be placed upon Langley’s machine that will be fully acceptable to everyone. The appended suggested label departs from the customary brief title in two respects. In the first place, it is much longer and goes more into the history of the exhibit than is customary. In the second place, in view of the facts that: the exhibit deals with the border line between success and failure of man’s effort to fly, and that the original Wright machine, a purely American product and the first to fly, is destined to a museum in another country, we have suggested that the iabel on the Langley machine, also a purely American product and capable of flight but not successfully flown, contain an explicit and definite statement, which would be unnecessary under other circumstances, giving to the Wrights the credit due them as the first to fly, on December 17, 1903. It is our earnest hope that this proposed restatement of the label will prove satisfactory both to yourself and to Mr. Orville Wright, with both of whom we have had such friendly relations on the Na- tional Advisory Committee for Aeronautics and in whose judgment and fairness of mind we have such implicit confidence. Respectfully yours, (Signed) Jos—EpH S. AMES Professor of Physics, Johns Hopkins University. (Signed) D. W. Tay tor Rear Admiral (C. C.) U.S. N., Retired. NO. 5 SMITHSONIAN INSTITUTION AND WRIGHT BROTHERS 13 APPENDIX I (Ames-Taylor Report) LABEL LANGLEY FLYING MACHINE THE ORIGINAL LANGLEY FLYING MACHINE OF 1903, RESTORED. IN THE OPINION OF MANY COMPETENT TO JUDGE, THIS MACHINE WAS THE FIRST HEAVIER-THAN-AIR CRAFT IN THE HISTORY OF THE WORLD CAPABLE OF SUSTAINED FREE FLIGHT UNDER ITS OWN POWER, CARRYING A MAN. THIS MACHINE SLIGHTLY ANTEDATED THE WRIGHT MACHINE DESIGNED AND BUILT BY WILBUR AND ORVILLE WRIGHT, WHICH, ON DECEMBER 17, 1903, WAS THE FIRST IN THE HISTORY OF THE WORLD TO MAKE A SUSTAINED FREE FLIGHT UNDER ITS OWN POWER. CARRYING A MAN. Langley’s machine was designed by Samuel Pierpont Langley, Secretary of the Smithsonian Institution, and completed in 1903. The original machine was never successfully launched into the air: attempts at launching with a catapult on October 7 and December 8, 1903, were failures owing to defects in the operation of the catapult launching device, and the machine was damaged severely. In 1914, using all available parts remaining, the machine was re- constructed, with certain modifications, and with hydroplane floats attached for the purpose of enabling it to rise from the water in- stead of being launched by a catapult. In that condition, and carry- ing a man, it was successfully flown with the original power plant. at Hammondsport, New York, June 2, 1914, and photographed in flight. With a modified and more powerful power plant, it was subsequently flown repeatedly. These tests indicated that the original airplane would have flown if successfully launched in the tests of 1903. After the Hammondsport flights the pontoons were removed and the airplane was restored in accordance with original drawings and data to its original condition, and is constructed in the main of the original parts. 14 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 81 Washington, D. C., Jane 32,1925. APPENDIX. 1 (Ames-Taylor Report) THE LANGLEY FLYING MACHINE. Memorandum for Dr. Charles D. Walcott, Secretary, Smithsonian Institution, 1. In connection with our letter to you of even date, concerning the label on the Langley Flying Machine in the National Museum, we beg to add the following remarks of an historical nature, and our views and conclusions in some detail. 2. Professor S. P. Langley became actively interested and en- gaged in the study of aeronautics in 1887, and was assiduous in the theoretical and experimental study of the subject till his death in 1906. The more important of his results were finally published in Volume 27 of “ Smithsonian Contributions to Knowledge,” Part I, issued in 1891, entitled “* Experiments in Aerodynamics ” ; Part 2, the “ Internal Work of the Wind,” 1893; and Part 3, the “Langley Memoir on Mechanical Flight,’ 1911. In the course of his study he became convinced of the possibility of “ mechanical flight,” 7. ¢., of constructing a heavier-than-air machine, to be driven by an engine, and sufficiently powerful and stable to carry a man. To this end he constructed certain models about 12 feet wide by 15 feet long, weighing approximately 30 pounds, each driven by a 14 horsepower steam engine which with its boiler weighed not over 7 pounds per horsepower. These models actually did fly, in one case as long as 1 minute and 49 seconds and for a distance of 4,300 feet. These two machines made _ successful flights on May 6, 1896, in the presence of Dr. A. Graham Bell, and on November 28, 1896, in the presence of Mr. Frank G. Carpenter. The model machines numbered 5 and 6 were placed on exhibition in the National Museum on April 21, 1g05. Finally, by the aid of a grant of $50,000 made by the Board of Ordnance and Fortification of the War Department in December, 1898, which was later sup- plemented by funds to the amount of $20,000 from the Smithsonian Institution, he constructed in the years from 1898 to 1903 a full- size flying machine (which he called an “ aerodrome’’), a repro- duction on a scale approximately 4: 1 of these steam models which had previously flown in 1896. The engine of this final machine was a radial 5 cylinder, water cooled, gasoline type, 5 inch bore by 53 inch stroke, developing 52.4 horsepower at g50 r. p. m., and NO. 5 SMITHSONIAN INSTITUTION AND WRIGHT BROTHERS 15 weighing 125 pounds, or 2.2 pounds per horsepower. This engine was designed and built by Mr. Charles M. Manly at the Smith- sonian shops. Two tests were attempted with this flying machine, Mr. Manly being the pilot in both cases. 3. The machine was designed to obtain its initial impetus by means of a spring-catapult propelling it along a pair of rails on top of a house boat. The first test was conducted in the middle of the Potomac River, opposite Widewater, Virginia; and suitable pro- vision was made for the flotation of the machine upon its landing on the surface of the river as it was intended to do. The second test was made on December 8, 1903, off the Arsenal Point in the Potomac River at the junction of the Georgetown Channel and the Eastern Branch. A full description of the machine and the tests is given in “‘ Langley Memoir on Mechanical Flight,” published in 1911. Both attempts to launch the machine failed. The first on October 7, 1903, failed because a lug on a pin projecting from the bottom of the lower front guy post hung in its slot on a support on the launching car or catapult, causing the front wings to be badly twisted from a positive angle of lift to a negative angle of depression, thus forcing the front end of the machine downwards instead of supporting it, and resulting in the machine striking the water about 150 feet in front of the house boat from which it was launched. The front wings and propellers were broken by the im- pact and the rear wings and control surfaces were destroyed by towing the machine through the water. The second test on Decem- ber 8, 1903, failed for reasons which were never absolutely deter- mined. Photographs of the operation show clearly, however, that the immediate cause was the collapse of the rear part of the machine. This was probably due to a sudden gust of wind striking it and throwing it against a stanchion as it passed down the launching track, while it was still in contact with the catapult. Thus, no evi- dence was obtained of the aerodynamic or other features of the machine itself. Further study at the time was not possible because funds were exhausted and the public prejudice against the work made it impossible for Dr. Langley to raise either public or private funds. 4. The machine was drawn from the water in its damaged con- dition the night of December 8, 1903. A few days later it was re- moved to the shops of the Smithsonian Institution where the frame was repaired and the engine, which had not been injured, was stored for further use till such time as additional funds might be- come available to build new wings and to defray the expenses of 16 SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL. 81 further tests. Official disposition of that part of the machine belong- ing to the War Department was made on March 23, 1904, when by formal letter of the Board of Ordnance and Fortification, signed by Major General G. S. Gillespie, President of the Board, and addressed to Dr. Langley, the Board stated that“... . all of the material procured for experiments with the aerodrome from allot- ments of this Board will be left in your possession, in order that it may be available for any future work which you may be able to carry on in the solution of the problem of mechanical flight ; unless, of course, the Board of Ordnance and Fortification shall otherwise direct, but until such action be taken there will be no necessity for a separation or distribution of the property so far as the Board is concerned.” 5. It would seem from the above that at that time there was expectation that further tests would be made with the machine. 6. The machine had in the meantime been cleaned and restored to its original condition, except for the necessary wings and con- trol surfaces. The ribs and cloth covering on the original wings and control surfaces had been so damaged as to require replacement, but the metal fittings were all saved for rebuilding the wings when it might become possible. 7. The engine was shipped to New York in 1906 and exhibited at the first aeronautical show which was held at the Grand Central Palace by the Aero Club of America. It was then returned to Washington and placed on temporary exhibition in the National Museum, but the rest of the machine remained in the Smithsonian shops and was not then placed on exhibition in the National Museum. 8. It appears that as early as 1908 the Smithsonian Institution contemplated making further tests with the Langley Flying Machine. This is evident from a memorandum of September 14, 1908, signed by Cyrus Adler, addressed to Mr. Rathbun, at the Smithsonian Institution, which reads as follows: “September 14, 1908. “For Mr. Rathbun: ‘Thad a talk today with Mr. Chanute, the gist of which I should like to put on record. “He spoke of Mr. Manly’s desire to fly Mr. Langley’s flying machine just as it was constructed in order to demonstrate that it could have flown. Mr. Chanute said that in his opinion Mr. Lang- ley’s machine could fly just as it was constructed, and this had been NO. 5 SMITHSONIAN INSTITUTION AND WRIGHT BROTHERS 17, demonstrated by the fact that a Frenchman has built a machine exactly like Mr. Langley’s which has flown, but he believed further that the machine would be wrecked in alighting. “T thought you might care to have this because it is more than likely that before very long, through the War Department or in some other way, the question of trying the machine will be forcibly brought up. Very truly yours, Cyrus ADLER.” This is further evidenced by the following correspondence between Dr. Walcott and Dr. Octave Chanute, one of the pioneers in flying experiments : “ November 16, 1908. “Dear Dr. Chanute: ‘In a letter received during the summer while I was away from the city, Mr. Charles M. Manly says: The Langley machine is today capable of more than any other machine yet built, and is apt to remain so for some time. The engine is now seven years old and still is the peer of the world. “Mr. Manly has suggested that he be permitted to make trial tests of the Langley machine at some future time. I write to ask whether in your judgment it would be wise to have an attempt made to fly with it. Sincerely yours, Cuas. D. WA cortt.” * Chicago, Illinois, November 20, 1908. “Mr. Chas. Walcott, Secy., Smithsonian Instn., Washington, D. C. Dear. Sir: ‘‘T have your letter of the 16th, asking whether in my judgment, it would be wise to make an attempt to fly with the Langley machine. “ T have never seen this machine but I suppose that I understand it fairly well from descriptions. “My judgment is that it would probably be broken when alight- ing on hard ground and possibly when alighting on the water, al- though the operator might not be hurt in either case. “Tf the Institution does not mind taking this risk and suitable arrangements can be made about the expense. I believe that it 18 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 81 would be desirable to make the test, in order to demonstrate that the Langley machine was competent to fly and might have put our gov- ernment in possession of a type of flying machine, which, although inferior to that of the Wrights, might have been evolved into an effective scouting instrument. Yours truly, O. CHANUTE.” “November 27, 1908. , Dear Sirk “ T wish to thank you for your letter of November 21, in relation to the Langley machine. I will talk the matter over with Mr. Manly the next time I see him. Very truly yours, Cuas. D. WaLcortT.” ‘* Doctor OcTAVE CHANUTE, 61 Cedar Street, Chicago, Illinois.” g. In 1910, the Smithsonian Institution made an effort to secure the original Wright machine of 1903, or a model thereof for ex- hibition in the National Museum. This is evidenced by the follow- ing correspondence between Dr. Walcott and Mr. Wilbur Wright : * Smithsonian Institution, Washington, U. S. A., March 7, 1910. “ My dear Mr. Wright: “The National Museum is endeavoring to enlarge its collections illustrating the progress of aviation and, in this connection, it has been suggested that you might be willing to deposit one of your machines, or a model thereof, for exhibition purposes. “The great public interest manifested in this science and the numerous inquiries from visitors for the Wright machine make it manifest that if one were placed on exhibition here it would form one of the most interesting specimens in the national collections. It is sincerely hoped that you may find it possible to accede to this request. “With kindest regards, | am Very truly yours, CHARLES D. Watccott, “Mr. WILBUR WRIGHT, Secretary.” Dayton, Ohio.” NOWS5 SMITHSONIAN INSTITUTION AND WRIGHT BROTHERS 19 ‘Dayton, Ohio, March 26, 1910. ‘“Mr. Charles D. Walcott, Washington, D. C. “My dear Dr. Walcott: “Your letter of the 7th of this month has been received. If you will inform us just what your preference would be in the matter of a flier for the National Museum we will see what would be possible in the way of meeting your wishes. At present nothing ‘s in condition for such use. But there are three possibilities. We might construct a small model showing the general construction of the airplane, but with a dummy power plant. Or we can recon- struct the 1903 machine with which the first flights were made at Kitty Hawk. Most of the parts are still in existence. This machine would occupy a space 40 feet by 20 feet by 8 feet. Or a model show- ing the general design of the latter machine could be constructed. Yours truly, Wivpur WRIGHT.” “ Smithsonian Institution, Washington, U.S. A., April 11, 1910. “Dear Mr. Wright: “ Yours of March 26th came duly to hand, and the matter of the representation of the Wright airplane has been very carefully considered by Mr. George C. Maynard, who has charge of the Division of Technology in the National Museum. I told him to indicate what he would like for the exhibit, in order that the matter might be placed clearly before you and your brother. In his report he says: The following objects illustrating the Wright inventions would make a very valuable addition to the aeronautical exhibits in the Museum: 1. A quarter-size model of the airplane used by Orville Wright at Fort Myer, Virginia, in September, 1908. Such a model equipped with a dummy power plant, as suggested by the Wrights, would be quite suitable. 2. If there are any radical differences between the machine referred to and the one used at Kitty Hawk, a second model of the latter machine would be very appropriate. 3. A full-size Wright airplane. Inasmuch as the machine used at Fort Myer has attracted such world-wide interest, that machine, if it can be repaired or reconstructed, would seem most suitable. If, however, the Wright brothers think the Kitty Hawk machine would answer the pur- pose better, their judgment might decide the question. 20 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 81 4. If the Wright brothers have an engine of an early type used by them which could be placed in a floor case for close inspection that will be desirable. “ The engine of the Langley Aerodrome is now on exhibition in a glass case and the original full-size machine is soon to be hung in one of the large halls. The three Langley quarter-size models are on exhibition. The natural plan would be to install the different Wright machines along with the Langley machines, making the exhibit illustrate two very important steps in the history of the aeronautical art. ‘The request of Mr. Maynard is rather a large one, but we will have to leave it to your discretion as to what you think it is prac- ticable for you to do. Sincerely yours, CHARLES D. WaLcorTT, Sectetaryas “Mr. WiLBur WRIGHT, 1127 West Dhird: Street, Dayton, Ohio.” 10. Apparently, nothing developed from the above correspon- dence. Dr. Walcott’s last letter quoted above was never replied to. It is a matter of grave regret that at that time the Wright brothers did not see their way to reconstruct and deposit in the National Museum their original full-size airplane, the first machine ever to fly successfully with a man, because then, in 1910, it would have been the only full-size flying machine on exhibition in the National Museum, the Langley machine being still in the shops of the Smith- sonian Institution awaiting further tests. 11. In September, 1911, the Smithsonian Institution secured and placed on exhibition in the National Museum the original Wright airplane that was tested at Fort Myer in 1908, and purchased by the War Department, being the first military airplane purchased by the Government. 12. In January, 1914, the late Lincoln Beachey, one of the pioneer aviators, and others, again suggested that it would be of interest to determine by actual test whether the essential features of Professor Langley’s aerodynamic theory, as illustrated in his 1903 machine, were correct. Finally, at the initiative of the Smithsonian Insti- tution, the Curtiss Aeroplane Company was invited to submit a bid to refit the machine and to make tests. The formal letter to the Curtiss Aeroplane Company was dated March 31, 1914, and the reply offering to undertake the work for a price of $2,000, was NO. 5 SMITHSONIAN INSTITUTION AND WRIGHT BROTHERS 21 written by Mr. G. H. Curtiss on April 1, 1914. The machine was thereupon sent to the shops of the Curtiss Aeroplane Company at Hammondsport, New York, on April 2, 1914, and the engine was shipped on April 13, 1914. 13. In preparing the machine for flight with the original engine, certain modifications and additions were made. These were due, in the main, to the fact that, whereas the original machine was fitted for use with a catapult, these new tests were to be made from the surface of a lake, using hydroplaning floats. Therefore, certain changes were necessary to attach these floats to the machine and to properly inter-brace them and the supporting surfaces together. 14. It is perfectly clear from the correspondence between the Smithsonian Institution and the Curtiss Aeroplane Company that no emphasis was placed upon the use of the original machine, as such, but that what was desired was knowledge concerning certain features of the Langley design, which was expressed in Dr. Wal- cott’s letter of March 31, 1914, previously referred to, in the fol- lowing terms: “Tn connection with the reopening and development of work under the Langley Aerodynamical Laboratory, it seems desirable to make a thorough test of the principles involved in the construction of the Langley heavier-than-air man carrying flying machine, espe- cially the question as to the tandem arrangement of the planes, and general stability, especially longitudinal stability.” 15. A brief interesting account of the Hammondsport tests is contained in the Annual Report of the Smithsonian Institution for IQI4, pages 217 to 222. 16. After the flights were discontinued in November, 1915, the machine was returned to the Smithsonian shops on June 26, 1916. There it was completely overhauled. New wings and control sur- faces were built to the same form and size (with solid instead of hollow ribs to save the expense of the latter) so as to refit the machine for exhibition purposes in the National Museum and restore it as nearly as possible to its original condition as it was in 1903. As much of the original material was used as possible. When this overhaul was completed, it was placed on exhibition in the National Museum on January 15, 1918. 17. It is seen that up to 1915 the Langley machine was used solely and properly for the purposes intended by Professor Langley himself, for which it was originally turned over by the Board of Ordnance and Fortification which had defrayed the major portion of its cost. When all had been done to this end that was possible, ho No SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 81 the machine became properly an exhibit in the National Museum. It was never an exhibit until 1918. 18. Previous to this date, there had been placed on exhibit in the Museum the two Langley steam-driven models which had success- fully flown in 1896, and the quarter-size model of the large machine equipped with its 3 horsepower radial gasoline engine. The first two of these are approximately, and the latter exactly, one-fourth the linear dimensions of the full-size machine. It is thus clear that, when in the letters from the Smithsonian Institution to Messrs. Wilbur and Orville Wright, of March 7 and April 11, 1gto, the request was made for models of their successful machines, it was the hope to have both Langley and the Wright brothers represented in the Museum by exhibits of the same character. 19. The question whether the original Langley machine of 1903 was capable of flight under its own power and carrying a pilot has been a controversial one since, subsequent to the Hammondsport trials of 1914, there was litigation to which the Smithsonian Insti- tution was in no way a party, involving infringement, or alleged infringement of the Wright patents by other manufacturers, and since, in 1921, the English patent attorney for the Wrights published a violent attack, with allegations of fraud, etc., in connection with the Hammondsport trials. 20. There are just three questions involved, which must be answered before it is possible to determine the capability of flight of the original Langley machine. These questions are: First, was the power plant adequate? Second, did the machine embody the proper aerodynamic principles to enable it to balance and maintain itself in the air? Third, was it sufficiently strong structurally to carry its weight and the stresses due to flying? 21. As regards the power plant, there seems no question that, in the Hammondsport trials the original Manly engine never developed the power of which it was demonstrated to be capable in 1903. Furthermore, during the Hammondsport trials with the original engine, the weight lifted into the air, including the pontoons, was 40 per cent greater than that of the machine as of 1903 with a pilot. Moreover, the bracing and supports to the pontoons and the pontoons themselves must have added materially to the resistance of the machine. If under these circumstances, the Langley machine was capable of arising from the water, which was demonstrated, there is no question in our mind that the 1903 machine had an adequate power plant. NO. 5 SMITHSONIAN INSTITUTION AND WRIGHT BROTHERS 23 22. With reference to the second question, although there were some changes in the supporting and guiding surfaces in the Ham- mondsport machine as compared with those of the 1903 machine, they were not, in our judgment, material, either as regards the Hammondsport machine when fitted with the original Manly engine, or subsequently when modified by a more powerful engine with a tractor screw. Moreover, the machine as it stood was vir- tually an exact copy of a quarter-size model which had shown itself aerodynamically quite satisfactory. We conclude, accordingly, that the answer to the second of the fundamental questions above 1s also in the affirmative. 23. When it comes to the question of strength, the case is not so clear. There is no question that the changes made in 1914 provided additional strength. Additional strength was obviously needed if 40 per cent additional weight was to be carried. However, the fact that additional strength was provided renders it impossible to remove the third question from the realm of controversy. This is a question for technical experts. A complete wing, one-quarter of the sustaining area, showed, by sand load test, ability to carry a total weight of 260 pounds without damage, while one-quarter of the weight of the original machine and pilot was 2074 pounds, only. Subsequently, the Hammondsport machine with a much more powerful engine (a Curtiss 80 horsepower engine) and with only a moderate increase in strength, showed itself capable of flight carrying 1,520 pounds, or 8s per cent more weight than the original machine of 1903. These facts, in our opinion, establish a strong presumption in favor of the adequacy of the structural strength of the original machine. However, we have asked the disinterested head of the Design Section of the Bureau of Aero- nautics of the Navy Department, to study with his experts the original machine and give us their opinion as to the adequacy of the original structure They are of the opinion that structurally the original Langley flying machine was capable of level and controlled flight. 24. It should not be thought that the original Langley machine was, in any sense, a finished product. Langley himself regarded his machine as only a beginning ; numerous problems had occurred to him which needed solution before aviation could be considered practicable. Since Langley and the Wright brothers looked at the subject from such different angles it would have been an inestt- mable advantage to the science and the art of aviation if Langley had been able to continue his work. 24 SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL. 81 25. In conclusion, we beg to call attention to the fact that a careful examination of the Langley machine now on exhibition in the National Museum shows that there are four minor inaccuracies as compared to the original machine of 1903, which should be remedied, namely: (a) The safety flotation tanks should be installed ; (b) The fin forward of the dihedral rudder should be removed ; (c) The vertical surface at the rear of the dihedral rudder should be removed ; and (d) The catapult lugs should be fitted to the king post. Respectfully submitted, JosEPH S. AMEs, Professor of Physics, Johns Hopkins University. D. W. Taytor, Rear Adnural-(C> C.)) Ut S; NacWetireds: In October, 1925, Dr. Walcott directed that the label of the large Langley machine of 1903 should be altered to read as follows: LANGLEY AERODROME THE ORIGINAL LANGLEY FLlyinc MACHINE OF 1903, RESTORED IN THE OPINION OF MANY COMPETENT TO JUDGE, THIS WAS THE FIRST HEAVIER-THAN-AIR CRAFT IN THE HISTORY OF THE WORLD CAPABLE OF SUSTAINED FREE FLIGHT UNDER ITS OWN POWER, CARRYING A MAN. THIS AIRCRAFT SLIGHTLY ANTEDATED THE MACHINE DESIGNED AND BUILT BY WILBUR AND ORVILLE WRIGHT, WHICH, ON DECEM- BER I7, 1903, WAS THE FIRST IN THE HISTORY OF THE WORLD TO ACCOMPLISH SUSTAINED FREE FLIGHT UNDER ITS OWN POWER, CARRYING A MAN. The aeronautical work of Samuel Pierpont Langley, third Sec- retary of the Smithsonian Institution, was begun in 1887. By fundamental scientific research he discovered facts, the publication of which largely laid the foundation for modern aviation. Langley designed large model aeroplanes which repeatedly flew in 1896 with automatic stability for long distances. The U. S. War De- partment, impressed by his success, authorized him to construct a man-carrying machine which was completed in the Smithsonian shops in the spring of 1903. Attempts made to launch it on October NO. 5 SMITHSONIAN INSTITUTION AND WRIGHT BROTHERS 25 7 and December 8, 1903, failed owing to imperfect operation of the catapult launching device. In these trials the wings and control surfaces were badly damaged and lack of funds prevented other tests at that time. The aeroplane was left by the War Department with the Smithsonian Institution for further experiments. In 1914 (following the foundation by the Institution of the Langley Aero- dynamical Laboratory) the experiments were resumed, using all available parts of the original machine. The frame and engine were the same as in the first trials; the reconstructed wings were used without the leading edge extension ; the control surfaces were reconstructed; and launching pontoons with necessary trussing were substituted for the original catapult. Thus equipped, and weighing over 40 per cent more than in 1903, with Glenn H. Curtiss as the pilot, it was successfully flown at Hammondsport, N. Y., June 2, 1914. With a more powerful engine and tractor propeller it was subsequently flown repeatedly. These tests indicated that the original machine would have flown in 1903 had it been success- fully launched. After the Hammondsport flights the machine was restored in accordance with the original drawings and data under the supervision of one of the original mechanics, using all original parts available. In 1918 the machine thus restored was deposited in the National Museum for permanent exhibition. (Its 52-horse- power gasoline engine was designed by Charles M. Manly, who superintended the construction of the machine and piloted it in 1903.) THE MODEL AERODROMES DESIGNED BY LANGLEY, THE LANGLEY- MANLY ENGINE, AND PHOTOGRAPHS OF THE MACHINES IN FLIGHT ARE SHOWN NEARBY. 6. As regards the sixth point as given on page 3 | do not know the basis for Mr. Wright’s feeling that the Smith- sonian has failed to recognize properly the abilities of him- self and his brother as research men. The Institution has published two articles, one by Wilbur Wright on ‘Some Aeronautical Experiments” and the other by Orville Wright on “ Stability of Aeroplanes ” (see Smithsonian Annual Reports, 1902, pp. 133-148, and 1914, pp. 209-216). Such publication by the Smithsonian Insti- 26 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 81 tution is in itself definite recognition of the status of the Wrights as discoverers of new truths. The Smithsonian Institution has borne charges in which have occurred the words “ hostile,’ ‘* insidious,” ‘ false propaganda,” in consequence of the events I have described. In order to show that the Institution’s officers have not been insincere I quote the following passage from a letter which I sent to the Editor of the Journal of the Royal Aero- nautical Society, April 27, 1928: 1. Langley himself said after the two unsuccessful launchings in 1903: “ Failure in the aerodrome itself or its engines there has been none; and it is believed that it is at the moment of success, and when the engineering problems have been solved, that a lack of means has prevented a continuance of the work.” He died in the same belief. 2. Manly twice risked his life in this faith, and eagerly wished to risk it thus again. From conversation I had with him in 1925, I am certain that he also died in the same belief. 3. Chanute on several occasions stated that “ he had no doubt ” that Langley’s machine “ would have flown if it had been well launched into the air.” Such, then, in brief review are statements that have been made. In concluding this account, I express, on behalf of the Smithsonian Institution, regret: 1. That any loose or inaccurate statements should have been promulgated by it which might be interpreted to Mr. Wright’s disadvantage. 2. That it should have contributed by the quotation on page 23 of the Smithsonian Annual Report of 1910 to the impression that the success of the Wright brothers was due to anything but their own research, genius, sacrifice, and perseverance. 3. That the experiments of 1914 should have been con- ducted and described in a way to give offense to Mr. Orville Wright and his friends. I renew to Mr. Wright on behalf of the Smithsonian In- stitution, my invitation of March 4, 1928, to deposit for NO. 5 SMITHSONIAN INSTITUTION AND WRIGHT BROTHERS 27 perpetual preservation in the United States National Mu- seum the Kitty Hawk plane with which he and his brother were the first in history to make successful sustained human flight in a power propelled heavier-than-air machine. [*1- nally, as a further gesture of good-will, I am willing to let Langley’s fame rest on its merits, and have directed that the labels on the Langley Aerodrome shall be so modified as to tell nothing but facts, without additions of opinion as to the accomplishments of Langley. This label now reads as follows: LANGLEY AERODROME THE ORIGINAL SAMUEL PIERPONT LANGLEY FLYING MACHINE OF 1903, RESTORED. DEPOSITED BY THE SMITHSONIAN INSTITUTION 301,613 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLUME 81, NUMBER 6 \ STUDY OF BODY RADIATION BY L. B. ALDRICH CITY OF WASHINGTON PUBLISHED BY THE SMITHSONIAN INSTITUTION DECEMBER 1, 1928 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLUME 81, NUMBER 6 A STUDY OF BODY RADIATION BY L. B. ALDRICH @O0eO8Se00, (PUBLICATION 2980) CITY OF WASHINGTON PUBLISHED BY THE SMITHSONIAN INSTITUTION DECEMBER 1, 1928 bse 2 yo ew. a- e @ eR. On eee 2g ue Qe 8 QF o & a 2 aya » ASTUDY OF BODY RADIATION By L. B. ALDRICH CONTENTS ohce TPE ee UIC GE TIG aes UE tg entero ese ic ts Pietenepecs union renene eeieaicke Pe Giese 2 WMeeAbboL Ss: Experiments) im. 1O2T os. 4-25. oes eos eee ce ei a on 3 Preliminary experiments in still air........---.-++seeee eee e ersten 7 Preliminary calorimeter experiments. ......---.++++se seer err tess r eset 14 Galorimeter tests’ with’ cloth’ wallso 7... 2. .22- 5-2-1 se einer ns 16 Relationship between melikeron and thermoelement results........-..++-: 17 Experiments on ten subjects in still and in moving air.......------+eeee+> 19 GTA IAHGE USS ITN Ney oe cle a tate et etee suri tc ere ceca inirns ro tassel 20 Suma yOr reStltse speek psn aeons rr mean eae 24 INGE) ELI GOS Ne aeeitio te pial elo mn ae ipinice ce icin icic rs ime Nea essa kona ec 206 LIST OF TABLES TABLE PAGE A. Abbot-Benedict observations in 192T.........--+--+ see sere eerie: 27 Br-1o. Observations and results of preliminary experiments on ten subjects. 29 C. Observations and results of preliminary calorimeter tests............-- 39 D. Observations and results of calorimeter tests with cloth walls....:..-.- 40 Er-10. Observations and results of experiments on ten subjects in still and in MOWATT AIG Sate es Se ie ie oc aoe eyes ate eee a eos 41 F. Summary of cloth-covered calorimeter tests. ...----.-.+++errsrtrtrees 51 G. Summary comparing thermoelement anid imelikenOtles sce eck lsiterda lei SI H. Summary of table E, varying air Talore alse ooweedoveunece coo oode Asc 52 J. Summary of table E, SEMA er Oe ke he tones Pree oeeeen mick: se pce iayea ope 53 K. Summary of changes in skin and clothing temperatures in moving air.. 54 L. Condensed summary of tables B and E.......-+----20ss eter r tte: 54 Lisr OF TEXT FIGURES FIGURE PAGE TeeNMeliicero tirana motintimer sts senic ale 4 cis le ery. = ete rials eel apes azole ahs 4 DubhermoclementGevice iss feais elstem «aise oo a omtenene siete folehle dient cesct enc ch 8 3. Thermoelement electrical COMMECLIONS asec cbse leek eee el teeta orient 8 4. Bath for constant temperature jtimetlOnale ee citar ecm aa ct fect 9 Cea@alibnabinee Dati mgt. coc ke op one ete aaa Oe aia | 10 6. Thermoelement calibration curve. .......--++ssesr ess ee settee tess II 7. Sketch showing body position MUTI DEES acter seo cree ne crneere ae sre ef sperr 12 @ Vertical and horizontal calorimeters. 2-22.02 *)sin? 6 where R= (constant Melik. No. 2) x C? = 4.0 X (current in amperes )” COON alOr T,=absolute temperature of radiator T)= absolute temperature of melikeron shutter Ae is i (13832) 1-0 == = SOL a = 0000 pe Pte? 83)? +G2) AOE” é ieee Ee Sone 16" From this equation, the value of 7,, the absolute temperature of the surface measured, is determined. In examining tables A and B, we find that the 4th power formula applied to the measurements on either skin or clothing yields values as great or slightly greater than the observed temperatures. This is evidence that the skin and clothing radiate as a black body at the low temperatures measured. Cobet and Bramigk (Ueber Messung der Warmestralung der menschlichen Haut und ihre klinische Bedeutung, Deutsches Archiv ftir klinische Medizin, Vol. 144, p. 45 to 60) con- firm this result on the skin, and Leonard Hill (The Science of Ven- tilation and Open Air Treatment, British Govt. Report, 1919, Medi- cal Research Commission) finds both skin and clothing nearly black body radiators for low temperature radiation. In table B, the values in the Radiation Summary were obtained by the application of Stefan’s formula to the mean temperatures given under Temp. Summary. For example, in table Br, we have given Estimated wall temp.=21°0 Mean skin temp. = 2307 Then R=a(T4=To*) = 8.20 x 10°" | (273+ 33.7) *— (273+ 21.0) *| = .1131 calories per sq. cm. per minute. 14 SMITHSONIAN MISCELLANEOUS COLLECTIONS voL. 81 The value of the Total Radiation of the subject is obtained some- what empirically as follows: The total surface area (see Notes on Tables, p. 26) is divided into sections skin area, girls 8%, boys 7% hair 5% clothing, girls 78%, boys 70% shoes 9% (if boots, 10% and clothing 78%) The average skin radiation per sq. cm., as just determined, is mul- tiplied by the corresponding number of centimeters of exposed skin, and similarly for clothing, hair, and shoe areas, and the sum taken. Since part of this total is ineffective, due to the area between the legs and under the arms not radiating to a full hemisphere of wall, this total radiation is reduced 8%. Dividing this result by the number of sq. m. surface area gives the value recorded under Total Radiation. PRELIMINARY CALORIMETER EXPERIMENTS The total radiation values of table B appeared much too large when compared with the basai metabolism values. The total energy pro- duction or metabolism must at all times equal the total energy loss. Exclusive of a small loss through urine and faeces and the warming of air and food taken in, there are three ways in which the body loses heat, namely, by radiation, by convection (including conduction), and by evaporation of water from lungs and skin. Du Bois states (Basal Metabolism in Health and Disease, ed. 1927, p. 400) that for a room temperature 22° to 25° C. and relative humidity 30 to 50%, the loss by vaporization of water from lungs and skin is about 24% of the total loss. By analogy with work done on the cooling of wires and blackened spheres we would expect the body convection loss to be at least as great as the radiation loss. For example, on p. 251, Smith- sonian Physical Tables, 7th Ed., McFarlane finds the total loss of least as great as the radiation loss. For example, on p. 251, Smith- to a blackened enclosure at 14° is .00266 gram calories per second, or .1596 calories per minute. On page 247 the difference in radiation between a black body at 24° C. and one at 14° is 918—801=117 gram cal. per sq. cm. per 24 hours or .0813 calories per sq. cm. per minute. Then the per cent of radiation loss of the blackened sphere (which, to be sure, at these low temperatures radiates decidedly less than the “black body ’’) is 0813 .1596 It is of course true that the actual energy production of each of the ten subjects was materially greater than that shown by the basal = 51%, convection loss = 49% No. 6 BODY RADIATION—ALDRICH 15 metabolism values. Yet even when adequate allowance is made for this, the radiation loss seemed to be an unexpectedly large proportion of the total energy production. After conference with Dr. Abbot and several members of the New York Commission on Ventilation, a series of experiments was started with the hope of shedding some light on the amount of body convection loss. These experiments proved that convection was, in- deed, less than had been anticipated, but the close approach of total radiation to basal metabolism remains surprising. HORIZONTAL VERTICAL _Fic. 8—Cylindrical copper calorimeters, each 38 cm. long and 30.5 cm. diameter, filled with water and completely covered with tight fitting jackets con- sisting of one thickness of brown canton-flannel cloth. A—Thermometer. B—Stirring device. Ss C—Electrical heating element. Two calorimeters were prepared of thin sheet copper, cylindrical in shape, each 38 cm. long by 30.5 cm. in diameter. One was mounted vertically and the other horizontally, each supported on four rubber blocks on the top of four metal rods. This permitted free convection and radiation on all sides but the rate of cooling of the vertical calorimeter might well be less than that of the horizontal because the warm convection currents rising from be- low would more closely bathe the sides in the vertical form. Appro- priate stirring and heating devices were inserted as shown in figure 8. Each was filled with a known amount of water, and the 2 16 SMITHSONIAN MISCELLANEOUS COLLECTIONS voL. 81 outside completely covered with a tight fitting jacket consisting of a single thickness of brown canton-flannel cloth. The purpose of the shape and covering of the calorimeters was to simulate the clothed human body. Heat was lost from the calorimeter only by radiation and convection, and the total loss of heat per hour could be accurately determined from the rate of change in temperature of the water and the water equivalent of the calorimeter. A series of tests was made with each calorimeter, and in each test the radiation loss was determined with the melikeron and with the thermoelement, following exactly the method described above as applied to human subjects. p 0 Fic. 9—Diagram for computing solid angle exposed to melikeron. The results of the preliminary tests are given in table C. Several interesting points appear. First, the radiation loss of the horizontal calorimeter is 6 or 7% less than in the vertical. This indicates that the shape of the calorimeter is important in determining the amount of the convection and helps to account for the difference in convection between the sphere 50% and the cylinders 70 to 80%. As noted above, however, this discrepancy is also in part due to the less perfect radiat- ing properties of lamp-black than of porous cloth. Second, in the test of March 3, with air motion of about 300 feet per minute the radia- tion is only 47% and the convection increased to 53%. Third, with no cloth cover, the test of March 26 shows only 34% radiated from the copper surface in still air. This is an indication of the low emis- sivity of the metal surface as compared with the cloth. CALORIMETER TESTS WITH CLOTH WALLS A weakness in the experiments thus far has been the impossibility of accurately knowing the mean temperature of the walls to which the subjects or calorimeters are radiating. A very helpful letter from Prof. Phelps of the New York Commission dated March 27, 1928, suggested the possibility of standardizing the wall conditions by sur- rounding the subject with cloth draperies whose temperature, closely that of the air in the room, could be determined with the same thermo- NO. 6 BODY RADIATION—ALDRICH Ay element used on the subject. This suggestion seemed especially feasi- ble since our results indicate that cloth radiates nearly as a black body. Accordingly, brown canton-flannel cloth was hung forming a cur- tained room 24 meters high and 14 by 2 meters in area, enclosing the calorimeter and with the melikeron mounting projecting through the curtain. The same cloth also formed the ceiling and floor. For part of the tests a current of air of known velocity from an electric fan outside the curtain was admitted through a hole in the cloth. The air velocity at the calorimeter was measured with a Katathermometer, an instrument invented by Prof. Leonard Hill, of England (see The Science of Ventilation and Open Air Treatment, British Govt. Report, 1919), and serving admirably for this purpose. The motor of the electric fan was run on storage batteries to insure a more constant air current. The Katathermometer was kindly furnished by Mr. Duffield, of the New York Commission. The results of these tests are found in table D. Table F is a con- densed summary of both tables C and D. From these tables a num- ber of conclusions can be drawn: (1) The amount radiated from the horizontal cylindrical calori- meter is about 7% less than from the vertical cylindrical calorimeter. (2) The estimated wall temperatures in the preliminary calori- meter experiments are too low. From this cause the amount radiated should probably be lowered at least 10%. Much more weight can be placed in the measured wall temperatures of the cloth walls. (3) For air motions greater than 75 feet per minute, the melikeron is unsatisfactory for use. An irregular drift of the galvanometer zero due probably to small fluctuating convection currents makes the in- strument unreliable. (4) In the preliminary experiments the melikeron gives appre- ciably higher results than the thermoelement, and in the second set of experiments this discrepancy disappears. The cause of this is explained in the following section. RELATIONSHIP BETWEEN MELIKERON AND THERMO- ELEMENT RESULTS In the preceding experiments, recorded in tables B, C, D and E, we have 265 comparisons of temperatures determined directly by thermoelement and computed from the radiation as measured with the melikeron, and including skin, clothing, hair, shoes, wall, and cloth-covered calorimeter temperatures. By a study of these com- parisons we can determine their relationship, with a view to using only the thermoelement in a new series of experiments. The thermo- 18 SMITHSONIAN MISCELLANEOUS COLLECTIONS voL. 81 element is much quicker and easier to use than the melikeron. Also it offers no difficulty in air currents where the melikeron becomes unusable. Table G is a summary of this kind. The first trials with the water jacketed melikeron indicated a minus correction (see page 8) when the water jacket temperature was less than room temperature. The water jacket temperatures are therefore given in all the tables and the differences Room Temperature minus Water Jacket Tempera- ture are recorded in table G. Examination discloses a rough equality between the differences Melikeron minus Thermoelement and Room Temperature minus Water Jacket Temperature. In the wall tem- peratures no difference is noted between melikeron and thermo- element—which is as we might expect since the wall is always close to room temperature, and the melikeron reading is very small. In the measurements at air velocities greater than 130 feet per minute there is decided disagreement —which we know is due to the unsatisfactory performance of the melikeron in air currents. Of the remaining ob- servations, the difference Melikeron minus Thermoelement on the skin is much larger than in any other group. For comparison, all the other groups, viz., clothing, hair, shoes, and calorimeter, are united in one group at the bottom. From the algebraic mean of Room T. minus Water J. T. in this group, when Room T. minus Water J. T.=°82, Melik. minus Therm.=°8o0 that is, the melikeron calculated temperature is in error by as much as the water jacket differs in temperature from the wall. Hence we may conclude that on the skin, when Room T.—Water J. T.=0, Melik.—Therm.=I?91 —0°67=1°24. Again from the arithmetical mean of Room T.—Water J. T., when Room T.—Water J. T.=1°14, Melik—Therm. = °80 from which on the skin, when Room T.—Water J. T.=0, E31 1.14 A mean of all clothing, hair, shoes, and calorimeter (Melik.—Therm. ) differences whose (Room T.—Water J. T.) differences are less than 10 gives Melik.—Therm. = 1°91 — x 2S0= 1.100 No. of Melik.—therm. Room T.—water J. T. observations difference difference (algebraic) 30 251 °39 Calorimeter tests alone give 8 -20 .24 = me Cp eh SMM © Steer ety er i ay NO. 6 BODY RADIATION——ALDRICH 19 These results confirm the preceding conclusion that the melikeron computed temperatures are in error by just as much as the water jacket differs in temperature from the wall. Summarizing the above evidence, it appears that when the Room T. minus the Water J. T. is zero, the Melikeron and Thermoelement temperatures on clothing, hair, shoes, and calorimeter agree with each other within °1. On the skin, however, when the Room T. minus the Water J. T. is zero the melikeron computed temperatures are approximately 1°1 greater than the thermoelement temperatures. Dr. Abbot’s skin measurements of 1921 at Boston (see table A) give evidence of the same thing—that on the skin the melikeron computed temperatures are higher than those measured directly with the thermoelement. A mean of 53 of his values in table A gives Melikeron minus Thermoelement =1°9 As an explanation for the persistently larger melikeron temperatures on the skin, Dr. Abbot suggests that since the skin is porous and the internal temperature of the body is higher than that of the surface, the melikeron sees into a deeper layer than that reached by the thermo- element. EXPERIMENTS ON TEN SUBJECTS IN STILL AND IN MOVING AIR A second series of experiments on human subjects was begun on May 30, 1928. It included 8 children of school age and 2 adults. Three similar sets of observations were taken on each subject, first in still air, second with moderate air motion, and third with faster air motion. As before, the air motion was produced by an electric fan three meters away, the motor of which ran on storage batteries. Air velocities were again measured with the Hill Katathermometer. Each subject was placed inside the same curtained room described under the calorimeter experiments. Skin, clothing and wall temperatures were measured with the thermoelement device. Exactly the same body and wall positions were measured on each subject. A complete set included 7 observations on the exposed skin, 15 on the clothing, I on the hair, 2 on the shoes, and ro on the walls. The skin tempera- tures were then corrected to the melikeron scale by increasing the thermoelement skin temperatures I°I as explained in the previous section. The observations were grouped and summarized as shown in table E. Following exactly the method described on page 13, Stefan’s 4th power formula was applied to the various groups and values of the total radiation determined as given in table FE. 20 SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL. 81 Unlike the first series of experiments on 10 subjects (see table B), this second series was carried out in midsummer. Fortunately the room was equipped with refrigerating pipes so that the room tem- peratures were kept fairly normal and comfortable. These pipes were entirely outside of the curtained room where experiments were made. Attention is called to the summaries of table E data contained in tables H, J and IX. Table H divides the air velocities into four groups and shows how, with the room and wall temperatures remaining nearly constant, the calories of radiation loss progressively decrease as the air motion increases. Table J gives for each subject, in calories per sq. m. of body surface, the basal metabolism and the loss of heat by radiation; also the ratios between these two quantities. These ratios are also arranged according to increasing room temperatures. A marked decrease in the ratio occurs with increasing room (and with it the wall) temperatures. Table K is a summary showing the changes in skin and clothing temperatures with varying air velocities, dividing the changes into three groups, namely, temperature changes on the side of the subject toward the fan, perpendicular to it, and away from it. A drop in temperature occurs in all three groups, with the greatest drop on the side toward the fan. In the other two groups the drop is only about one half as great. On the side towards the fan the clothing temperature drop is about one third greater than the skin temperature drop. GENERAL DISCUSSION We have presented the results of three series of experiments on the radiation loss of human subjects, and a fourth series on the radiation loss from specially prepared calorimeters. ‘The first series gave the results of Drs. Abbot and Benedict on the radiation and skin tempera- tures of a nude subject when the room temperature was held at 15° and again when it was held at 26°. It is interesting to note the change in radiation loss in these two different cases. On March 31, when the room temperature was 15°, the thermoelement skin temperature (mean of 87 values, many of which are not included in table A) was 27-2, and on April 1, when the room temperature was 26°0 (mean of 40 values), was 30°8. The black body temperatures, computed from the melikeron values and the Stefan formula, were March 31 (mean of 20 values) 28°2, and on April I (mean of 12 values) 33°4. Dr. Benedict estimated the wall temperature on April I to be 26°, and it is probable that on March 31 the wall temperature was at least as low as 15°. (Outside temperature was 8°6.) Assuming these wall No. 6 BODY RADIATION—ALDRICH 21 temperatures we can compute from the Stefan formula the average radiation per sq. cm. per minute from the body. It results as follows: March 31, from thermoelement, .1003 cal., from melikeron, .1109 cal. April 1, from thermoelement, .0432 cal., from melikeron, .0673 cal. Thus the body actually radiated in the order of twice as much when the walls were at 15° as when the walls were at 26°. The best work in basal metabolism indicates that an individual’s metabolism remains practically unchanged through this range of room temperature. A very considerable readjustment, perhaps of water vapor loss, must take place to compensate for the large change in radiation. Let us compare the two series of experiments on human subjects recorded in tables B and E. Each series included 10 individual sub- jects, composed of adults and children of school age, of both sexes and all normally clothed. The first series was performed in midwinter, the second series in midsummer. During the first, the mean relative humidity was 43% and during the second 62%. In each series de- terminations were made on each subject of the total loss of heat by radiation in still air. The second of the two series deserves greater weight for two reasons : (1) Cloth walis were used and the mean wall temperature deter- mined from actual measurements with the thermoelement. (2) A greater number of skin and clothing temperatures were measured since only the thermoelement was used. In the first series the subject radiated to the walls, windows, and furniture of the room. Their mean temperature was estimated from the room temperature, after a study, on a typical day, of the relation- ship between the room temperature and that of the walls, windows, and furniture. From this study it was concluded that the mean wall temperature was probably °5 below room temperature. This arbi- trary correction was adopted for all the preliminary 10 subjects and also for the preliminary calorimeter experiments (see table C). It is remarked on page 17 under (2) that the estimated wall temperatures in the table C data are probably too low. The reason for this can be seen from the fact that the table C data were obtained in the spring, whereas the table B data were taken in midwinter. The mean outside temperature was T1°o for table C, and 3°5 for table B. It is evident that with a warmer temperature outside, the °5 difference between room and wall temperature was too great. On the other hand, on examining the data of the typical day from which the arbi- trary °5 correction was determined, I find that the outside tempera- ture was 2° and that the mean wall temperature was in reality I?o below room temperature. The arbitrary correction was made °5 22 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 81 because | erroneously thought the mean outside temperature was considerably above 2° It is probable then that since the mean out- side temperature was actually only 1°25 above that of the measured day, the wall temperatures of table B should have been several tenths degree lower. As explained on pages 18 to 19, there is also another correction to be made in table B, due to the difference Room Temp.— Water Jacket Temp. This correction requires a lowering of the skin and clothing temperatures of about the same magnitude as the wall temperature correction just mentioned. It is a fortunate accident that the difference in temperature between the body surface and the walls thus remains nearly the same and the mean radiation values in table B remain unchanged. Table L compares the means of the two series, tables B and E. The total radiation is greater in the first series due to the lower mean room temperature. The adult basal metabolism (determined from Du Bois’ chart) is higher in the first series because the 3 adults were two male and one female, average age 31, whereas in the second series the adults were both female and average age 43. The work of many investigators agrees in placing the basal metabolism per sq. m. of body surface of adults considerably lower than that of chil- dren. Yet the radiation losses in tables B and E show no such change as between adults and children. In table L the ratios Radiation loss Basal metabolism are in each case higher for adults than children. This is difficult to explain. At normal indoor temperatures, in still air and with the subject normally clothed and at rest, the major heat losses would be dis- tributed as follows: The loss by evaporation of water from lungs and skin (as stated by Du Bois, see page 14) is 24% of the total. The convection loss, assuming it is similar to that of the cloth-covered vertical calorimeter, is } of the radiation loss. Or, Water vapor loss = 24% of the total Radiation loss = 46% of the total Convection loss = 30% of the total It is interesting to compare this with a statement by Rubner (see page 20, Leonard Hill, The Science of Ventilation and Open Air Treatment) that “for an average man, in still air, the loss of heat is distributed as follows: Warming of inspired air, 35; warming the food, 42; evaporation of water, 558; convection loss, 823 ; radia- tion, 1181 ; total loss, 2700 kg. calories.” NO. 6 BODY RADIATION——ALDRICH 23 In considering the method by which the total radiation values are obtained in these experiments, there will perhaps be question con- cerning the correctness of the empirical division of the body surface into skin, clothing, shoe, and hair areas, as well as the 8% reduction for ineffective radiation between legs and under arms. Yet these factors may be altered through a considerable range and not materi- ally alter the final result. The radiation loss will still be nearly the same magnitude. It has been a matter of surprise to the writer that the literature covering calorimetry experiments on the total energy consumption of human subjects makes so little mention of the surrounding tem- peratures to which the subject radiates ; also that in the comparisons between direct and indirect calorimetry the temperatures used are nearly the same throughout. It was my privilege on March 21, 1928, accompanied by Prof. Phelps, of the New York Commission, to visit the Bellevue Hospital laboratory of Dr. Du Bois, to talk with him and see the operation of the Sage calorimeter which, under the skilful manipulation of Dr. Du Bois and his assistants, has added a new chap- ter to our knowledge of metabolism in health and disease. The visit was of especial interest in that Dr. Stefansson, the explorer, was pres- ent for a metabolism test to determine the effect of an exclusively meat diet. Dr. Du Bois explained that the reason all his experiments had been carried out at nearly the same temperature was because of the in- tricacy of the apparatus and the difficulty of redetermining all the con- stants for each set of temperatures. He agreed that it was important to compare direct and indirect calorimetry at other temperatures and indicated that he hoped to find opportunity to do so. Incidentally in the course of these experiments, rough tests were made with the thermoelement device to see how rapidly its tempera- ture falls off as the thermoelement recedes from the skin or clothing. The thermoelement has a bright metal surface and consequently its temperature is but little affected by absorption of radiation. The tests show that, in moving the device horizontally away from the body, as soon as actual contact is broken between thermoelement and skin or clothing, the thermoelement temperature falls rapidly almost to room temperature and then gradually declines to room temperature as the thermoelement recedes. At 30 cm. distance no effect of the presence of the body could be detected in still air. There would be a marked effect of course if the thermoelement were held over the body instead of at the side, or if the thermoelement had a better emissivity so that its temperature would be raised by a larger ab- sorption of radiation. 24 SMITHSONIAN MISCELLANEOUS COLLECTIONS voL. 81 SUMMARY OF RESULTS. The concrete results of these experiments are briefly summarized: (1) The radiation from the skin and clothing is approximately that of a “ black body ” or perfect radiator. (2) Skin temperatures computed from melikeron radiation mea- surements are about 1° C. higher than skin temperatures measured directly with the thermoelement. This is not true on clothing or calorimeters. Apparently the melikeron sees deeper into the pores of the skin. (3) A cloth-covered, vertical, cylindrical calorimeter at body tem- perature loses in still air 60% by radiation, 40% by convection. A similar horizontal calorimeter loses 54% by radiation, 46% by con- vection. The human body convection loss is probably similar to this, that is, the convection loss is roughly one third less than the radia- tion loss, in still air and normal room temperatures. (4) Increasing air motion rapidly decreases the percentage radia- tion loss and increases the convectional. With the vertical calorimeter : Air motion % radiation loss 0 60 75 ft. per min. 4I 130 ft. per min. 35 190 ft. per min. 25 . (5) Total body radiation similarly decreases with air motion: Air motion Radiation loss (mean for ro subjects) o to 50 ft. per min. 30.7 large cal. per sq. m. per hour 50 to 100 20.3 100 to 150 257 180 to 250 23.2 (6) Increase in room temperature (which also means increase in wall temperature) produces a progressive lowering of radiation loss. The ratio Radiation loss Basal metabolism decreases with increase of room and wall temperature : Radiation loss Room temp. Basal metabolism Se eee eee 80 (mean of Io subjects ) 24.1 .75 (mean of 10 subjects) Z22yal: .84 (mean of 3 subjects) ‘ablewalevan oe 24.5 .74 (mean of 4 subjects) 25.6 .66 (mean of 3 subjects) (7) Keeping room and wall temperatures unchanged, the tem- perature of skin and clothing decreases with increasing air motion, NO. 6 BODY RADIATION—ALDRICH 25 the decrease being greatest on the side facing the wind and about one half as great on the side away from the wind. The clothing tem- perature drop on the side towards the wind is about one third greater than the corresponding skin temperature drop. Summary of fo subjects : Skin temp. drop— Clothing temp. drop— : [ 2 Sees vas Air motion Away from Towards Away from Towards Perpendicular (ft. per min.) wind wind wind wind to wind 0 to 100 —°4 — °8 —°6 —1°3 —°5 100 to 250 ow —— Te. aA wiley) — 5 (8) At normal indoor temperature, in still air and with the subject normally clothed and at rest, body heat losses are distributed as follows : Rivaporations ok water...caemeeteee dees 24% Radiant mimaege craved torsts sve cio teapot aie ov econe 46% Comnvectionmrer reife eee oe nae 30% (9g) Tests with the thermoelement show that the air temperature falls to room temperature very rapidly as the distance from the body increases. That is, there is a steep temperature gradient in the first centimeter or so from the body surface. With the thermoelement 30 cm. away no effect of the presence of the body could be detected. (10) The Abbot-Benedict work (table A) indicates that the radia- tion loss from a nude subject is about twice as great for a room temperature of 15° as it is for a room temperature of 26° This evi- dence does not entirely support the “suit of clothes ” theory referred to by Du Bois. In explanation of this theory, he says (p. 385, 1927 ed. “ Basal Metabolism”): “A constriction of the peripheral blood vessels (occurs) and the amount of heat carried to the surface is rela- tively small in proportion to the heat produced. . .. The patient really changes his integument into a suit of clothes and withdraws the zone where the blood is cooled from the skin to a level some distance below the surface.” (11) Normal fluctuations in humidity indoors produce negligible effect upon the radiation loss. This is to be expected. Our bodies, about 300° Absolute, radiate almost wholly between the wavelengths 4p and 50p with a maximum at Ion. Water vapor absorption is so strong for much of this range and so nearly negligible near the maxi- mum, Ion, that its possible effect is nearly fully produced even by the humidity of an ordinary room. Thus the effect of changes of quan- tity of water vapor in the ordinary room is small. Were the air of the room exceedingly dry, changes might be noticeable. 20 SMITHSONIAN MISCELLANEOUS COLLECTIONS. VOL. 81 Interesting and important questions concerning the comfort and welfare of children in classrooms are inadequately answered today. It is hoped that this report may in some degree help towards a better understanding of these problems. NOTES ON TABLES Temperatures are given in centigrade degrees. Air velocities are in feet per minute. Surface areas are determined from Du Bois’ height-weight chart (Archives of Internal Medicine, Vol. 17, p. 865, 1916). Basal metabolism values are taken from Du Bois’ “ Basal Metab- olism in Health and Disease,” edition 1927, p. 145. In table E, Wall A, B, C, D, E, refer to definite places on the canton-flannel curtains hung around the subject and forming the walls to which the subject is radiating. Places A, B, and D are on the sides, C on the ceiling and £ on the floor. Position numbers followed by an asterisk are taken on the skin because of short sleeves or low socks, In table E also, skin temperatures in the three columns on the right are just as read from the thermoelement device. In the sum- mary on the left they have been corrected to the melikeron scale by the addition of 1°1 as explained in the text. No. 6 BODY RADIATION—ALDRICH 2, TABLE A.—A bbot- Benedict Observations SuBjEcT: Miss W, nude March 30, 1921 Observed Temp. Temp. Temp. Computed Water (thermo- from Time Pos. Jacket element) Radiation Remarks 133 19 20 aT 32°8 34-1 Pos. 14 cm. below 19. 38 ar 20.8 33.0 2250 45 19a =: 19.8 31.6 31.8 Pos. 14 cm. below 1ga. 2 05 55 19.3 32.7 40.7 10 i 19.1 32.5 36.8 18 aie 19.1 31.9 39.6 29 54 19.1 33.0 34.2 Pos. 10 cm. to left of 54. 35 Lhe i eiahs B27) 34.2 45 55 19-5 31.2 36.6 58 54 19.7 BA Bere Pos. 10 cm. to left of 54. 3 27 32 20.0 2758 28.4 Standing facing window, holding iron 27 3I 19.7 2722 29.3 post to steady herself. 38 30 1Q).2 27a, 30.6 Pos. 2 cm. below 30. me 34 18.9 2053 Boho 9/ Pos. 6 cm. above 34. 53 45 18.7 2750 3082 4 OI 46 18.4 TET, 29.9 12 Am OrY7; 22a0 34.6 20 28 18.8 26.6 26.4 March 31, 1921. Room Temp. 15.0 C. Outside Temp. 8.6 C. ets iO 30.9 2242 Dr. B.’s hand. shies 20.8 52 29.9 Floor. Bear ors 19.4 oe 28.7 45 toward wall and ceiling. 10 23 14 Te 22.0 33-4 Miss W., subject. 34 46 Wed: 26117, 29.8 39 46 Wyjie2 26.0 2750). = 45 14 Wise 29.6 32.5 53 46 172 25.1 26.5 56 14 fn 29.5 Qn 7, II 02 46 Meee 25.4 26.8 10 ie 17-3 cae 21.5 Toward ceiling. 17 14 1793 29.2 31.4 23 46 17.3 24.0 25.2 30 32 17.4 22.6 23.9 34 30 74. 24.5 26.1 fe) 3 7A 24.2 26.1 42 28 L750 22.9 24.5 47 34 17.6 27 7al 29.1 57 53 78 29.0 31.8 28 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 81 TABLE A (continued) March 31, 1921 (continued) Observed Temp. Temp. Temp. Computed Water (thermo- from Time Pos. Jacket element) Radiation Remarks I2 02 54 17°9 26°5 28°4 10 55 L729 25.3 2651 21 55 18.2 25.6 26.8 Subject lying down, melikeron held 27 54 TSa3 25/07. 28.0 over her. 30 53 L323 28.9 29.8 2 00 ft 20.8 35.4 2073; Dr. M., subject, Rt. hand is made into a tube resting in left. Hands taken apart for skin temp., so that both palms were exposed. eee a 20.8 3505 3787 Palms not exposed, position held. April t, 1921. SuspyectT: Miss W. Room Temp. held at 26.0 Outside 4.2 9 56 14 25.3 3327 35-0 Sitting. 10 OL 46 24.6 29.8 2147 Standing. 10 14 2220 2083 2550 Sitting. 17 46 23.2 20).7 Bitn7, Standing. 24 34. 22.0 31.0 38-7 ; : 30 2 22.6 20.2 2129 Standing on stool. 36 29 22.4 30.0 2S) 40 32 2252 28.1 30.0 53 a3 21.9 32-5 34-7 II 05 54 21.6 2162 3530 12 54) 203s BTA | a0 21 »'53-5 20.0, “30-9 ) “345 ; Die a 24.5 34.4 BAe, Dr. M, subject, Rt. hand made into a tube resting in left. Melikeron opposite hole made by hand. Ther- moelement with rubber back in- serted in hole made by hand. ec my apis 36.1 See Opened hands and clapped them together again with thermoelement between. 2 OM oi 23.4 i. 338 Hands in position of tube. 03 eS) 22.9 eee sa58 Hands in position of tube. 06 7 ae 34.5 Looe No rubber back. Mean of 13 values. 12 ae 2126 iia 34.2 14 xe Beer 257 anes No rubber back. Mean of 10 values. 4I Be 2OnT 35.0 3587. Hands made into tube. 50 ae 20.0 35.2 35.6 NO. 6 BODY RADIATION—ALDRICH 29 Tas_e B.—Observations and Results of Preliminary Experiments on Ten Subjects TABLE BI DATE: Jan. 18, 1928. Temp. Sungect: S. A! Ss ae? See Sere Sex: Male. Temp. Thermo- _ Stefan AGE: 7 yrs. Element Formula WEIGHT: 25.5 kg. < 3 ss HEIGHT: 124 cm. [Saas aces LOPS see On 3450 SURFACE AREA: .95 sq. m. 26a......... 19.5 27:3, 29-9 CLOTHING: Green, wool suit, cotton | 100.-.--.--. 19565 | VgOn4e 3352 stockings. Said cn. : QMO) 12300. e2S.0 Air temperature outdoors, 10°. aa he ed 5 Blas ae Relative humidity indoors, 40%. 34 Sang Fate 25.0 eo Room temperature, 21°. 5. FORD 19.5 23.8 eye TORE a: FORA) 277 GOSS TEMPERATURE SUMMARY ints walleey ». TOe4e se Sia ©2050 No. af : ted Kind alien eee Oa ea oe oo formula San 3 3 A 7 ToTaL RADIATION Glothing.-.....: 5 28.5 36.9 large calories per sq. meter per ante Sac te I 29.9 hour. Shoessene ena (est. ) 25.6 Wall (est.) 21.0 BAsAL METABOLISM RADIATION SUMMARY Calories per sq. cm. per min. SS clnlges eee Ao eters ror ede ake 1131 Glothinee ee ee 0653 MUL gen perso he a neaiciens SS. 2 0779 43 large calories per sq. meter per hour. 30 SMITHSONIAN MISCELLANEOUS COLLECTIONS voL. 81 TABLE B2 DATE: Jan. 21, 1928. Temp. SuBJECT: M. W. Place Tee ee SE ae Female Temp. ee eoteep GE: II yrs. ement Formula WEIGHT: 28.1 kg. a c 2 HEIGHT: 140 ae int. wall..... 20.7 2159 Zita, SURFACE AREA: I.07 sq. m. 18a......... 20.7 32-3. 33-4 CLotuinc: Red, wool dress, cotton | 26a..-...... 20.7 30.1 28.4 stockings. GOOMare atres.c 20.7" “31.9. | 3250 Air temperature outdoors, —2°.8. Deena fs 2057 PAO he eeee ; Sf aE Le Oy 1 ee ee 2029 525-7). 32027, Relative humidity indoors 32%. Room temperatireoe ont int. wall..... 2047 2129 217, Pp ns Aber serge 20.7 26.0 26.3 OMe ence, @2On7, 27.7, 28.6 TEMPERATURE SUMMARY TESA te tee 2017 214 gee : No. Temp. computed | 26a......... 20e7, 27.0 26.4 Kind Values” romsttag int. wall..... 20.7 “21Gs E2I.7 6 LOO teas Sane 20.8 30.2 33.6 Skinp aac 4 33.0 Sab eke ee 2018.9) 2427 BZOar Clothing... . oe 5 26.7 PAirinss aoe ae 2 27.4 Shoesi. cee (est.) 24.0 Tora RADIATION Walle oe ccs, Aree (est. ) 21.40 h 25.8 large calories per sq. meter per our. RADIATION SUMMARY Calories per BASAL METABOLISM sq. cm, per min. 44 large calories per sq. meter per Shima cise ie tang 4 pete ace oe ,10%2 hour. Clothing 5's! iach eieteda eine .0440 PLAT yee ate cho cee a ee .0502 BODY RADIATION SUMMARY Calories per sq. cm. per min. S Rolle estas oho Sasi teee ne eras arr loti er eer be esate ee ae . 0642 A QIKe mre Seniesa A rani! 0878 Shoesmntas stench onl eaia. 0420 A 19.2 RADIATION—ALDRICH 31 TABLE B3 DATE: Jan. 28, 1928. Temp. SuBJ EcT: J .S. Place yackee ene epee Sex: Male Temp. Thermo- Stefan AGE: 12 yrs. Element Formula WEIGHT: 46.2 kg. . 2 2 HEIGHT: 158 cm. DO arnt ctss aia: 19.2 30.8 32.5 SURFACE AREA: I.44 sq. m. TOO......... 19.2 30-4 32-3 - CLOTHING: Cotton waist, corduroy | 262.--..-.-- 19.2 24.1 29.4 trousers, high, red rubber boots.| !!--------- 19.t 28.6 25.4 (Snow storm outside) S4---+----- I9.I 30.I 29.0 Air temperature outdoors —4°.4. a8 She eae eae aes apes Boe Relative humidity indoors 37%. sone ie ; ae pene OWS een ee eH, | kere) ro>raltal sien ’o vie) 6: . . . Pons tore ate EOi 7. TO Coal ee LOML 19.4 19.5 LOA tees TO52) V22540 + 23.7 TEMPERATURE SUMMARY Sie c 1O.2) 2377) 6 2458 ; No. Temp. computed | int. wall..... 19.2 19.3 19.6 ee ee epee Stet NAB arene of: TO.1 | 20.5) 637s LOOR ieee. LOST | \20n7) Vs24 SUS ystems 4 32).2 Sens Mn eg ea 3 oe ToTaL RADIATION Sloesi | ceo ee 24.2 36.7 large calories per sq. meter per RNallies neve to (est.) hour. BasAL METABOLISM 44 large calories per sq. meter per hour. SMITHSONIAN DatTE: Jan. 31, 1928. SUBJECT: S. W. SEx: Male. AGE: 6 yrs. WEIGHT: 18.2 kg. HEIGHT: ITI cm. MISCELLANEOUS COLLECTIONS TABLE B4 | \ Place black velvet. SURFACE AREA: .76 sq. m. 18a......... CLOTHING: Cotton waist, wool trousers, | 100--------- cotton stockings. 2 Oe erica a Air temperature outdoors 0°. ot uae tk: Relative humidity indoors 46%. Bel. Sue, | GIG kd Room temperature 21°.8. be, De eee 5 TEMPERATURE SUMMARY Omer No. Temp. computed TOG neces eis ace Kind Values from Stefan 7 SOTA LM oe Pees oot ms BOQ = Skint esis ore 3 34.0 mt. wall <.; Glothingee eee 6 Dae Hain, 2s aback I 22)0 Shoes nee sees I 25.6 Walleye eee est. Die < ee 3 hour. RADIATION SUMMARY Caiories per sq. cm. per min. S inh ys Meee pe ee eee Shisal hour. Clothing: puss cae ners .0562 Giese iets pene Roe .0950 SHOES son ayant ape Me .0372 VOL. 81 Temp. Water Temp. computed Jacket by from Temp. Thermo- Stefan Element Formula 19.8 21,0 20.5 19.9 32.3 3320 19.9 30.6 33.6 19.9 30.6 32.0 19.9 2752 28N 7, 19.9 2872 28.7 19.9 25.6 272 19.9 2123 216 19.9 25.8 25.02) 19:0) — 2557 #2628 20.0 29.2 29.8 20.0 23.0 25.6 20.0 32.0 34.8 19.9 2Me2 Zang ToTAL RADIATION 33-3 large calories per sq. meter per Basa METABOLISM 44 large calories per sq. meter per NO. 6 BODY RADIATION—ALDRICH 33 TABLE B5 DatTE: Feb. 4, 1928. Tek tye Temp. : . . compute SUBJECT: E. L. Place Jacket ie ior SEx: Female. Temp. Thermo- Stefan AGE: 8 yrs. Element Formula WEIGHT: 24 kg. ¥ g HEIGHT: 127 cm. EO ierias ress AOsG) Resi a il SURFACE AREA: .93 sq. m. 100......... 20.9 31-5 34-3 CLoTHinG: Cotton dress and stockings. | 262-.-.----. 20.9 27-9 28.7 Air temperature outdoors 13°.4. oe fe gt Zea ‘ 8 a8 oe ° Relative humidity indoors 46%. saree. 5 aa Be i om 3 On eeriber te me I a eer inn Stroh ro _ . . Room temperature 22°.4. SA Ae ira Ba, Mactan anes TOs Bae ee 20. 206% air TEMPERATURE SUMMARY Sie B68 27 38.8 No. Temp. computed | int. wall..... 20.9 22.4 21.7 Kind Maldess ¢ von Sitan Lopes Sh 2010) 322) 35R7 f a LOT eee to 20.9 | 8To5 3387 SIS ee ae 4 BAe, Eeouune Bers 8s 2 ae ToraL RADIATION Se a uc Ae 35.2 large calories per sq. meter per WVeciilesnes OM Es Beste) 21.9 hour. RADIATION SUMMARY BasaL METABOLISM Calories per . 42 large calories per sq. meter per sq. ° min. our. Shiner sere & otereisieiae . 1152 Glothingheans ya ets 0592 LAT is catt OR ys eters ae .0598 34 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 81 Temp. Water Temp. computed Jacket by from Temp. Thermo- Stefan Element Formula fo) °o ° 20.6 33.0 35.1 20.5 30.8 251 20.4 27-6 28.4 20.6 25.9 28.6 20.8 28.0 27.37 20.9 27.5 272 20.9 21 <2 20.2 20.9 25.3 26.5 20.9 25-7 25.8 20.9 28.0 29.5 20.9 23.4 25.7 20.9 30-5 33-3 20.8 210 21.5 BasAL METABOLISM 42 large calories per sq. meter TABLE B6 Date: Feb. 4, 1928. SuBJECT: R. S. SEx: Male. azk AGE: IO yrs. WEIGHT: 31.8 kg. HEIGHT: 138 cm. 18......... SURFACE AREA: I.10 sq. m. TOO! «opie sie CLotHinc: Cotton waist, grey, wool | 26a-......-. trousers, stockings. O40 ++ eee Air temperature outdoors 8°.3. PRE ats. Relative humidity indoors 46%. : 2 een... Room temperature 21°.4. 29 ee ee Pa eS TEMPERATURE SUMMARY RO ay eeeye er eee No. Temp. computed SMe asas tes & Kind Values from Stefan TOT eee oe ee ; formula int. wall..... S lela ase een et 3 34°4 Clothingay.. a 6 Dies Pla Seis sca ee I 28.4 Shoes i.6 ad eae I 25:7. Wall (est. ) 20.9 hour. RADIATION SUMMARY Calories per sq. cm. per min. S kite Nao Mecanens-2, n eane eee 1203 Wlothing. 225-2 ae eens eee 0566 lat ee) ee ee .0652 hoesiaess 2 uien «2 eer eee 0420 ToTaL RADIATION 33.1 large calories per sq. meter per hour per NO. 6 BODY RADIATION—ALDRICH 25 TABLE B7 Date: Feb. 11, 1928. Temp. “ Wat ali 3 ted Supject: P. L. Place Jacket by from SEx: Female. Temp. Thermo- Stefan AGE: 8 yrs. Element Formula WEIGHT: 25.9 kg. 2 5 e HEIGHT: 129 cm. LOTeerercs 05: 20,9) - 202 7)(©)) 3562 SURFACE AREA: .96 sq. m. 18......... 20.9- 32-8 34.5 CLorutnc: Cotton dress, short sleeves, | 262--------- 20.9 27-9 27-7 cotton stockings. S4-- 2s eee 20:9 | 25.6 2627 Air temperature outdoors 6°. oe eo: 20:0) Ae Ag cere, Relative humidity indoors 46%. af Bee Bee Bee = : OF ee ae 0 Maetete)ialtavel eVinie . . . Room temperature 21°.9. 20 rarities; ans 20.9 25 il 24.8 TOW cre gee: 20) 29.1 O. TEMPERATURE SUMMARY ia ee 566 ae I ee No. Temp. computed Kind Values from Stefan formula ToTaL RADIATION Sine hes 2 34.8 28.1 large calories per sq. meter per @lothing....6- 4: 6 26.8 hour. Ate hee Cee I 277. eons Rpeater parteletote ys ( z , Ze t BasAL METABOLISM alli Aroxetece rare ts est. 2a 42 large calories per sq. meter per hour. RADIATION SUMMARY Calories per sq. cm. per min. Sines eae oF sano veo . 1204 Clothingere ee ts sce ae ate .0465 BUI Hi aiesh AY. epee rercyey Spee .0555 30 SMITHSONIAN MISCELLANEOUS COLLECTIONS — VOL. 81 TABLE B8 DaTE: Dec. 13, 1927. Temp. Supjecr: M. Mh, ae a SEx: Female. Temp. Thermo- Stefan AGE: 27 yrs. Element Formula WEIGHT: 61.3 kg. i ; c HEIGHT: 165 cm. UO acter rers 19-9) Y 3207) wai SURFACE AREA: Te 67 sq. m. 26a eae 19.9 2023 26.6 CLOTHING: Dark silk: dress, silk stockq1|/"100-<--~. +. - 19.9 34-4 tees ings. SA tiauancr ere 19.9 28.8 216 Air temperature outdoors Dg See ae aoe ee Relative humidity indoors 59%. a Rupa ue eet on Room temperature 22°.6. SO ete eo ae i cn TEMPERATURE SUMMARY No. Temp. computed Kind Values from Stefan formula Skinh.c one seer 2 21.4. Clothing 22-2. 4 29.8 Hainan. . yc e I 26.6 Shoesic, ase ete a: (est.) 27.30 Wall (est. ) DOT RADIATION SUMMARY Calories per sq. cm. per min. SEINE cake hide Lee ee Oe: 0830 Clothing’ wee..42e2 o. see 0676 Paar wats aecacce aap cee .0390 Shoes 7 azote cesta eee ee 0426 TOTAL RADIATION 35-9 large calories per sq. meter per hour. BasAaL METABOLISM 37 large calories per sq. meter per hour. NO. 6 BODY RADIATION—ALDRICH 37 TABLE Bo DaTE: Dec. 9, 1927. Temp. : Wat Temp. ted SUBJECT: KerB: Place Take Son aretie SEx: Male. Temp. Thermo- Stefan AGE: 21 yrs. Element Formula WEIGHT: 61.3 kg. 7 . + HEIGHT: 173 cm. TOReeeriar sc 204) 20-5, |) Bisa SURFACE AREA: I.73 sq. m. 18......... 20.3 27-1 29: CLOTHING: Woolen shirt and trousers, T9Q......--- 20.4 25.2 27-5 thick leather boots. II........- 20.4 25 4 25-5 Air temperature outdoors, —2°.8. Saar oe ae 20. Relative humidity indoors 38%. ee accel cae Boe 23 - Room temperature 18.4. rae. Bare oo ee LOR ce teouceners 2250) DAs 26.1 TEMPERATURE SUMMARY (Seca. 22166 325) 23808 No. Temp. computed Kind Values from Stefan formula TOTAL RADIATION Sire ee 5 Paes . 38.2 large calories per sq. meter per Clothing Gea. .6. 4 26.0 one aire sees (est.) 270 Shoess.-ef ere I 23.6 BasAL METABOLISM Wall..........-. (est.) Tig 39.7 large calories per sq. meter per hour. RADIATION SUMMARY Calories per sq. cm. per min. SHeireeeeae 3c Ae ee eens ees BOVE, Glochin cae sere oie acter .0681 Rlettend ate RA ee ese eyk .0766 38 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 81 TABLE Brio DatTE: Dec. 21, 1927. ee Temp. e emp. compute SUBJECT: L. A. Place Jacket by from SEx: Male. Temp. Thermo- Stefan AGE: 45 yrs. Element Formula WEIGHT: 74.6 kg. . . e HEIGHT: 179 cm. MSA y eee tes 19.37 3329 23G.0 SURFACE AREA: I.93 sq. m. 26a......... 19.3 31.7 29.4 CLOTHING: Dark wool suit. Ila......... 19.3 25.0 27.2 Air temperature outdoors .. Relative humidity indoors 37%. Room temperature 21°.3. TEMPERATURE SUMMARY No. Temp. computed Kind Values from Stefan formula ° Skins. Haaaeen ie I 31.6 Clothing. a-.-e" I DTD Haine na eee I 29.4 Shoésic 5.000 oe (est.) 24.5 Walt o8 eech eae (est. ) 20.8 RADIATION SUMMARY Calories per sq. cm. per min. Sling antares a6 Ce eee .0950 Clothing ani. ese .0552 lair, SE a Sch uee thereat ete .0752 SHOeSH imide cee SS es .0317 ToTAL RADIATION 31.4 large calories per sq. meter per hour. BASAL METABOLISM 39.7 large calories per sq. meter per hour. NO. 6 BODY RADIATION—ALDRICH 39 TABLE C.—Preliminary Tests of Cylindrical Copper Calorimeters, Cloth-covered Vertical Horizontal PAINOUMEOLCOP Pela em eee joc 6s sl ee 2.91 kg. 2.60 kg. PAUMOUNtHOn FASS ee seers tere oe 2 wa ss eae 1.50 .50 PAIMO LI te OleWallenewn iene criertte ke oicucce = 255 jee 26.70 2720) Motalawaterrequivalent™a 64-4 sn. o..0 5. 27.05 kg. 27.48 kg. reavotscalonimeteneerieeninee eee os kcne 5LO3.. sq..cm. 5103. Sq-/em. Air Mean Melik. Loss of Heat Velocity Temp. Water in calories Date Calor- Room Outside (feet Gal Jacket per hour 1928 imeter Temp. Temp. per min.) Water Temp. (large cal.) Feb. 29 «Vert. BO D7, Oo 204 2028 21°36 Mar. I Vert. 2ayniT 8. oO BO M22 e2 19.57 Mar. 3. Vert. 22.9 6.7 about B2nOmm2On4 38.42 300 Marrs) Elonz, 2455) 14).4 O Bo OMAN 19.63 Mar t6) ) -Eloriz-) 2250 23 oO BoA 27.75 Mize 20 Ielosh, ABE Tee O BAS B26) 12.86 (no cloth Apr. 21. Vert. 24.1 8.3 O Be Mn Ree, cover) Apr. 25 Vert. 2OP Om Ia nO oO Bb LOLO Apr. 25 Vert. 22ST SO 0 eee a6 Mean Mean Temp. Loss by Temp. Cal. Radia- Esti- Cal. Surface tion (in mated Surface (computed large % Wall No. (Thermo- No. from cal. per Rad- spate Temp. Values element) Values Melik.) hour) iated Feb. 29 21.4 II 27.0 5 28.2 17.07 80. Mar. 1 23e2 12 29.0 4 29.1 15.83 81. Mar. 3 22.4 17; 25.6 6 29.2 18.25 Ave Mar. I5 24.1 18 29.0 6 29.3 14.00 lee Mar. 16 215 18 28.4 6 29.5 21.45 Tae Mar. 26 25.1 18 BON, 4 26.7 4.41 Bar Apr. 21 aes 28 B2e2 4 27a) Lua Bony: Apr. 25 AE 27 28.1 4 BO) 1 sk as Apr. 25 a OF 28.6 4 2O MAS Mey cvosene 40 SMITHSONIAN MISCELLANEOUS COLLECTIONS voL. 81 TaB_Le D.—Tests of Vertical Calorimeter, Cloth-covered, Surrounded by Cloth Walls Air Vel. Mean Melik. Loss of No. of Out- (feet Temp. Water Heat in Mean places Date Room side per Cal. Jacket large cal. Wall wall temp. 1928 Temp. Temp. min.) Water Temp. per hour Temp. measured Apr. 27. 21°6 9°4 Oo 3422 21.3 31.80 2250 19 Apr. 27 21.4 9.4 oO 2220 2a: 24.70 22.2 16 Apr 28° 224 8.9 oO 30.9 22.5 21.95 22.9 18 May YF 23-9 20:6 O 34.5 23.1 25.40 24.7 13 May I 23.9 20.6 130 3359 2202 33.50 24.6 II May 3 24.7 = 2722 oO 212 24.2 15.70 26.0 10 May 3 25.0 27.2 75 30.9 2AC2 16.75 25.0 9 May 5 28.5 33-9 19 39-5 28.5 34-95 29.9 12 May 5 28.8 33.9 130 38.8 28.5 25.15 29.7 1 May 7 24.4 20.0 190 36.4 25.6 47.60 25.0 12 May 7 24.4 20.0 190 35.6 25.4 43.80 25.0 12 Mean Temp. Mean Cal. Loss by Radiation Temp. Surface in large cal. per Cal. No.of (com- No. of hour % Radiated Surface places puted places —§_———-+————— —— Date (Thermo- meas- from meas- Thermo- Thermo- 1928 element) ured Melik.) ured Melik. element Melik. element ° fo} Apr. 27 29.5 26 29.1 4 18.95 20.06 59.5 63.2 Apr 27 2801 24 28.1 4 15.70 15.70 (6305 63.5 Apr. 28 27.9 22 28.2 4 14.13 13.40 64.4 61.2 May I 30.5 16 30.6 4 16-07 15:64 163.3 61.6 May I 29.2 22 30.6 4 16.43 12.54 49.0 B7aa May 3. 28.9 8 29.3 4 8.88 7193" 5605 50.5 May -3 928.5 9 2342 4 Gay 6.94 36.8 41.5 May 5 34.7 16 34.7 4 13.84 13.84 40.7 40.7 May 95) 9033.7 16 33.8 4 TT 55 eli sOmn 2256 3252 May. 7 20.1 16 21.4 3 Le 7Oe | LlsO4 angie 2212 May 7 29.2 16 BTa7, 3 18.50° 1.42: (42:2 26a1 “NO. 6 BODY RADIATION—ALDRICH 41 Taste E.—Observations and Results of Experiments on Ten Subjects in Still and in Moving Air TABLE EI DaTE: May 30, 1928. Place SUBJECT: S. A. a SEx: Male. LOOM eee 19 AGE: 7 yrs. 5 mos. walhAn es. 19 WEIGHT: 24 kg.’ Beis: 20 HEIGHT: 127 cm. Cae 21 SURFACE AREA: .93 sq. m. Dee 20 CLOTHING: Woolen sweater, cotton es 19 trousers, socks. 2 Goer «- 34 Air temperature outdoors, 21°.1. a ; 2 Relative humidity indoors, 56%. eae Be TEMPERATURE SUMMARY ca nek a Kind No. Temp. at air vel. Ser hs eh 209 Values $ 0 ree Re ay 38 Skanes. Te eS A 32235 Seb) | 2327: G 29 Glochinga se else 2564) 12765, 2623 | . 22-2 = 27, ann ees ee Geary. 3284 .| Ure ces 23 DOES caer 4: Pee eos G27 eA 2522)! Talis. 1. 32 Wall. LOM Ey 2250) 273 )|\ Lfisene ote e: 33 ROOMY S:; 5.4). 2 TOS 2h 3 2029) | 17a 34 OSs aie 26 RADIATION SUMMARY oe ote 2 (CaloriesiperSd=xcll.mni)|| | seaiaen Gln 2 per min. at air vel. 25 130 180 BOG eran 33 Snipers -II57 .0915 .0865| 35--------- 29 Glothimg: 22.5 - 0641 .0457 .0433| 36.-.-.---. 28 aires he estes, II3I .1094 .0990] 29...-...--.- 30 SHOES (0 4s 512% 0444 .0468 .0337] 30--------- 30 Bn een a: 27 ToTaL RADIATION a Ne Bf Air vel Calories per hour Bow 22 per sq. meter Cheek 23 g 38.6 Pe ee. 21 $30 eae Heer ee 19 180 Zig TOON sash 19 O ON DH HORA AH DAOCOO HAH AN Din OW DH DAL OI DON ° 21. 2. 21 23. 21 20. 35: 30. 29. 26 24. 25. 28 Die 28. 27. 30. Bon Bie BOR 27e 26. Qin Die 34- 2p 27 30. Parke 27 2a ate 22 2) - 21 21 MH DO: OM OWN OWWNNRHWOMNY ONOMUMH SUIWMWNODADARORH SO Temperatures at Air Velocity 0 13 180 ° 2Or Zi ile 21. Zien 19. Bee >) ° to . . . . . . . . . . . . © ODP HOH ONWUNHUO ONNUO OO COM DADHUH AP HON OW) N ° 42 SMITHSONIAN MISCELLANEOUS COLLECTIONS voL. 81 TABLE E2 DaTE: July 12, 1928. Place Temperatures at Air Velocity SuBJEcT: S. W. o 15 30 Sex: Male. TOO: |) 249 232 23510 AGE: 6 yrs. 6 mos. cy nA ee, eee 23.9) 523.57) WEIGHT: 18 kg. Pena ate oe nae oS 2372 HEIGHT: 116 cm. Go eta: 26.0 (24031) 2357 SuRFACE AREA: .78 sq. m. Deere 25.6. | 24,00 2304 CLOTHING: Tan cotton suit, socks. es 52 25,11 220 2375 Air temperature outdoors, 27°.0. 2 OF ea 35.6 35-5 35.0 Relative humidity indoors, 68%. TOO) res e152 32.0 31-7 31-9 Sa Ee Ol eyes 58: 23°01 "23. 3ap oac4 TEMPERATURE SUMMARY oe Winks: er aoe oo Kind No. Temp. at air vel. 54 i 30.2 31.7 22.4 Values 0 15 S ORMn pete aad ne : 5 a 3 BePNRi ccc we 2 29.5 30.7 28.8 SkitiaGntswe il 4-4 23seS baaea Basie > sek 32.0 30.0 28.7 Clothing :.4.. FX 3958, 3Y.A. 30.25 25-7 ce > 32.5% “3hsS sy, le Hairs oc24 8. 1 A561 633.6 22 e722 nee ck: = 7 32:6 = 325s WS le a7, Shoes! s.... 2 | (2884) 228-2" 2 2Sici elo ert: iri 34.8 34.4 33.8 Wall sac. oss TO 26.0. 245° “Se soup tOe. ox scis-i 33°9° 33-00) 33-3 Room. ..... 2. “95.1. 2303 Satna el seme - 35-1 33-3 34.6 M7 Aes yt aa esis B43 3308 a Rapration SumMARy rofl) jag 31g 31.0 Calories per eaacnt Stpeerecs 28.7 20:0 2822 fr pune ie 50 a Bee dh eas 28.2 25) — ee Sldnivsustahcee ;0766. -.0838: “= et | Soa, Hair alas. 083s “0838 one Dou? a Dott 132 OEE Rae Glothing.. 2... .0521 .0619 .0552 36 arae ed ee =0 ie i oe 5 Shoessa tne, 28 .0215 .0327 .0362 ai wis Se See a8 7) fay Eee 3146) 930.9 31,0 TOTAL RADIATION 2 Oe an ee Bil43 32 30.5 Air Vel. Calories per hour | wall A...... 26.6 25.4 24.7 per sq. meter Breese oe 2086) S25 ar 24.1 oO 2023 Crees 2087 25.7 24.9 15 34.4 Dees 26.2 24.3 23.9 50 22001 eee 25.8 24.3 24.1 ROOM. pa 25.30 23.13) eae 16 BODY RADIATION—ALDRICH 43 TABLE E3 DaTE: July 12, 1928. Place Temperatures at Air Velocity Supyect: M. W. g 1 a Sex: Female. roomie mae! 25300" 238 2352 AGE: II yrs. 5 mos. wallA.o...: ee a8, ahiG WEIGEr be 7 5 eeu Be 6) Caz aaa HEIGHT: 143 cm. Canes 26.1 “2458 24.6 SURFACE AREA: 1.07 sq. m. D B56. age Om 2806 CLoTHING: Light oon ‘dress, socks. Een A 5. SUES i 6 23.0 Air temperature outdoors, 27.0. 26 fd lS OW BOAOwe esas Relative humidity indoors, 68%. TOOM ete aa07 iaanse 6320s LOTaetea a eis: 2250 2205 32.0 TEMPERATURE SUMMARY 198 eee ih 32.6 33.9 33.0 Kind No. Temp. at air vel. LO rece ae 33-1 32.6 31-9 Values 0 15 50 Arey ate O49) 33-4 33-4 ° ° ° DIR eae ete 31.6 28.4 28.2 SO ee a 7. 35-0 35-8 34.0 Be er cio $9.3 28.0), 25a8 ee Teja Gal Ole Sti WSO). 21) Dane en hes. BlO) 304! VESIFO Hair.. Mee S oO Go a4 5h. 0 | 2m ayia a B10, Bile. 4 at Shoese.. 4. - BREST ARE S2O- On 277 Orc. Hei eee), 33.6 33.6 32.7 Wall... 2. -:. DOME Syme 24a i224 TOs doe 24.2) (3381) eae0e IRooment er OeeeIeAN 5 t2302) (2302 qe eee Baa. asso 285 Tareas. B42) | B52 B20 RADIATION SUMMARY ON ain SUNS C FET SNL) Gaioticn pee eq. ca! LOsmoaemenss Br294 3272) aio per min. at air vel. OREM aca ia 30.0 Zhe 6 . 0 15 BOR A 7s Raat eas Bil sO 29.2 27.6 S)ai ane FOSS HOOOS 0882 ||| 56am. 33.8 23.4 @r.8 Clothing SransnHen eye LO525 0 0587 f 0528 Bek Acie - 33.4 Boe 30.5 Pellettityeeches coeeoxene BO7GO ee OSOSm 64007 7) = 360 ier. 2000) 3380) 3i5 SROCSa ace si- cuter -04908:) -0447 0296 || “2910108: 29.8 20.1 27.6 TTT Tic COL ate ZOO) E2042 2509 TotaL RApIATION a See aa ae a sae E : bell 2 Seek apse) : : ; eh cae eats wall Ae a. 26530) e252) 2406 Oo 31.0 Beene D7) DEO) hak 15 34.2 Chen 26.1 252° 25010 50 30.2 Dae 25.3 BAN 24.1 | Dusen erate 25.2 2A 23.9 FOOMPA y- BAO? JE 2BF Or 23h2 44 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 81 TABLE E4 DaTE: July 14, 1928. Place Temperatures at Air Velocity SuBJEGI: alae. 0 84 136 Sex: Male. - % AGE: 14 yrs. 3 mos. ae Sera ess pe ae 22.8 WEIGHT: 43.5 kg. pee es pee a a3 Hercana teach Bon) 2a git 22) SuRFACE AREA: 1.36 sq. eG oe 3-3 253 CLOTHING: Cotton waist, ihlack woolen ies Se St) WEEE Space trousers, canvas shoes, bgt socks. Bee Bou oe Bee Air temperature outdoors, 27°.8. TOG Ue tet ae! 31 oS 30.7 8 Relative humidity indoors, 68%. OT eae 31.7 31.1 30.0 Qa eyes 2 31.2)'30..50 = 35.0 TEMPERATURE SUMMARY NO ge eigca® 31.1 30.6 30.5 Kind No. Temp. at air vel. Saisie 9 oes oe 29.2 30.1 29.4 Values 0 84 136 Die eaten 28.7 28.2 2827 . ° ° ° Beals attelhes 27.77 26.9 28.6 SIM eee oe 7 2 33423-9334) SZ ea Gays ss 31.1% 30,8) 32066 Clothing:. ... “15,' 73053: 929:3. 202A oe a. 2.6 940nl Siler Fait is. 52 sme Tet Sil th, 1.31. COsOM mT Sty shes. t 23.52). 1) 4341 S2iet HOES! sea e ess 2) a 282A)020,1, 270ml Salen fe ae 33.3. 32"5) 13078 Wally. feee- 10-2328) 2325. “23 2ueay fc... 34.7 33-7 32.0 Room...... 2 A 122.5 2227. 22d erase ates 34.6 32.7 30.1 Oise th ees 6 30.8 30.5 30.5 RADIATION SUMMARY OT ees 31-7 31-0. 30.7 Caloric ere cue Shree eee 28.2 29.1 28.2 per min. at air vel. FEES hice 28.6 29.1 277, 0 84 TS OME 200% ea” 2 ois BT 31.4 290 Skin's ea cue orate 0043) | 20802 FO700N/I 35am. ae. = 20.3. 128.1 28.1 Clothing....c..0.< .O569 “.O500" .O5274)) s0.s ene se 212 29.6 28.1 Flaite seme sn tte {0047 120708 "O55Sr0te9rce saa. < 2807. (27 Sere SHOES wee hat 0405“ sO401 9 sO4TAGIE BOlrr a. at. - 20.0) 12755 0 nee 7s 2 ee ee Zs eee BO G3 9 e2O nA a Deena es 20.5 | 30ar 8 12oe7 ee Toray ay nee wallvAte a. 2ATO. W232 540) 2367; per eq eee : pa eS a poe oa “3 - a ete a ae ay 84 fo ee oo : : : ee 2 eee 24/0 24:3 2406 3 Ooi TOOM..4.... 2216. 82.19) 92208 NO. .6 BODY RADIATION—ALDRICH 45 TABLE E5 DATE: July 18, 1928 Place Temperatures at Air Velocity SUBJECTS bale 0 97 235 SEx: Female _ s. i : LOOM Eee 242 2A e 24.5 AGE: 8 yrs. 5 mos. walltA vas: a>. 25,0. 24:6 2407, RICE eo ee ae 24.9 24.5 25.0 HEIGHT: I3I cm. Cu 250) -a5i5e eae SURFACE AREA: .98 sq. m. : dD ae 9 5 a Be CLoruinG: Light, cotton dress, high Eee oa aa see noe ° ° BOW eas cin bcs 34.5 34.6 34.4 ir temperature outdoors, 28° to 33°. Tae E : se Relative humidity indoors, 61%. On eran erie Se a : ; TQameieemicn: 32.0), 29.9 220.5 TEMPERATURE SUMMARY TOmser nee 25 eS liege Slee Kind No. Temp. at air vel. SA eraey: anaes of: 30.8 30.6 Zi? Values 0 97 235 Dei ers ee: 28:9) 3052)" 930-5 : . ‘s Sell |.) Sauer ees fs ‘ : Shin... 7 Ge 34-4533.6) Oe tee cus Clothing.... 15 20n6n 3043) 2989 Ai 32.0 30.9 30.1 Telaire tener I d2-5mme2- 2) (30-4) gu ao) age geet Shoess-ne ae - 2 20n45 62052) 28e1 onal cee B26) 93318 aai6 Well 66.6 oe TOMER 25m Te P25 e 5 ene Bee 33) Sue ans IRoom—er eer 2 Aleit Phe Bats 17a ee aang eaanole Baek Oona ee 29.8 30.5 30.0 RADIATION SUMMARY MOG cata oa: 3I-I 30-5 31.2 lentgcieaeer caters Sirians hee AQ) A) Ahh per min. at air vel. cote Baad: 29.4 29.2 28.0 97 2355) “DGarieneue se Z205 0 S252 ee Sie Skane eae BOSTOMMOS5 ONE O7 340) | 35oqee eee 21 OG OnOMmEZORS Clothing 44.25% EO5026 0460355 20350) || 3Oneee nee BPG Oth Bit ats) citer tee eccsi SONS Ole BORRYO|| Osoucansec 4OO 29 AGG Shoeste eae ent. 587A OZ05 O23" || 3Oesee ene 20040 2792 On9 ee 2 30.0 2080) 29,2 2Sir Nore che sy 30.1 29.8 29.8 eee Torta. eek ee wealllpAVerwe = 2650) 25860" 25,4 : per Eaeactes 5 eer Se Bee ae 9 29-0 Dee. ZERT) 2560) 2503 97 2s Eee PNAS) Dl) IS). 235 oad, MOON. 6 o06es 24.0 24.5 24.8 40 SMITHSONIAN TABLE E6 DaTE: July 18, 1928. Place SuBjEcT: M. A. 0 SEx: Female. i AGE: 55 yrs. eich Fat a a aH WEIGHT: 64.5 kg. ue ae HEIGHT: 167 cm. Cotas. = SURFACE AREA: 1.73 sq. m. nae 5 CLOTHING: Light, cotton dress, black ie a stockings. BGR ts hat 4 Air temperature outdoors, 28° to Ba Pag ee Relative humidity indoors, 61%. Botte hay 32 TEMPERATURE SUMMARY I9a......... 30 Kind No. Temp. at air vel. 19.......-. 30 Values 0 97 235 By Uaorecre ate 31 S o ° a Neg et he 2 Skin........ 72) 39-0) (3400) 34 eA a ae. ec Clothing. «2915. | Syl 30:7 (3O25iaay is 31 Flaite 18 oleae, 28h e26.9° 27 15a aa fy. 30 Shoes (608) 22 52022. +2053: 28 3a ars Ste 34 Wall 5. c2ceus 10° 2525 25.3. 25igul pea... ks 33 Room." .)..t (2 924.0" (242° SAGE 35 RADIATION SUMMARY DES eae > Calories per sq. cm. 3 net el 23 perimin: atair.vel:> Miwa ee 3 0 97 Scere 29 Skin. cadsassts SOSSSeu0853. 5.082 Lo Bie eal ola are 29 Clothing....... .0504. .0490 .0464 | 26a.......-. 28 VAI, «wi Sa 2s teyeke .0262 .0126 .0194] 35---:-:--- 32 Shoes......... .0332 .0356 .0268] 36.---..--. 3! Oe oie 3 ToTaAL RADIATION Oia ree 3I Air vel. Calories per hour Dive ey aeons 31 per sq. meter 20eere ee 20 oO 28.0 wal lA erenaer 28 97 26.8 G Se 26 2 HoNeagamsy ate 26 aor Di eee 25 Ee oe 24 LOO Nee: 24. OM HN ON NH DWONTOR HN ERM DANOH OHOUDARH DWHUHOWO MISCELLANEOUS COLLECTIONS 97 BOW 1 WNIENW AHORUNUUH DAN ONO HOKNOONOHANOAN VOL. 81 Temperatures at Air Velocity 235 ° 24. 2A 25. 26. 24. 24. 33- 30% MAN CO RRUWODANUNNMNUUM HHH HAO ANH AONMOMO 5 7 suc) BODY RADIATION—ALDRICH TABLE E7 Date: July 19, 1928. Place au Suspect: M. B. x Sex: Female. fOOM: $45. « 24.5 AGE: 4 yrs. 8 mos. wallsAe. @.).: Zoe WEIGHT: 15 kg. Beaver! 25.1 HEIGHT: 100 cm. Cites BGT SuRFACE AREA: .65 sq. m. Reis 25.1 CLotuinc: Light, cotton dress, no 130 ee 24.8 sleeves, high socks. sGEe fa: 32.2 Air temperature outdoors, 29° to Bane MO Olag nia han ce 3250 Relative humidity indoors, 58%. TORR eect: Bod. VQaeae oie: 32.0 TEMPERATURE SUMMARY a NS aes See Kind No. Temp. at air vel. Peis th IN ; , 6 Values 0 140 235 Cie ea ae ee oe k ° ° ° eR ipehet tense ens 30. 8 Shanice. + ees le Ome On RADIATION SUMMARY a wees os ; oa 8 oa a een eee iNew a cee 2 3050" 2s SP EETGONS 0 140 235mm 20d. cceuiarr: = 31.9 Bilbo 30.4 Skinsaiews 24 se sOS2100 0738 y -OOSON Eee mayen fcc 32.6 31.5 31.0 Clothing....... .0502 .0437 .0380| 36--------- 32-7 31-8 31-5 Haire: cms. 1.05340 20403. SO39amlnco seam ace as 30.1 26.5 26.9 Shocsaistoade JORIS" 420341) OZOGN RO” eae 29.8 27.8 27.4 2) Bt re aie 2163, 3065, 2059 BB epee kya au8 122 r.2 Oo. ToTaL RADIATION wallA...... S66 Bee ge Air vel. Calories per hour Bet ace ee2On5 26.6 267.3 Deed Eucrcy Coe 271 “27 J eee Oo 28.9 Dy fee 25.8 26.0 26.1 140 25.2 Bee teens 25 A 25 Aye 2 5c 235 22.6 TOOM nee ae 24-0 5530 ean NO. 6 BODY RADIATION—ALDRICH 49 TABLE Eg DATE: July 20, 1928. Place Temperatures at Air Velocity SuBjEcT: Es L. g i. 2 SEx: Female. 4 ROM 6 og aae Ao 25.9 26.2 AGE: 8 yrs. 5 mos. wall A...... 26.6 26.4 26.6 WEIGHT: 24 kg. 27.1 26051 26 OT era yietse) ere. e . . -9 HEIGHT: 129 cm. Cm, 2EsAn e27EAy aaa SURFACE AREA: .94 sq. m. D Ba) ae ee ee sts, Fel | aD : -4 26.8 CLOTHING: Light, cotton dress, no E 265) meaGisn CONG sleeves, high socks. a6 ce 35.1 34.6 34.7 Air temperature outdoors, 32° to 35°.6. | too......... 3350) 32°40 43-0 Relative humidity indoors, 60%. TOMPe ae 221 B25 33.0 eT | POR se gE 30 NON sO. OM On2 TEMPERATURE SUMMARY TQ) i 31.6 30.7 31.4 Kind No. Temp. at air vel. Oo oa 33-0 eo 32-5 Values 0 145-245 Dee eerie As 3G 3058) 3054 7 ° ° ° Saree 3048 | 20: 0) aGan SRGDE ere Om S4-91 3443) 1341.0)" aero tas 32 sve G22) 68205 Clothing = =-413, §431.8 30:8 30.7) 22%) |. 32) Ou Bon. 3386 Poteet Lee Se 33022 13029) ]. phen, non S40t- 34045 sar HOGS a ern 2) SLO: 30). 5>) 300 | TBai ss 34-4 1342. 3400 Walleeeerc lO) PGigil PXN5@) 29/672 Iefaee ee: 34.8 2257, Bano IRGC Goegoe 2 ZOROMEZ On te Zon: 17 e ee 4500 33.8 2346 |] 9......... 22) 5e" ao Ga hed RADIATION SUMMARY aa oe Soe ae 3 S oe Bee is) WSR Nee : : : Deenae eater Tete 3160) aie 8iimg0-9 145 245 20am hie 2363 30.2 30.9 Sinan sete tic HO720) 20073) 50583) 35eece ane a: Ome Lege 2053 Clothing. 12... ¥O420) 40352) £0320:| 30642. 0508 SES) |) B07, ag hss iar ea eee 50567) £0294) ©0340) .200.0-5. . 32.6 27.7 28.0 Shoesei.ch0, 46: HOAGO! §y03TO) | 7.0307%|| BOsvs. ak I-12 28255 2972 ae i a ee gE Wi RSy ren hore Bb.Om Qleape aha: Deharaer Stine Soh Tig eo siLenn aT 5S ToTaL RADIATION wallAs. 3. Dia 27 27 Air vel. Calories per hour Be ue 2780 27.1 2705 Dan ore Curves 27.9 28.0 28.3 Oo 25.8 De ee 26.8 26.9 27.0 145 20.8 esate A 26,5) 206 2703 245 Ee FOOM! Ae)... 26535) 264) 2606 SMITHSONIAN 50 ’ TABLE Ero DaTE: July 20, 1928. Place Se Gc. 0 SEx: Male. % AGE: IO yrs. II mos. See Ree = WEIGHT: 35 kg. wa eee oP HEIGHT: 135 cm. Guin 2 SURFACE AREA: I.14 sq. m. ee ae CLOTHING: Cotton waist, golf knickers, ate Be high socks. pate 2 Air temperature outdoors, 32° to 35°.6. | 22°77 34 Relative humidity indoors, 60%. ROPE ae ace 25 34 ROW teicaeieasne 23 LOA ee ne Ace 33 ‘TEMPERATURE SUMMARY TO wer aor ae 33 Kind No. Temp. at air vel. A eateries 33 Values e i se De, lee 32 Skinuanseuce 07 35525 3408) (345 Oe ee es 3I Clothing. 5.3228. 3252 3imoupe eee 33 Hatt. ic.cnte ls (BAe. 9 34a 328e BA eye 32 Shoesiaos oa 2 22-0. 2109 3282 ; 34 Wall jcnc are 10 26.9 27.2 27.3 ee Reacg ea ts a CRE es 2) 25-8. 26, i cane Room 558: 26.3 26"7, eee oe aA Pe Reh ore 2 RADIATION SUMMARY ms Aree s Calories per sq. cm. See ewe 22 per min. at air vel. Fi Serta A 21 2 as TENN 26a oe a 3 A 34 SID Ateraead ners 0768 .0703 .0664] 35......... 34 Clothing... 2 0533. -0457 -O425)1) 36......... 34, Plait cc neck aioe 0647 .0629 .0458] 29......... 32 Shoes.... -0462 .0431 .0451 | 30......... 32 DT Ea aie 31 TotaL RADIATION 2 One tae 31 Air vel. Calories per hour wall A sila caer tel 1s ahh per sq. meter Bee ee oii Oo 30.0 Cee ae. 27 145 26.3 De a 26 245 24.4 Eee oes 26 LOOMP a sae ae 26 ONT CODOWORWTHU HOD HHO DHONDO HO NAWUD OMDB OAD MISCELLANEOUS COLLECTIONS 145 ° 20% 26. 26. 27. 26. 26. DON ABA HUB HOH WON COWHER DOU NOW OANA OMNUO COU VOL. 8I Temperatures at Air Velocity 245 26. 26. 27m 28. oo . . do . . . er he . . e . . io OH HHUWNRONN O DOWN DION DUH ON DOW HPMNOODOOONON No. 6 BODY RADIATION—ALDRICH 51 TABLE F.—Summary of Cloth-covered Calorimeter Tests Calorimeter Wall Air Per cent Radiated by No. of A Temp. Vel. Melik. Thermoelement' Tests Preliminary tests— Vertical Estimated ..... oO 80 72 5 Horizontal Estimated..... : O 74 65 2a Final tests with cloth walls— Vertical Measured...... oO 61 60 5 Vertical Measured...... 75 39 41 2 Vertical Measured...... 130 41 35 2 Vertical Measured...... 190 40 25 2 Taste G—Summary Comparing all Thermoelement Temperatures with Cor- responding Temperatures Computed from Melikeron Values Given m Tables B, C and D Average difference Room temp.—Water Jacket temp. No. of Average difference Algebraic Arithmetic Place Observations Melik.—Thermo. Mean Mean Siig: ooh ok 37 I°9I 67 Tse @lothinesane. 49 Tet LG 1.45 laity cick < 9 30 1.61 1.61 Shoeseeceen se 8 1.39 5hh@ 1.20 VVialllersrserers ote 15 .03 137 1.37 Cloth-covered Cal. in still Alige en cree 52 . 56 .82 .95 moving air N30 ft... 16 53h) —.4 avi SS TOOMtecae 12 228 "2 T6 Clothing Hair 67 1.10 1.12 1.40 Shoes Clothing Hair 135 .80 82 Tots Shoes Calc for plants and animals, using host-parasite data as “ crucial evidence.” 1The porcupines are a peculiar group whose relationships to other rodents are not understood. It would be especially interesting to know what, if any, Mallophaga are found on African porcupines. 2 Geographical, not taxonomic, groups, in the four lands named above. 12 SMITHSONIAN MISCELLANEOUS COLLECTIONS voL. 8&1 One quotation, showing his effective use of hosts and parasites to- gether as sources of evidence, may well be given here. After dis- cussing again the Crayfish and their external parasites, the temno- cephaloids, he writes: The acceptance of Matthew’s hypothesis of four separate dispersal streams of crayfishes from the northern hemisphere potamobiids [northern crayfish] into Madagascar (through Africa), Australasia, New Zealand and South Amcrica implies: The presence in the past of temnocephaloids upon northern potamobiids, for which there is no evidence. The extinction of both crayfishes and temnocephaloids in Africa, where there is no evidence that either ever existed and no obvious or plausible reason... . why either or both should have become extinct. The general distribution in the past of both crayfish and their parasites in the tropical belt, for which again there is not positive evidence. Moreover, since opportunity has been afforded for the southern crayfish to migrate into the tropical belt, and since they have not done so to any marked degree, it would seem that the tropics do not afford a congenial environment for crayfishes. The extinction of temnocephaloids upon Asiatic and North American potamo- biids, for which there is no evidence, and which should not, I think, be assumed without some justification or explanation. These considerations seem to me to rule out Matthew’s hypothesis completely. If the parastacids [southern crayfish] have been derived from potamobiids, the only possibility seems to be that such derivation took place in America, and that the parastacids, as such, first appeared in South America, and must have reached the other southern land masses by a southern route of dispersal, carrying their temnocephaloid parasites with them. In 1928 two more papers by Harrison appeared. One (1928) presents important additional evidence as to antarctic zoogeography. A genus of lowly segmented worms, Stratiodrilus (one of the Histriobdellidae) occurs on fresh water crayfish in Tasmania, in New South Wales, and in Uruguay, and in this paper Harrison describes a fourth species on a crayfish from Madagascar. He discusses the family Histriobdellidae and its three genera and he prophesies that one or more species of the southern genus Stratiodrilus will be discovered on the gills of other South American crayfish (Parastacus) and of New Zealand crayfish (Paranephros). The second of Harrison’s papers in this year (1928) is an excellent general review of the whole host-parasite method. He had not learned of von Thering’s thorough-going use of this method of illuminating problems of genetic relationships of hosts, of geographical distribution of both hosts and parasites, and of former intercontinental connec- tions. Also he failed to realize the extent of Kellogg’s appreciation of the wide applicability of the host-parasite method. Harrison’s own realization of the broad value of such data apparently came from reading two of my papers and from correspondence with me in the year 1921, a correspondence which, though brief, was very valuable no. 8 PARASITES—METCALF 13 to me. But his grasp of the importance of parasites as indicating relationships of hosts was reached independently of von Ihering and Kellogg and much antedated my own. The following quotation shows Harrison’s grasp of the wide extent of the usefulness of the host- parasite method: The ostriches of Africa and the rheas or nandus of South America are commonly supposed by ornithologists to have arisen from quite distinct stocks. But their lice are so similar, and so different from all other bird-lice, that these must have evolved from a common ancestor, and so also must the birds them- selves. Evidence derived from lice is confirmed by the cestode and nematode parasites of the two groups of birds. Thus a phylogenetic relationship may be established by means of parasites. Equally, a supposed relationship may be refuted. Their lice prove that the penguins are in no way related to any northern group of aquatic birds, but belong in an ancient complex which includes the tinamous, fowls and pigeons; that the kiwis of New Zealand are modified rails, and not struthious birds at all; that the tropic-birds are not steganopodes but terns, and so on. A third use is to refute suggestions of convergent resemblance, which are often very lightly made, and which are so exasperating to the zoo- geographer since they are usually incapable of either proof or disproof. Lepto- dactylid frogs are found in South America and Australia. Did they evolve separately, or are they derived from common ancestors? The herpetologist cannot say with any certainty, but the parasitologist discovers that they share a genus, Zelleriella, of ciliate protozoan parasites, and must have had common origin. This same example will serve to illustrate a fourth use for the host- parasite relation. The genus Zellericlla can, and does, infest frogs other than Leptodactylids. It is not found, however, anywhere except in Australia, South and Central America, so that its distribution affords strong presumptive evidence that South America and Australia have been joined in past time in some way which excluded the northern land masses. These examples indicate the nature of the host-parasite relation, and its possible usefulness. In 1926 Harrison discussed before the Australian Association for the Advancement of Science “ The Composition and Origins of the Australian Fauna, with Special Reference to the Wegener Hypoth- esis.” This paper, in press but still unpublished in 1928, I have, of course, not seen. S. J. Johnston, of Sydney, Australia, had heard Harrison present before the Sydney University Science Society his first discussion of the biting lice (Mallophaga) of birds as furnishing evidence of the genetic relationships of their hosts (Harrison 1911) and two years later Johnston (1913) wrote of the frog trematodes of Europe, Amer- ica, Australia and Asia and their bearing upon possible former con- nections between these now separate lands. He concluded that the trematodes of Australian frogs find their nearest relatives in those of Asiatic frogs, and Grobbelaar, writing in 1922 upon African frogs I4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL. 81 and their trematodes, accepted this judgment of Johnston’s. Harrison questions this conclusion and predicts “ with the utmost confidence ” that future additions to the then very scant knowledge of Asiatic frog trematodes (six species) and the Trematoda of Australian frogs will show “that . . . . the closest affinities of Australian frog trematodes .... lie with those of South American frogs.” In this 1913 paper Johnston refers to trematodes of Australian sea eagles, sea gulls and herons and he points out also that two flukes of the genus Harmosto- mum found in two Australian marsupials are so closely related to another Harmostomum from a South American opossum that they “must be considered as derived from common ancestors.” Johnston must have had in mind the bearing of these parasite data upon prob- lems of former connection of Australia with Asia and with South America, but neither in this nor in two subsequent papers (1914, 1916) upon Australian trematodes and cestodes in general did he bring out clearly the paleogeographic importance of his data. He emphasized chiefly their bearing upon the genetic relationships between the hosts. Metcalf, the author of the present paper, was the fourth student of parasites to come independently to a‘realization of the important aid which parasites may give in solving problems outside the field of parasitology proper, and he used the host-parasite method in his earlier papers * much more extensively than it had been used before ; but really he added nothing essential to the conception of this method which von Ihering had in 1891 and 1902. Kellogg too seems to have realized the applicability of the host-parasite method to other problems than genetic relationships of hosts, though he made but scant, if any, application of it to them. Harrison carried Kellogg’s work upon bird relationships further and also in his papers subsequent to 1924 used parasite data extensively in problems of zoogeography and paleogeog- raphy. Priority is of very little interest, but, for what it is worth in this matter, the priority is clearly von Thering’s. Metcalf in his chief paper (1923) purposely overemphasized his data, endeavoring to bring out even slight suggestions which could not be established without corroboration from other sources.” His desire *Seven papers from 1920 to 1924; also one in 1928. 2““The endeavor will rather be to present the known data from the Anura and the Opalinidae and note their implications. Even very scant data, insufficient to have any real weight as they stand, will be stated and their implications noted, with the thought that even very minor items, of slight moment by themselves, may sometime be correlated with other data and then be of interest. The endeavor is, therefore, to have the treatment of this theme inclusive rather than critical.” (From Metcalf, 1923.) no. 8 PARASITES—METCALF 15 was not so much to prove certain particular taxonomic, zoogeographi- cal and paleogeographical propositions as to illustrate and emphasize the method of using parasite data in the study of such problems. That, indeed, is the chief purpose of the present paper also. Metcalf studied the opalinid parasites found in the preserved Anura (frogs and toads) in the United States National Museum, including species from all parts of the world. He was already familiar with those occurring in Europe. Other species were obtained from the Indian Museum at Calcutta and a few more from South America. Assuming the general correctness of a set of Mesozoic and Tertiary maps compiled by himself, chiefly from Arldt, von Ihering, Scharff and Schuchert, and based upon geological and biogeographical evi- dence, not including parasites, he studied conjointly the taxonomy and distribution of the Anura and their opalinid parasites and applied these data from biogeography, paleogeography and from the host- parasite studies, to problems of the place and time of origin of differ- ent hosts and groups of hosts, of different parasites and groups of parasites, to the routes, times and directions of dispersal of both hosts and parasites, and in the discussion pointed out evidence bearing on the correctness of the maps used, and upon problems of ancient climates. Before applying the data from the study of the opalinid parasites he tabulated the available data from both hosts and parasites under six items as follows: “‘ Species of opalinid; Host species; Family or subfamily of host; Known geographical occurrence of opalinid in the species of host named; Known occurrence of host; Known occurrence of genus of host.’ This tabulation, used in connection with maps of the present day oceans and of the continents in the several geologic periods, was of great aid in studying present and former distribution of both hosts and parasites, places and times of origin of each and routes and directions of dispersal. The publication of similar tables may properly be urged upon those who undertake comprehensive studies of any group of parasites. They will make the author’s data most easily available to other students and so should extend the general use of host-parasite data. Where data from fossils of either hosts or parasites are known and are sufficiently extensive they should be tabulated, say under such items as these: Geographic locality of fossils of the host family ; Geologic period of such fossils ; and, if fossil remains of the parasites are known, similar data as to them should be tabulated. Of course preservation of parasites as fossils will be rare, but their spoor may be found and may be quite specific, as, for example, in the case of the Peridermiums of pines, 16 SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL. 81 rusts which produce swellings of possibly specific character. Other examples would be bone lesions of recognizable cause. Let me here merely list a few of the things that seemed to be indicated with a greater or less degree of probability by these earlier studies of Metcalf. Having assumed paleogeographical maps showing certain intercon- tinental connections, he applied to them the data from Anura and their opalinid parasites and found they fitted in such a way as to be in general confirmatory. Protoopalina, the most ancient genus of the opalinids, was present in Equatoria (Australia plus Africa and South America) as early as the Triassic period, and its most archaic subgeneric group of species have persisted in these three continents, with only slight modification, until the present day. Other subgeneric groups of species of Protoopalina arose as follows: Group II in Australia at a time not indicated by the data. Group III before the separation of Australia from Asia in Jurassic or early Cretaceous times, in Australia or southeastern Asia, spreading to Europe during the Cretaceous or early Tertiary by a route north of the Himalayas, and to Africa in the late Tertiary, entering from the northeast. Group IV, in the Jurassic period, in Australia or southeastern Asia. Group V, in Cretaceous times in Australasia, their presence in Australia and Java but not in Sumatra indicating that Java retained connection with Australia longer than did Sumatra. The absence of members of this group from South America is one of several bits of evidence indicating that migration between South America and Australia was chiefly westward. Group VI, in the Jurassic period in Australia. Group VII, in Precretaceous or Cretaceous times * in South Atlantis which united Patagonia to South Africa. Group VIII, during the Tertiary period in western North America. Group IX, in Jurassic times in Lemuria (the Indian Ocean land connecting Madagascar and India, see fig. 3), with a Tertiary dis- persal to eastern Asia, Formosa and Java. The opalinids of the earliest Anura were apparently of the genus Protoopalina, as evidenced by structure, life history and distribution, since Protoopalinae occur in all families of Anura whose habits permit infection with opalinids. *Later studies tend to place this South Africa-Patagonia union somewhat later, in the early Tertiary. no. 8 PARASITES—METCALF 17 The genus Zelleriella arose in Patagonia, before the separation of Patagonia from Antarctica. This separation occurred probably in the middle Miocene. Zelleriella did not arise until Patagonia had lost its African connection, for the genus does not occur in Africa. In the early or middle Tertiary it spread to Australia ; in the late Tertiary to Tropical America. Its original hosts were southern frogs (lepto- dactylids). Its presence in South America and Australia, and its absence from Euro-Asia is, when carefully studied, as already noted, evidence of former southern land connection between these continents. To continue merely listing the things indicated by Metcalf’s host- parasite data from Anura and their opalinids would be wearisome, so we will omit reference to the genera Cepedea and Opalina and their subgenera, whose times and places of origin and times and routes of dispersal were discussed, and will note further only some of the types of conclusions suggested. Evidence was found as to the places and times of origin of the several families of frogs and toads, and the routes by which, and the times at which, they spread to the lands they now occupy. There are similar indications as to a number of genera of the hosts, Bufo, Polypedates and Rana, for example. Spread of true frogs (Raninae) from the north into South America has not occurred, except for one species, and there are no indications of any southward wandering of Anura across the Isthmus of Panama since its formation in the Middle Pliocene. On the other hand, there has been extensive spread of Anura northward across this Isthmus. The Sonoran desert of northern Mexico and the southwestern United States has been a hindrance to northward wandering of southern frogs since the middle Pliocene, but has not held back the tree frogs (Hylidae). Negative as well as positive evidence is often given. For example, the absence of Zellericlla—the characteristic opalinid of the southern frogs—from Euro-Asia indicates that southern frogs were never in Euro-Asia. The absence of the genus Opalina from South America, though it is present in the toads (Bufo) in Central America, shows that toads have not passed south across the Isthmus of Panama since Opalina, a Tertiary immigrant from Asia, reached Central America. Again the only Euro-Asian species of tree frog (/yla arborea) with its several subspecies is not endemic in Euro-Asia, but is an immigrant from North America, for it carries a North American Opalina. This recital of a few of the indications from Metcalf’s studies is sufficient to emphasize the point here in view, namely, that host- : : SS 658 * > eg ES SEE ne i bee ae SE Set man 18 SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL. 81 parasite data may be applied to a great range of problems. This, which we might well name the von Ihering method, gives decisive results in many cases, while in other instances it furnishes merely corroborative evidence or evidence to be joined with that from other sources. Metcalf subsequently published several papers discussing the host- parasite method or host-parasite data, as noted in the appended bibliography. Darling (1921, 1925) used data from the hookworms of man to indicate human origins and migrations. Before the publication of this earlier paper Darling had very likely not read the papers of von Ihering, Kellogg, Harrison, Johnston and Metcalf, which had made somewhat similar use of parasite data, for he does not refer to these authors. It seems probable, therefore, that Darling may have been another independent discoverer of the broad significance of such data from parasites. The following quotation will show Darling’s sug- gestions: . . man of the Holartic regions [is] parasitized exclusively or almost ex- clusively by Ancylostoma duodenale, while man of the Oriental and Ethiopian regions [is] parasitized exclusively or almost exclusively by Necator americanus. This .... suggests the possibility of .... there having been two primitive races of man, each one originally parasitized by a particular species of worm. Certain it is that N. americanus is found more exclusively among biack and _ brown-skinned races, while 4. duodcnale is found exclusively or greatly pre- dominates at the present time among Caucasian and Mongoloid stocks. It may be that a Eurasiatic race of men, possibly the Pithecanthropus of Trinil, Java, became split off and furnished the stock from which man of oriental and Ethiopian regions sprung. Proliopithecus emerging from Holarctic Africa may have been not only the parent form of man, gibbon, chimpanzee, gorilla and the orang-outang, but he may have harbored the parent form from which have arisen the different hookworm species found in the various species of anthropoids of today. Possibly the ancestral tree of the primates can be revised after a study of the host relationships of their respective obligate nematode parasites. At any rate we can say that it seems likely from the present distribu- tion of A. duodenale and N. americanus as determined in surveys recently made of selected groups that there were originally races of man parasitized exclusively by A. duodenale and inhabiting the Holarctic region, that is Europe, Asia, north of the Oriental region, and northern Africa; and that there were other races of man parasitized exclusively by N. americanus and inhabiting the Oriental region, that is the southern peninsulas of Asia and Indoasia or the Malay Archi- pelago; and also the Ethiopian region, that is, Africa south of the Sahara Desert. Ewing (1924) in a study of biting lice of the family Gyropidae discusses the significance of their geographical and host distribution arguing in favor of a crossing over between rodent hosts and primate no. 8 PARASITES—METCALF 19 and ungulate hosts rather than descent from common ancestors. In a second paper (1924a) Ewing discussed the host-parasite relations of human and louse races and the hybridization of both and he includes in this discussion prehistoric races of men and of their head lice, and he mentions again the probability that the tropical American spider monkeys (Ateles) acquired their head lice (Pediculus) “ originally from man but not from recent man.” Two years later the same author (Ewing, 1926) discusses further the significance of the geographical and host distribution of the genus Pediculus. Four paragraphs of his summary may well be quoted: 1. In America two distinct groups of Pediculus exist, one of them confined to man and one to monkeys. 2. The forms infesting man are apparently largely hybrid races of head lice, the pure strains of which were originally found on the white, black, red, and yellow races of man living in their original geographic range. 6. The monkey-infecting pediculids of America, so far as known fall into distinct species according to the hosts they infest, thus indicating, to a certain degree at least, a parallel host and parasite phylogeny. 7. If these monkey hosts (Ateles, species) procured their lice from man it was not from recent man but from human hosts that lived tens of thousands of years ago—long enough to allow a species differentiation to develop among the monkey hosts. Ward (1926), ina presidential address before the American Society of Parasitologists, has mentioned the importance of such uses of data from parasites and refers in this connection to some of the work reviewed in the present paper. Hegner (1928) discusses the protozoan parasites of monkeys and man and concludes with the following statement : _... the protozoan parasites of monkeys and man belong for the most part to the same species or are so similar in their structure, life-cycle and host- parasite relations as to be practically indistinguishable. This situation is par- ticularly striking when the protozoa of monkeys are compared with those ol other animals associated with man. If the proposition that close relationships of parasites indicate a common ancestry of their hosts is valid, then the facts available regarding the protozoan parasites of monkeys and man furnish evidence of importance in favor of the hypothesis that monkeys and man are of common descent. This shows Hegner’s recognition of the importance of host-parasite data in studies of phylogeny. Some few students have attempted to minimize the importance of parasite data in problems of biogeography (Noble, 1922, 1925 ; Dunn, 1925). Harrison (1924, 1926) has sufficiently answered their criti- cisms. Noble’s criticisms are based largely upon his new classification of the Anura, a classification not as yet accepted by herpetologists. 20 SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL. 81 The present writer thinks improbable Noble’s idea that the southern frogs of Australia and those of South America evolved independently from the archaic toads, and developed along parallel lines. So far as I can learn, the papers mentioned cover the use thus far made of data from parasites in connection with the three classes of problems here considered. So little has been done in this field and so little has that little been known that each successive student has thought himself a discoverer and a pioneer. It has been probably a unique incident in biological and geographical science. There have been instances of double or triple discovery—mutation, for example— but sixfold independent discovery of a concept with wide significance and capable of important application in further research has probably not before occurred. We have described in outline the use that has been made of this “von Ihering method.” It seems well before closing this paper to suggest possible further applications of the method, using other groups of parasites, and to mention some specific problems needing study by this method. Harrison (1928) has reviewed from this point of view different groups of animal parasites considering their availability for host-parasite studies. Let us include plants as well. Protozoa—There are, of course, many groups of Protozoa part or all of whose members are parasites or commensals, having at any rate an obligatory association with definite animals or plants. Among the Sarcodina are many parasitic Amoebae and a few Heliozoa are inter- nal parasites. I know of no use of data from these forms in studying such general problems as we have had in mind. Our knowledge of the taxonomy of these parasites, of their host-occurrence and of the geographical distribution of both parasites and hosts is inadequate, but the material for such host-parasite studies in these groups seems to be available. There is a considerable degree of specificity in the host relations of the Endamocbae and they are found 1n many groups of animals. Multitudes of the flagellates are parasitic and probably no other group presents more advantageous material for host-parasite studies. Some flagellates are parasitic in plants. Although knowledge of flagel- late parasites is extensive, it is very fragmentary, being almost nil for many regions of the earth and far from complete for most regions and for most hosts. In some groups we have enough records to begin tabulating the host occurrence and geographical occurrence and scru- tinizing the tables for what they may indicate. Probably the finest groups for host-parasite studies are the termites (white ants) and the no. 8 PARASITES—METCALF 21 flagellates living in their intestines. Approximately fifteen hundred species of termites are known and from all tropical and many tem- perate parts of the world. They have a highly elaborate taxonomy with four families, subfamilies, genera, subgenera, species and sub- species, and the genetic relationships and the phylogeny seem capable of successful study. Forty-six genera comprised in 12 families of termite flagellates have been described from less than 40 species of termites, this being but a meager beginning of the taxonomic and phylogenetic study needed for this truly vast number of mostly unde- scribed species. It seems unlikely that any other organisms will lend themselves so favorably to host-parasite studies as will the termites and their flagellates. Every individual termite is richly infected. The wealth of species of these hosts and of their Protozoa is so great as to be somewhat awesome. ‘“‘ There are probably more flagellate Protozoa in the intestines of termites than in all other animals com- bined.”? It is a bold student who attacks these groups with the idea of employing them by the von Ihering method, but the one who does so should reap a rich reward. The termites are a peculiarly favorable group for such studies be- cause, in addition to their varied internal fauna of flagellate parasites, they harbor, either customarily or occasionally, representatives of every other group of parasitic Protozoa ( Amoebae, Ciliates, Sporozoa ) so that one studying them through their flagellates would often be able to check up results from some of their other parasites. The Chlamydozoa are but little understood. It seems not unlikely that when better known, especially if they prove to be associated with mozaic and other filtrable virus diseases, they may prove of much interest. The Sporozoa offer much fine material for host-parasite studies, all being parasitic. Most species of terrestrial and fresh water animals harbor representatives of one or more of the numerous groups of Sporozoa, and they infect also very many marine animals. Many Sporozoa, perhaps most of them, show a high degree of specificity in their selection of hosts, being confined each to one species of host or to one taxonomic group of hosts. This renders their evidence in some instances peculiarly convincing. Among ciliate Infusoria are numerous parasitic species. Balantidium and Nyctotherus, parasites of man and other mammals, should be valuable for host-parasite studies. The “Astomata,” which include several perhaps unrelated families, should also furnish favorable ma- 1 Cleveland, L. R., quoted from a letter. 5 22 SMITHSONIAN MISCELLANEOUS COLLECTIONS voL. 81 terial. But best of all ciliates for such studies seem to be the archaic group, the prociliates,’ including only the Opalinidae, the basis of Metcalf’s studies, to which reference has already been made. The inability of their hosts, the frogs and toads, to endure salt water makes their evidence as to land routes of dispersal peculiarly cogent. Opalinids have remarkably clearly indicated phylogenetic relationships (Metcalf, 1926), probably more clearly indicated than in any other group of Protozoa. These two groups, the Anura and their opalinids, are thus peculiarly favorable for studies by the host- parasite method, especially studies of the phylogeny of the Anura and of their geographic dispersal. The Ophryoscolecidae, a group of ciliates which live in the stomachs or intestines of ungulates, anthropoid apes and some South American rodents, have a highly diversified taxonomy, with relationships well indicated, are almost world-wide in distribution and seem, from our present inadequate knowledge, to be specific as to their hosts. They and their hosts should furnish important host-parasite data. No animals are better represented in fossil records than are the Ungulata. Among the flatworms the Temnocephaloidea, with the crayfish on whose gills they are parasitic, have been used very effectively in host- parasite studies by von Ihering and Harrison, as already noted. Von Thering and Johnston have made similar use of data from the flukes (trematodes), the tapeworms (cestodes), and some of their hosts. But the important results already obtained by aid of evidence from the flatworms are but a very minor fraction of the harvest that may be reaped by adequate study of this group. Darling’s studies of the origin and spread of human races in the light of their hookworm parasites are an example of the use of data from round worms (Nematoda). Among the Nematoda there are innumerable free-living forms, and great numbers of parasitic species infesting almost all kinds of animals and very many kinds of plants. A parasitic nematode is even known from a ciliate infusorian—a metazoan parasite in a protozoon. There is in the parasitic members of this group and their hosts a wealth of material which should prove an inexhaustible mine for working by von Ihering’s method. The nematodes rival the trichonymphs of the termites as a source of data for such use, indeed because of their universal abundance and the huge number of their species they must surpass the trichonymphs in the number and variety of problems their evidence will help solve. *Using Wenyon’s (1926) modification of Metcali’s name “ protociliates ”’. no. 8 PARASITES—METCALF 23 Harrison, as already described, has made use of parasite data from Stratiodrilus, a genus of archaic annelids, to indicate intimate relation between Australasia, Madagascar and South America. The annelids as a class, however, are poor in parasitic species. Among the Crustacea the parasitic copepods may perhaps give light upon some interesting problems, though their host relations and especially the specificity of these relations need further study. The parasitic species of copepods are apparently chiefly ancient and reached for the most part their adaptation to parasitism long age, having undergone little modification in later geologic periods. Others, however, seem to have adopted parasitism more recently. A thorough analysis of the parasitic copepods from this point of view would be worth while for its own sake and would give added significance to their host-distribution and geographical distribution. Among the Arachnoidea (spiders, mites, ticks, etc.) several groups are parasitic, but the parasites are not confined each to one individual host or even to one species of host. They are free to pass from one host to another. This makes them far less useful for host-parasite studies than are more restricted parasites, but, in some instances at least, they present usable data. The true insects include many groups among whose members parasitism is more or less well developed. Examples of insect parasites of terrestrial vertebrates and of insect parasites of insects at once come to mind, but with these insects, as with the mites and ticks, specificity of host-infection is in general not highly developed, though there are numerous exceptions in which there is constant relation between kind of host and kind of insect parasite, as, for example, some moths parasitic in bee colonies and some beetles restricted to ant nests. Many insects parasitic upon plants have closely specific host limita- tions, being confined each to a single host species or to a related group of species, however freely they may pass from host individual to host individual. One thinks at once of the plant lice (Aphides), but many even of the larger insects have similarly restricted plant prey—eé. 9.) the potato beetle, the squash bug, the plum curculio, the hessian fly, the cotton boll weevil, grape Phylloxera, some butterflies, some moths. many gall-flies, etc. Molluscs, echinoderms, vertebrates and other chordates, show few examples of parasitism, commensalism or obligate association of any kind. It is doubtful if the few cases known (shark-Remora, fish living among the tentacles of jelly-fish, fish living within sea cucumbers, fish 24 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 81 living in the mantle cavity of molluscs, and some others) will prove of much interest from the point of view of the present paper. Parasitic plants have never been used, so far as I can learn, in such studies as those in which we are here interested, though they present a great mass of usable host-parasite data, but in all the groups which Fic. 1.—The Atlantic Ocean and the adjacent land areas. The dotted lines indi- cate 2000 fathoms depth. (Modified from a map by W. & A. K. Johnston.) furnish these data much further study is needed. A good degree of specificity between host and parasite is a desideratum and this we find in a good many cases. The rusts are very favorable in some regards. Most of them are restricted in their hosts, many cause lesions which can readily be recognized, as, for example, the Peridermiums of pines and the branch “nests” of cedars. Many of the rusts of the conifers produce distor- tions in the hosts which could be identified in fossils. The two hosts, no. 8 PARASITES—METCALF 25 intermédiate and definitive, necessary for each species of rust, present a most interesting condition for distributional studies. The necessity for two hosts in the life cycle of a rust, presents a complication, but one which makes the evidence from the rusts and their hosts more than doubly significant. On the other hand the rusts lack one ad- Fic. 2.—The Pacific Ocean and the adjacent land areas. The dotted lines indicate 2000 fathoms depth. (Modified from a map by W. & A. K. Johnston.) vantage—their taxonomy is not well understood. This disadvantage 1s only partly compensated by their large number of forms and their numerous and diverse hosts. When the rusts are more widely and more thoroughly known they will present data of peculiar value in host-parasite studies. The smuts of grasses, especially of uncultivated grasses, might furnish data; so also the powdery mildews (Erisiphaceae) and the downy mildews (Peranosporaceae) , especially those infesting unculti- vated species. 26 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 81 Mycorrhizae, commensal root fungi, of pines and many other groups might be of especial interest, first because the data they furnish might be compared with those from rusts and other fungi, and, second, because they produce lesions which possibly might be recognizable in fossils. Fic. 3—The Indian Ocean and the adjacent land areas. The dotted lines indi- cate 2000 fathoms depth. (Modified from a map by W. & A. K. Johnston.) The fungi in general should be scrutinized for groups fitted for such studies. Fungus diseases of plants are being more and more studied and new data are thus being offered. Plants and their parasites, when studied by the von Ihering method, will surely give very important results, but such study must be accom- panied by further and laborious study of the structure, life history and taxonomy of the,parasites. Fossil records of the hosts are of especial interest in biogeographical problems and if these can be joined with fossil records of the parasites no. 8 PARASITES—METCALF 27 also it is still more fortunate. This cannot be expected in many cases, but there is prospect of some success in such study of bones of Vertebrata and their lesions (Moodie, 1923; Rupper, 1921), of conifers and their distortions caused by Peridermiums and My- corrhizae, of some other plants and their scars from fungus diseases, of many plants and their insect galls and probably of still other groups of animals and of plants showing fossil records of parasites. This paper may well close by suggesting as samples one or two special problems favorable for attack by the host-parasite method. We have already noted crucial data presented by parasites of several groups as to the problem of. east and west routes of dispersal in the Southern Hemisphere. The parasites of both plants and animals which show families, genera, and especially species, common to different southern lands, and southern lands only, may well be studied further. Such studies should finally determine not only the question of the former existence of such east and west migration routes, but also their position, their connections and their geologic time. On the other hand, if in some groups the dispersal was southward from northern lands, this fact will be demonstrated beyond dispute. Let us note here a partial list of species, genera, and families of southern occurrence whose parasites of all kinds should be studied (cf. figs. 1, 2 and 3). Mammalia The marsupials of Australia and of America (mostly tropical America). Their biting lice (Mallophaga) have been somewhat studied, so also their flukes (Trematoda) and tapeworms (Cestoda). The porcupines (Hystricomorpha) of America (mostly tropical America) and of Africa. Edentata (sloths and anteaters) in South America, South Africa, southern India, Malaysia. Birds Struthiornidae (ostrich family) with species—2 in New Zealand, 2 in Australia, 1 in Papua, 2 in South America, 1 in Madagascar. Trogonidae (the quetzal and its relatives) in South America, Central Amer- ica, Africa, and southern India. Chionidae (sheathbills) Antarctic Islands Psittacomorphae (parrots) in the Southern Hemisphere, with “stragglers ” in North America and some in India. Paristeropodes (a group of fowls) in Australia and South America. The Ocydromine Rallidae (rails) 3 in Australia, Heterochloa in New Zealand and also in Madagascar. Avocets and stilts in Australia, New Zealand, South America and Africa. Penguins in Australasia (including New Zealand and its Antarctic islands), South America, Africa, Antarctica, Antarctic islands in general, including St. Paul in the Indian Ocean. It is interesting to note 28 SMITHSONIAN MISCELLANEOUS COLLECTIONS voL. 81 that six species of fossil penguins were found in Graham Land by the Swedish Antarctic Expedition. In this connection note also that fossil Spheniscidae are known from New Zealand and Patagonia. Reptiles Giant tortoises in Galapagos Islands and in Malaysia. Amphibia Anura Leptodactylidae (southern frogs) in South America, Central America, West Indies, Australia, Tasmania, Papua, South Africa. Hylidae (tree frogs) in America (mostly tropical America), Australia, Tasmania, Papuasia, I species with several subspecies in Euro-Asia. Pipidae (Surinam toad, etc.) in Guiana and South Africa. Archaic Bufonidae, of other genera than Bufo, in Australia, north- western South America, Central America, tropical Africa, southern India, Ceylon, Malaysia. Gastrophrynidae, in Papuasia, tropical America, Africa, Madagascar, southern India, Ceylon, Siam.* Dendrobatinae in northwestern South America, southern Central Amer- ica, western Africa, Madagascar. Urodeles Coeciliadae (blindworms) in tropical America, tropical Africa, southern India, Ceylon, western Malaysia. I'reshwater fishes Cichlidae in tropical South America, Central America, Cuba, Africa, Mada- gascar, southern India, Ceylon. Characinidae in tropical America and tropical Africa. Galaxiidae in New Zealand, Australia, South America, the Falkland Islands, southern Africa. The genus Galavias occurs in New Zealand, Tasmania, southern Australia, the southern extremity of South America, the Falkland Islands. Osteoglossidae in South America and South Africa. Haplochitonidae in South America and South Africa. Dipnoi (lung fishes) in South America, tropical Africa, Australia. Molluscs Tertiary fossil species common to New Zealand and South America are named by Chilton (1909) as follows: Epitonium rugulosum lyra- tum, Crepidula gregaria, Turitella ambulacrum, Cucullaea alta, Venericardia patagonica, Brachydontes magellanica. This com- munity of species is of much interest and suggests a review of modern littoral mollusca and their parasites from the two regions. Arthropods Insects Ants—Notomyrmes in New Zealand and Chili, Prolasius in New Zea- land and its close relatives, Acanthoponera and Lasiophares, in South America. The following annotation from Emery (1895) is worth noting: _ Chili is, however, an isolated country, which we may call “a con- tinental island,” although it is not surrounded by water. If we should take the Chilian fauna as a standard for the primitive fauna of *The report of a gastrophrynid from Samoa is questioned. No. 8 PARASITES—-METCALF 29 von Ihering’s Archiplata, that should have been a very poor one, like the fauna of New Zealand, with which it offers a striking resemblance. The most characteristic feature of the Chilian ant fauna is the occurrence of peculiar species of Monomosium, like those inhabiting Australia and New Zealand, and of the genus Melophorus found only in Australia and New Zealand. These facts corroborate the hypothesis of a Cretaceous or Eocene connection between South America and Australia. New Zealand appears as a bit of old Australia, quite free from later Papuan or Indian intrusions, like Madagascar, which as an isolated part of old Africa, had received but a few immigrants, when, at the Pliocene epoch, a stream of Indian life entered into the Ethiopian continent. Probably Chili may be considered as a part of ancient Archiplata, secured from Guyanean and Brazilian immigrants by the heights of the Cordillera, but having preserved only an incomplete set of the original Archiplatean fauna. Beetles—Longicornia in Australia, New Zealand, South America; Buprestidae in Australia, New Zealand, South America. Flies—Zaluscoides in the Auckland Islands; the closely related genus Zalusca in Kerguelen. Peripatus—in Australia, South and Central America, South Africa, Peripatus (sensu stricto) in South America and South Africa. Arachnoidea (spiders, etc.) Myro (a spider) with species—2 in the Antarctic Islands of New Zea- land, t on Kerguelen Island, 1 at the Cape of Good Hope. Rubrius (a spider) Antarctic Islands of New Zealand, Tasmania, South America. Pacificana cockayni (a spider) in the Antarctic islands of New Zealand ; a related species in Tasmania and a closely related species at Cape Horn. Cryptostemma westernmanni (?) in tropical America and_ tropical Africa. Cercoponius (a scorpion) in Australia, South America. Crustacea—Land and freshwater forms: Parastacid crayfishes in Australia, New Zealand, South America (with one “ wanderer ” in California), Madagascar. Their gill flukes have been studied by Harrison, so also their Histriobdellidae. Trichoniscus, a subantarctic genus. One species occurs in the subant- arctic islands of New Zealand, Fuegia, Falkland Islands. Deto in Australia, New Zealand, Chatham Islarids, Auckland Islands, Chili, Cape of Good Hope, St. Paul Island. The species D. auck- landiae occurs in New Zealand and Chili. Idotoea lacustris in New Zealand, Campbell Island, the Straits of Magellan. Annelid worms. Many of the commonest New Zealand polychaetous annelids are identical with those of Magellan Strait, Fuegia and Chili. A comprehensive study of these worms and their parasites from these regions should prove of much value. Chilton (1909) says “of 13 species in the subantarctic islands of New Zealand only 2 are endemic in New Zealand, 8 are found in South America or the Falkland Islands, and 2 extend to Kerguelen”. 30 SMITHSONIAN MISCELLANEOUS COLLECTIONS voL. 81 Plants have not been studied through their parasites by the von Ihering method. On the chance of possibly interesting some botanists it may be worth while to list a few plants of interest in connection with southern dispersal. The forms listed seem to indicate: some a dispersal from northerly lands southward, but many more a dispersal! eastward or westward between southern lands, some by way of Ant- arctica. Omitting less conspicuous forms, note the following ferns and flowering plants: Ferns Polytrichum vestitum—Australasia, South America, islands of southern Pacific Ocean. P. richardi—Australasia, Southern Pacific islands. Asplenium flaccidum—Australasia, South America, Africa. Blechnum penamarina—Australasia, South America. B. capense—Australasia, South America, Africa. Hesteopteris incisa—cosmospolitan in the tropics. Pteridium esculentum—Australasia, South America. Polypodium billardieri—Australasia, Malaysia, South America, Africa. Hymenophyllum ferrugineum—Australasia, South America. H. tunbridgense—Australasia, South America, Africa. Dryopteris punctata—Australasia, South America, islands off South Africa. Polystichum adiantiformae—Australasia, South America, southern Pacific islands. Asplenium adiantoides—Australasia, Africa, islands of southern Pacific. Poesia scaberula—Australasia, Africa. Flowering plants Cypresses: Callitris in Africa, Madagascar, Australasia; Fitzroya in Chile, Tasmania. Hierochloe redolens (grass), Australasia, South America, southern Pacific islands. Monimiaceae: Tasmania, New Caledonia, New Zealand, Madagascar. Saxifragaceae—35 genera in Australasia, Madagascar, South Africa and South America, only 2 of which cross the equator. Proteaceae: 48 genera, 950 species in South America; 32 genera, 250 species in South Africa. Verbenas: Petraca in South America, Timor, Java; Petraeovitex, (a close relative) in Bouru and Amboina. Species common to Australasia and South America: Sedges as follows: Scirpus inundatus (extending to islands of the south Pacific), Carex darwinii, and its subspecies urolepsis, C. trifida; Luzula racemosa; Luguriaga parviflora (Liliaceae) ; Colobanthus quitensis; Crassula moschata; Geum parvifloruwn; Sophora tetraptera (the kowhai tree) ; O-xalis magellanica; Geranium sessiforum; Pelargonium australe, (New Zealand, Australia, Tristan da Cunha); Coriaria ruscifolia, C. thymifolia; FEpilobium conjugens; Veronica elliptica. no. 8 PARASITES—METCALF an Genera common to Australasia and South America: Drimus (3 species in New Zealand, 1 in Tasmania, 1 in Fuegia, 1 Tertiary fossil, D. antarctica, in Graham Land); Araucaria (1 Australasian, 2 South American, Norfolk Island 1, New Caledonia several, 1 fossil, A. imponens, in Antarctica, also 2 related fossils Araucaritis and Dadoxylon); Lomatia (6 species in Australia and Tasmania, 3 in Chili, also 4 Tertiary fossil species in Antarctica) ; Embothryum (1 Australian, 4 South American); Prionites (1 each Tasmania and Fuegia) ; Eucryphis (1 Tasmania, 1 Australia, 2 Chili) ; and others—Leptocarpus, Orites, Aristolochia, Drapetes, Terpnatia, Myosotis, Phyllaceus, Lagenophora, Leptinella, Enargea, Luzuriaga, Geranium, Azarella; Oreomyrrhis, Pernettia, Plantago (subgenus Plantaginella), Oreobolus, Carpha, Uncinia, Gaimarcia, Marsippospermum, Rostkovia, Libertia, Nothophagus (Tertiary fossils, 4 species in Antarctica), Caltha (Psychrophila), Drosera (one subgenus), Eucryphia, Gunnera, Prionotes, Tetrachondra, Pratia, Donatia, Abrolanella. Genera common to New Zealand and South America: Griselina (4 species in Chili, 2 in New Zealand) ; Ourisia (19 in South Amer- ica, 8 in New Zealand) ; Discaria (18 in temperate South America, 1 in New Zealand, 1 in Australia) ; Gaya (10 in South America, 1 in New Zealand) ; Fuchsia (60 American from Mexico to Fuegia, 3 in New Zealand) ; Jovellana (2 in Chili and Peru, 2 in New Zealand) ; Phrygilanthus (20 in South America, 2 in New Zealand, 4 in Australia) ; Muehlenbeckia (10 in South America, 4 in New Zealand, 7 in Australia, one of them extending to New Zealand, 1 in the Solomon Islands) ; Laurelia (2 in southern Chili, 1 in New Zealand, 1 fossil in Graham Land, Antarctica) ; Dacrydium (many in New Zealand, 1 in Chili) ; Pseudopanax (5 in New Zealand, 2 in southern Chili). Two paragraphs from Cheesman (1909) might be quoted: Of 37 species of flowering plants and ferns known from the Kerguelen-South Georgia region, 20 are found also in the subantarctic islands of New Zealand while 27 are found in Fuegia and the Falkland Islands. The total number of Fuegian plants found in the subantarctic islands of New Zealand is 29, 14 of these extending also to the Kerguelen and South Georgia groups of islands. These figures deal only with the specific identity; if we consider the genera, we find that, out of 88 genera found in the subantarctic islands of New Zealand, there are no less than 56 with representatives in Fuegia. Eleven species of plants found in the subantarctic islands of New Zealand are found either in the Tristan da Cunha group in the South Atlantic or in the Amsterdam Island group in the Indian Ocean, the flora of these two groups possessing many points of agreement notwithstanding their wide separation and showing also undoubted traces of affinity with those of Fuegia and Kerguelen. Two of these 11 species, however, do not occur in Fuegia or the Kerguelen-South Georgia group of islands. What parasites, if any, can best be studied to test and extend the significance of the distribution of these and other southern hemisphere plants? Will they be some group or groups of fungi? Will predatory 32 SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL. 81 insects of restricted food habits help? Will gall-forming insects give some light? How about nematodes? Will plant-feeding snails help? The last few pages have noted a few sources of data for but one set of problems connected with the biogeography of the Southern Hemi- sphere. There are many other problems and groups of problems. Let us mention only one other. It is thought that in Cretaceous times there was a strip of land running north from Japan, Korea and Kamchatka, crossing the Fic. 4.—Hypothetical composite map of the Pacific Ocean and adjacent lands during Cretaceous times, showing the land-strip bounding this ocean on the north and east and extending westerly from South America across the southern Pacific to Papua and Australia. Not all parts of this land-strip were in existence at any one time, the northern portions being mostly earlier, the South-Pacific bridge being later, perhaps early Tertiary. (Compiled from several authors, chiefly Arldt.) northern Pacific Ocean and running down the west coast of America to Ecuador and the Galapagos Islands (fig. 4). This circumpacific land strip may have connected at its southwestern end with the northern Malayan region (cf. fig. 2). It is thought to have connected with Eastern Asia in perhaps numerous places. It may have included the Aleutian Islands or may have lain mostly to the south of them. The mountainous islands of western Alaska, Vancouver Island, the Olympic mountains and the Siskiyou mountains of Northern California were probably included ; so also may have been Mount Tamalpais, the Pre- sidio Hill, the southern California islands, the tip of Lower California no. 8 PARASITES—METCALF 33 and the middle portion of Central America where the mountain ranges have an east and west trend. Upon the American portion of this circumpacific land strip is a very interesting relict fauna and flora including, to name but a very few, the bell-toad Ascaphus (an immi- grant from Euro-Asia who brought with him his characteristic Euro- Asian bell-toad parasite, Protoopalina, of an ancient subgenus) and a number of plants, conspicuous among which are several pines—the Monterey Pine, the Torrey Pine, Pinus jeffrey. Study of these western relict pines and their rust and other parasites and comparison with East Asian pines and their parasites might prove of much im- portance. We should remember, too, that the Peridermiums of pines produce lesions which should be recognizable if preserved as fossils. LITERATURE €lkeD ANDREWS, Justin M. 1927. Host-parasite specificity in the Coccidia of mam- mals. Journ. Parasitology, Vol. 13, No. 3, March. CHEESMAN, T. F. 1906. Manual of the New Zealand flora. 1909. Systematic botany of the islands south of New Zealand. The Subantarctic Islands of New Zealand, Vol. I, Wellington, N. Z., John Murray, Govt. Printer. CHILTON, CHARLES. 1909. The biological relations of the subantarctic islands of New Zealand. (Summary of results). The Subantarctic Islands of New Zealand, Vol. II, Wellington, N. Z., John Murray. Govt. Printer. CocKAYNE, L. 1907. Report on a botanical survey of Kapiti Island. Dept. of Lands, New Zealand Govt. 1908. Report on a botanical survey of the Tongariro National Park. New Zealand Dept. of Lands. 1908a. Report ona botanical survey of the Waipoua Kauri Forest. New Zealand Dept. of Lands. 1910. New Zealand plants and their story. N. Z. Board of Science and Art, Manual No. 1, Wellington. John Murray, Govt. Printer. This edition is preferable to the later one. 1921. The vegetation of New Zealand. W. Engelmann, Leipzig. Dar.ine, S. J. 1921. The distribution of hookworms in the zoological regions. Science, Vol. 53, April 8. 1925. Comparative helminthology as an aid in the solution of eth- nological problems. Amer. Journ. Trop. Med., Vol. 5. Dunn, E. R. 1925. The host-parasite method and the distribution of frogs. The American Naturalist, Vol. 50, No. 663, July-August. Emery, C. 1895. On the origin of European and North American ants. Nature, Vol. 52, Aug. 22. Ewrne, H. E. 1924. On the taxonomy, biology and distribution of the biting lice of the family Gyropidae. Proc. U. S. Nat. Mus. Vol. 63, Mch. 4. 1924a. Lice from human mummies. Science, Vol. LX, No. 1556. 1926. A Revision of the American lice of the genus Pediculus, together with a consideration of the significance of their geographical and host dis- tribution. Proc. U. S. Nat. Mus., Vol. 68, June ro. 34 SMITHSONIAN MISCELLANEOUS COLLECTIONS voL. 81 Forses, H. O. 1893. The Chatham Islands. Their relation to a former southern continent. Abstract in Nat. Sci. Vol. 3. Gapow, Hans. 1909. Amphibia and Reptilia. Cambridge Natural History. Harrison, LAUNCELOT. 1911. The taxonomic value of certain parasites. Ab- stract in Ann. Rep. Sydney Univ. Sci. Soc., 1911-1912. 1914. The Mallophaga as a possible clue to bird phylogeny. Austral. Zoologist, Vol. 1, No. 1. 1915. Mallophaga from Apteryx, and their significance. Parasitology, Vol. 7, No. 4: 1915a. The relation of the phylogeny of the parasite to that of the host. Abstract in Rep. Brit. Assoc. Adv. Sci., 1915. 1916. Bird-parasites and bird-phylogeny. Ibis, April, 1916. Abstract and discussion in Bull. CCXII. Brit. Ornith. Club. 1922. On the Mallophagan family Trimenoponidae, with a description of a new genus and species from an American marsupial. Austral. Zoologist, Vol. 2, No. 4. 1924. The migration route of the Australian marsupial fauna. Austral. Zoologist, Vol. 3, No. 7. 1926. Crucial evidence for antarctic radiation. Amer. Naturalist, Vol. 60. 1928. On the genus Stratiodrilus. Records Austral. Mus., Vol. 16. 1928a. Host and parasite. Presidential address. Proc. Linn. Soc. New South Wales. 1928, No. 1. “in press” in 1928. The composition and origins of the Australian fauna, with special reference to the Wegener hypothesis. Rep. Austral. Assoc. Adv. Sci., Perth, 1926. Hecner, R. W. 1928. The evolutionary significance of the protozoan parasites of monkeys and man. Quat. Rev. Biol., Vol. 3, No. 2, June. IHERING, H. von. 1891. On the ancient relations between New Zealand and South America. Trans. & Proc. New Zea. Inst., Vol. 24. 1902. Die Helminthen als Hilfsmittel der zoogeographischen Forschung. Zool. Anzeig., Vol. 26, Oct. 27. JouNsSTON, S. J. 1913. Trematode parasites and the relationships and distribu- tion of their hosts. Rep. Austral. Assoc. Adv. Sci., Melbourne. 1914. Australian trematodes and cestodes; a study in zoogeography. Med. Journ. Austral., Sept. 12. Abstract in Proc. Brit. Assoc. Adv. Sci., Australia. 1916. On the trematodes of Australian birds. Proc. Roy. Soc. N. S. Wales, Vol. 1. KELLOGG, VERNON L. 1896. New Mallophaga, I. Proc. Calif. Acad. Sci., Vol. 6. 1905. Insects. New York, Henry Holt. —— 1913. Distribution and species-forming of Ectoparasites. Amer. Nat- uralist, Vol. 47. 1913a. Ectoparasites of the monkeys, apes and man. Science, N.S., Vol. 38. 1914. Ectoparasites of mammals. Amer. Naturalist, Vol. 48, May, r1or4. Kettocc & Kuwana. Mallophaga from birds) Proc. Wash. Acad. Sci., Vol. 4, Sept. 30. LONGMAN. 1924. The zoogeography of marsupials. Mem. of Queensland Mus., Vol. 8, No. 1, Jan. 30. no. 8 PARASITES—METCALF 35 MattTHEw, W. D. 1915. Climate and evolution. Ann. N. Y. Acad. Sci., Vol. 24, Feb. 18. Metcatr, M. M. 1920. Upon an important method of studying problems of relationship and of geographical distribution. Proc. Nat. Acad. Sci., Vol. 6, No. 7, July. 1922. The host-parasite method of irvestigation and some problems to which it gives approach. Abstract in Anat. Record, Vol. 23, No. 1, January. 1922. Animal distribution and ancient distribution routes. Lectures upon evolution and animal distribution, Univ. Buffalo Studies, Vol. 2 a. No. 4, Dec. 1923. The opalinid ciliate infusorians. Bull. 120, U. S. Nat. Mus., June 15. 1923a. The origin and distribution of the Anura. Amer. Naturalist, Vol. 57. Sept.-Oct. 1923b. The host-parasite method and problems in which it is useful. Abstract in Anat. Record, Vol. 26, No. 5, Dec. 20. 1924. The opalinid parasites and the geographical distribution of the bell-toads (Discoglossidae), with a criticism of the age and area hy- pothesis. Abstract in Anat. Record, Vol. 29, No. 2, Dec. 28. 1926. Larval stages in a protozoon. Proc. Nat. Acad. Sci., Vol. 12, ING, 12, IDEe, 1928. The bell-toads and their opalinid parasites. Amer. Naturalist, Vol. 62, Jan.-Feb. j 1928a. Trends in evolution. A discussion of data bearing upon “ ortho- genesis.” Journ. Morphol. & Physiol., Vol. 45, No. 1, March 5. Moopir, R. L. 1923. Paleopathology. Univ. Chicago Press. Nose, G. K. 1922. The phylogeny of the Salientia. Bull. Amer. Mus. Nat. Hist.., Vol. 46. 1925. The evolution and dispersal of frogs. Amer. Naturalist, Vol. 50, May-June. Oriver, W. R. B. 1925. Biogeographical relations of the New Zealand region. Journ. Linn. Soc., Vol. 47, No. 313, Sept. 16. OrTMANN, A. E. 1806-1899. Tertiary invertebrates. Rep. Princeton Univ. Expedition to Patagonia. 190t. Theories of origin of Antarctic fauna and flora. Amer. Naturalist, Wol, 5: 1902. The geographical distribution of freshwater decapods and its bearing upon ancient geography. Proc. Amer. Philos. Soc., Vol. 41, No. 171, April-Dee. Rurrer, M. A. 1921. Paleopathology of Egypt. Univ. Chicago Press. ScHENCK, H. 1905. Ueber Flora und Vegetation von St. Paul und Neu Anister- dam. Wiss. Ergeb. Deutch. Tiefsee-Expedition, Vol. 2, T. 1, Lief 1. 1905a. Vergleichende Darstellung der Pflanzengeographie der sub- antarktischen Inseln inbesonders ueber Flora und Vegetation von Kerguelen. Same reference as Schenck (1905). 1907. Beitrage zur Kenntniss der Vegetation der Canarischen Inseln. Lief. 2, of same publication as Schenck (1905). SKOTTSBERG, CARL. 1915. Notes on the relations between the floras of subant- arctic America and New Zealand. Plant World, Vol. 18, No. 5, May. 36 SMITHSONIAN MISCELLANEOUS COLLECTIONS voL. 81 Warp, H. B. 1926. The needs and opportunities in parasitology. Science, Vol. 64, No. 1654. Sept. Io. Wenyon, C. M. 1926. Protozoology. London, Bailliere, Tindall & Cox. WuHueEeELer, W. M. 1927. The ants of Lord Howe Island and Norfolk Island. Proc. Am. Acad. Arts & Sci., Vol. 62, No. 4. May. Wituiams, H. U. 1909. The epidemic of the Indians of New England, 1616- 1620, with remarks on native American infections. Johns Hopkins Hospital Bull., Vol. XX, Nov. ZsSCHOKKE, F. 1899. Neue Studien an Cestoden aplacentaler Sangetiere. Zeitsch. f. w. Zool. Bd. 65, Heft 3, Feb. 14. 1907. Monezia diaphana n. sp. Ein weiterer Beitrag zur Kenntniss der Cestoden aplacentaler Sangetiere. Central bl. f. Bakt., Vol. 44. ‘ . _ SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLUME 81, NUMBER 9 A SECOND COLLECTION OF MAMMALS FROM CAVES NEAR ST. MICHEL, HATT (With TEN PLATEs) BY GERRIT S. MILLER, JR. Curator, Division of Mammals, U.S. National Museum (PUBLICATION 3012) Ra Le CITY OF WASHINGTON ~. PUBLISHED BY THE SMITHSONIAN INSTITUTION MARGH 30, 1929 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLUME 81, NUMBER 9 A SECOND COLLECTION OF MAMMALS FROM CAVES NEAR ST. MICHEL, HATTI (WitH TEN PLATES) BY GERRIT S. MILLER, JR. Curator, Division of Mammals, U.S. National Museum (PUBLICATION 3012) GITY OF WASHINGTON PUBLISHED BY THE SMITHSONIAN INSTITUTION MARGH 30, 1929 The Lord Gaftimore Press BALTIMORE, MD., U. 8. A. = A SECOND COLLECTION OF MAMMALS FROM CAVES NARS Si MICHEL ELA By GERRIT 'S. MILLER, TR: CURATOR, DIVISION OF MAMMALS, U. Ss. NATIONAL MUSEUM (WitTH 10 PLATES) Six years ago I published a short account of some bones of mammals from the floor material of two caves situated near St. Michel, north- central* Haiti (Smithsonian Misc. Coll., Vol. 74, No. 3, pp. 1-8, October 16, 1922). The small collection on which that paper was based had been made in 1921 by Mr. J. S. Brown and Mr. W. S. Burbank with the object of determining whether the caves contained deposits sufficiently rich in the remains of extinct vertebrates to justify a special expedition for their careful study. The few specimens brought home proved to be of so much interest that I visited Haiti in the spring of 1925, spending about four weeks at the plantation of l’Atalaye near St. Michel. A general account of this field-work ap- peared in Smithsonian Miscellaneous Collections, Vol. 78, No. 1, pp. 36-40, April 8, 1926. The following pages contain descriptions of the remains of mammals which I collected. Concerning the caves themselves there is nothing important to add to the notes made by Mr. Brown and Mr. Burbank. Two smaller caves were found close to the large cavern near the town of St. Michel. One of these has the entire roof fallen in so that very little of the original floor material could be investigated. The other was in good condition for working, and the deposits which it contained proved to be exceptionally rich. Locally the region in which this group of caves is situated is known as St. Francisque. The cave in the dry valley north of the Atalaye plantation had been completely worked out for guano since it was examined by Mr. Brown and Mr. Burbank. Thus the interesting bone deposit which it contained had been almost totally destroyed. Nevertheless I succeeded in finding a few speci- mens scattered among the sifted limestone fragments with which the 1Not “at the northwest end” of the republic as I wrongly stated in my general account of the region. SMITHSONIAN MISCELLANEOUS COLLECTIONS, VoL. 81, No. 9 SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL. 8&1 tN floor is now covered to a depth of nearly 1o feet. On the top of the ridge bounding the west side of this valley are situated at least three caves which had not been previously examined. One of these has no opening other than a hole in the roof about 6 x 10 feet in diameter. The chamber beneath this hole appeared to extend down- ward more than 20 feet. Its lateral extent could not be determined, and J made no attempt to explore it. One of the other caves is unusually deep, while the third is of more normal form, but rather narrow and crooked. In both I found abundant remains of extinct mammals. In all of these caves the deposits began at or near the surface and continued downward to a depth of about four feet. The bones then ceased, and further digging proved so fruitless that it was nowhere continued to rock bottom. Wherever bones occurred the deposit could be discovered in a few minutes’ work with shovel or trowel; and at any spot where the first few minutes’ digging revealed nothing the result of further excavation to a depth of six feet was fruitless. Mr. Brown and Mr. Burbank had previously found this to be the case. Before going to St. Michel I spent a day working in a cave at Diquini, near Port-au-Prince. The conditions there appeared to be exactly the same as in the large cave at St. Francisque, but no bones were found other than a few remains of domestic goat and pig in the most superficial layers, and recent bats and introduced rodents in fresh owl pellets. Why this cave should have been barren of the extinct fauna which occurs so abundantly in those near St. Michel is a question to which I cannot suggest an answer. Since this paper was written the St. Michel caves have been again visited in the interests of the National Museum. The generosity of Dr. W. L. Abbott enabled Mr. Arthur J. Poole, Scientific Aid, Divi- sion of Mammals, to spend the period from December 15, 1927, to March 15, 1928, in carrying on excavations which have probably re- sulted in exhausting the bone deposits. Inspection of the very rich material which he brought back to Washington shows that, in general, these new collections confirm the conclusions which I had reached after study of the specimens that I obtained myself. Such additional facts as they bring to light pertain chiefly to details concerning some of the new forms which I had already described in manuscript. I have therefore concluded to publish this paper as it was originally written, except for the account of the ground sloths, animals for whose understanding my material proves to have been so inadequate as to have led to conclusions which I now believe to have been wrong. NO. Q MAMMALS FROM CAVES IN HAITI—MILLER 3 INSECTIVORA Insectivores of the genus Nesophontes are abundantly represented in the Haitian caves. They have not previously been recorded from the island of Hispaniola. In the superficial layer of the cave floors the bones of these animals occur in undisturbed material along with remains of Epimys rattus and Mus musculus. This association is so intimate that there appears to be no reason to doubt the simultaneous occurrence of the insectivores and the introduced rodents. Some of the jaws of Nesophontes are more fresh in appearance than some of the jaws of Rattus near which they were found. Whether or not Nesophontes now exists alive is a question which for the present cannot be answered. No bones of insectivores have been found in any of the numerous fresh owl pellets which I have examined. It seems not improbable, however, that if any part of the island remains uninvaded by the roof rat, the native animal might now be found to exist there. It is a noteworthy fact that up to the present time no remains of Solenodon have been found in any of the caves. This animal is so much larger than Nesophontes that its absence from deposits which are mostly owl-made might at first be thought to be due to this circumstance. Its size, however, is no greater than that of several of the rodents which were freely eaten by the extinct giant barn owl, of whose refuse the bone deposits chiefly consist. While it is there- fore impossible to suggest any reasonable explanation of the absence of Solenodon bones, the fact of this absence is an important one because of its bearing on the question of the completeness of the faunal record preserved in the caves. NESOPHONTES PARAMICRUS sp. nov. Plater West Type.—Skull, lacking postero-inferior portion of occiput; the fol- lowing teeth in place: pm, pim*, m' and m? of right side, m* and m* of left side. No. 253063, U. S. Nat. Mus. Collected at front of large cave near St. Michel, Haiti, March, 1925, by Gerrit S. Miller, Jr. Diagnosis —Size and general characters of skull and teeth as in the Cuban Nesophontes micrus G. M. Allen. Upper molars without the well-defined sulcus which, in NV. micrus, lies between the base of metacone and posterior commissure of protocone ; lower molars with metaconid and entoconid obviously less nearly terete than in the Cuban animal. ————————— 4 SMITHSONIAN MISCELLANEOUS COLLECTIONS voL. 81 Skull—tThe skull appears to be similar to that of Nesophontes MICcrus. Teeth—As compared with those of Nesophontes micrus the larger maxillary teeth are more robust in general form, a character resulting from the less rapid narrowing of the base of the protocone toward the lingual side of the tooth crown. This peculiarity is especially evident in mt and m?, but it is also visible in pm*. This general tendency toward robustness of the cusps appears to be responsible for the main dental peculiarity by which the Haitian and Cuban members of the genus are distinguished from each other. In Nesophontes micrus there is always present, up to the time when this portion of the crown is destroyed by wear, a distinct and often wide notch at the point where the posterior margin of the protocone joins the base of the metacone. In N. paramicrus the bases of the two cusps are so large and well filled-out that they come together directly and smoothly or with at most a faintly developed intervening transverse crease. The same general features are present in the mandibular teeth, where the cusps show a uniform tendency to be heavier and less nearly terete than in the Cuban animal, characters best appreciated on comparison of the mataconid and entoconid of the two species. The heels of the lower molars are quadrangular (longer than broad) rather than squarish in outline and the bottoms of the central convexities tend to be rather broadly rounded instead of narrowly infundibuliform. Measurements——Type: greatest length, 32.4+; palatal length, 15.0; glenoid breadth, 12.4; interorbital breadth, 7.4; palatal breadth including molars, 9.2; front of canine to back of m*, 12.2; four molariform teeth (alveoli), 7.2. é Specimens examined. ——Skulls, 2; left maxilla, 1; mandibles, 18; humeri, 9; femora, 10; innominate, 1. Reimarks.—This species is sharply differentiated from the Porto Rican N. edithe by its much smaller size, and from the Cuban N. micrus by the peculiarities of its teeth. NESOPHONTES HYPOMICRUS sp. nov. Plate 1, fig. 2 Type—Nearly perfect skull (lacking auditory parts, incisors, canines and right median premolar) No. 253077, U. S. Nat. Mus. Collected in the deep cave near the Atalaye plantation, Haiti, March, 1925, by Gerrit:S.. Millers: NO. 9 MAMMALS FROM CAVES IN HAITI—MILLER 5 Characters —Like Nesophontes paramicrus but constantly smaller (see pl. 1 and detailed comparisons under “remarks’’) ; triangular outline of m1 and m? in palatal aspect narrower ; heels of mandibular molars shorter, their concavities narrowly funnel-shaped at base as in N. micrus. Skull—Except for its smaller size the skull appears to be essen- tially similar to that of N. paramicrus. Teeth._—The upper teeth in four individuals differ constantly from the two specimens of N. paramicrus in the narrower triangular outline of the crowns of mm! and m?. In the mandibular teeth the heel of each molar is shorter, though this character is usually more pronounced in mt, and 1M. than in Mz. Measurements—Type: greatest length, 27.6; condylobasal length, 26.8; palatal length, 12.8; glenoid breadth, 10.6; interorbital breadth, 5.8; palatal breadth including molars, 7.2; depth of braincase (median), 6.4; fronto-palatal depth behind molars, 5.2; front of canine to back of m*, 9.8; four molariform teeth (alveoli), 6.0. Specimens examined.—Skulls, 4; left maxilla, 1; mandibles, 24; humerus, 1; femora, 6; innominates, 3. Remarks.—That the original series of Nesophontes skulls from Porto Rico presents a range of variation in size which is unprec- edented among other dilambdodont insectivores is well known. This fact has been attributed to sexual dimorphism and as such has been made a part of the diagnosis of the family Nesophontide (see Anthony, Mem. Amer. Mus. Nat. Hist., n. s., Vol. 2, p. 365 “ june = October 12, 1918; Bull. Amer. Mus. Nat. Hist., Vol. 41, pp. 633, 635, December 30, 1919). The same conditions, though less well marked, were noticed by Anthony in a series of 33 skulls and 150 mandibles of the Cuban Nesophontes micrus (Bull. Amer. Mus. Nat. Hist., Vol. 41, p. 633, December 30, 1919). Through Mr. Anthony’s kindness I have had the opportunity to examine the entire series of Nesophontes jaws in the American Museum of Natural History, and as the result of this examination I am convinced that the differences in size shown by the Cuban and Porto Rican series are probably not due to the same causes as those which have produced the analogous differences that occur in the Haitian material. Among 26 jaws of the Porto Rican Nesophontes edithe in suffi- ciently good state of preservation to give the two most important measurements, namely, distance from articular process to anterior face of first molar, and depth through coronoid process, these vary from 16.2 to 22.2 mm. and from 9.0 to 13.2 mm. respectively. This 6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 81 is an unusually wide range of variation, but the steps by which it is accomplished are so small and the numbers of individuals are so evenly spaced along the series that the measurements present no features which suggest the inclusion of two species. The same is true of 48 jaws of the Cuban Nesophontes micrus. Here the range of variation in length from articular process to front of m, is from 12.2 to 14.6 mm. and that in coronoid depth is from 6.0 to 8.8 mm. One individual (teeth slightly worn) appears to be abnormally small, with the measurements I1.0 and 5.8 respectively ; but apart from this exception the variations are remarkably uniform, and again there is nothing to suggest anything else than purely individual variation. The series of 42 jaws from Haiti, in striking contrast, is readily separable into two lots on the basis of either one of three different measurements :* (a) distance from articular process to front of my, larger form (13 specimens), 13.2-14.0, smaller form (18 specimens ), 10.0-11.6; (b) depth through coronoid process, larger form (15 speci- mens), 7.6-8.8, smaller form (17 specimens), 6.0-7.0; (c) combined length of m, and mz, larger form (12 specimens), 4.50-4.85, smaller form (20 specimens), 3.70-3.85. The teeth in the smaller form are definitely reduced in size as compared with those of the larger indi- viduals, a character which is immediately appreciable on comparison of specimens. In the Cuban and Porto Rican series the teeth tend to remain more constant throughout the series. Therefore in those smaller Cuban jaws which approach in size the maximum of the smaller Haitian form the teeth are obviously larger than in the latter. Finally there is no difference in the structure of the heel in the teeth carried by the large and small Porto Rican or Cuban jaws, while in the Haitian specimens an obvious difference is present.” Turning now ‘to the skull and the maxillary dentition we find that the contrasts in size between the extremes of specimens from Cuba is about the same as that seen in those from Haiti. The teeth from Cuba, however, are alike in form from the largest to the smallest of 13 specimens, while in those from Haiti there is an obvious difference in the form of the triangular crown outline in the two largest as compared with five others. A final interpretation of these facts must await the ‘Owing to the fact that some of the mandibles are imperfect it is impossible to obtain all three measurements from every individual. * This is so constant that I made only one error in identifying, by this char- acter alone, 26 jaws (18 hypomicrus and 8 paramicrus) submitted to me one at a time by an assistant. The specimens were examined under a magnifying power with which I was unfamiliar, this having the effect of destroying all sense of relative size. NO. 9 MAMMALS FROM CAVES IN HAITI—MILLER i accumulation of much more abundant material; but it now appears obvious that the variation in Haitian Nesophontes is different in char- acter from that which is presented by the members of the genus occurring in Porto Rico and Cuba, and that the course which does least violence to probability may be followed by recognizing two species among the larger Haitian specimens, separated from each other by absolute differences in size and by easily appreciable structural characters of both maxillary and mandibular molar teeth, a condition which is not known to be due to sexual dimorphism in any insectivore. NESOPHONTES ZAMICRUS sp. nov. Plate 1, fig. 3 Type.—Anterior part of skull with complete palate (teeth lacking except pm’ left and the molariform teeth of both sides) No. 253090, * U.S. Nat. Mais. Collected in large cave near St. Michel, Haiti, March, 1925, by Gerrit S. Miller, Jr. Characters —Size much less than in any hitherto known member of the genus; palatal length, 10.6; four largest maxillary teeth, 5.0; four largest mandibular teeth, 5.6. Skull—Except for their smaller size the two imperfect skulls of this animal do not show any appreciable characters by which they can be distinguished from those of Nesophontes hypomicrus. The type gives the impression of greater slenderness, but this may be due to its small actual size. The ratio of palatal width to palatal length in the type is 54.7 and of palatal depth (at posterior margin) to palatal length is 37.7. In both of the two skulls of WM. hypomuicrus these ratios are 55.4 and 4o respectively, a difference which appears to be wathin the limits of reasonably looked-for individual variation. A greater difference is seen in the ratio of length from hamular process to depth including hamular process: 39.3 in N. gamicrus, 42.7 in N. hypomicrus. Still greater is that between the ratio of rostral width at level of canine to palatal length: 24.5 in N. zamicrus, 30.7 in N. paramicrus. Whether or not these peculiarities are anything more than individual is a question which must for the present remain open. Teeth.—The teeth, except for their smaller size, resemble those of Nesophontes hypomicrus in all the characters which distinguish this animal from N. paramicrus. Measurements—Type: palatal length, 11.0; glenoid breadth, 7.8; interorbital breadth, 5.0; palatal breadth including molars, 5.8; front of canine to back of m*, 8.2; four molariform teeth (alveoli), 5.0. 8 SMITHSONIAN MISCELLANEOUS COLLECTIONS voL. 81 Two mandibles: articular process to front of mm, 9.0 and 8.8; depth through coronoid process, 4.8 and 4.6; four molariform teeth (alveoli), 5.2 and 5.2. Specimens examined.—Anterior portion of skull, 1 (type) ; median portion of skull (rostrum broken away at level of pm*), 1; mandibles, 2; humerus, f. Remarks.—In their extremely small size the specimens which I refer to Nesophontes zamicrus are sharply set off from all the other material which I have examined. In the type and one mandible the teeth are just beginning to wear; in the second skull and second jaw the process is distinctly more advanced. The series of N. hypomicrus includes individuals representing exactly the same stages but showing no approach to the diminutive size of the smallest animal. CHIROPTERA Many bones of bats occur in the deposits. While some of these must have come from individuals which inhabited the caves and died there, most of them were probably dropped in owl pellets. The species are all, with one exception, known to be present inhabitants of the island. The one exception is a local form of a genus not hitherto found living elsewhere than in Cuba. There is no reason to suppose that it is extinct in Hispaniola. CHILONYCTERIS PARNELLII PUSILLUS G. M. Allen One skull from owl pellets in the cave at Diquini. MORMOOPS BLAINVILLII CINNAMOMEA (Gundlach) Three skulls from the larger cave near St. Michel. All in superficial deposit, one of them in a fresh owl pellet. MACROTUS WATERHOUSII WATERHOUSII Gray Three skulls and five mandibles from the large cave near St. Michel. One mandible from the small cave. All in superficial deposits. Four skulls from owl pellets in the cave at Diquini. MONOPHYLLUS CUBANUS FERREUS Miller A skull, lacking all the teeth except mm’ right and pm* and m+ leit, was found among the owl pellet material from the cave at Diquini. This specimen is unique among the many skulls of Monophyllus which I have examined in possessing the well-developed alveolus of a NO. 9 MAMMALS FROM CAVES IN HAITI—MILLER 9 simple premolar immediately behind the alveolus of the canine. The cavity is closely crowded between the alveolus of the canine and that of the anterior root of the normal anterior premolar. Its diameter is about .25 mm. In other respects the skull does not differ appreciably from those collected by Dr. W. L. Abbott at Jérémie. Measurements —Greatest length, 21.4; condylobasal length, 20.0; zygomatic breadth, 9.0; breadth of braincase, 8.8; postorbital con- striction, 4.0; breadth of rostrum across alveoli of canines, 3.8. BRACHYPHYLLA PUMILA Miller One skull from the steep cave near the Atalaye plantation. Its measurements are as follows: greatest length, 28.0; condylobasal length, 26.8 ; zygomatic breadth, 15.8 ; lacrimal breadth, 9.0 ; postorbital constriction, 6.2; breadth of braincase, 12.2; depth of braincase at middle, 9.6; mandible 19.0 ; maxillary toothrow (alveoli), 9.2; greatest width of palate including molars, 10.4; mandibular toothrow (alveol1), 10.2. This specimen and the two originally collected by Dr. W. L. Abbott near Port-de-Paix shows that the Haitian Brachyphylla is readily distinguishable from the large form inhabiting Porto Rico. From the small Cuban B. nana it appears to differ in slightly less reduced size. broader rostrum and palate, and larger molars. ARTIBEUS JAMAICENSIS JAMAICENSIS Leach Seven mandibles from the large cave near St. Michel, six skulls and nine mandibles from the deep cave near the Atalaye plantation and three skulls from owl pellets in the cave at Diquini. A large colony occupied the crooked cave in the group near the Atalaye plantation. When disturbed by the noise made by workmen digging in the cave floor the bats soon took refuge in small holes in the roof, where they remained almost completely hidden. On one occasion a half-grown young, unable to fly, fell from a roof cavity to the ground near where we were excavating. As it lay helpless it uttered chirping, bird-like cries. Immediately the air was filled with dozens of plunging and rising adult bats behaving in the manner of a flock of terns hovering over a wounded companion. Not one of them actually touched the young animal, and the confusion soon subsided, the adults retiring again to their holes. PHYLLOPS HAITIENSIS (J. A. Allen) Ten skulls, one left maxilla, 7 mandibles from the large cave near St. Michel; one skull from the deep cave and one mandible from the 10 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 81 crooked cave near the Atalaye plantation. One skull from owl pellets in the cave at Diquini. The skulls were found at all levels from the surface downward to a depth of about two feet. EROPHYLLA SANTACRISTOBALENSIS (Elliot) One skull and two mandibles from the large cave near St. Michel ; two skulls and three mandibles from the deep cave near the Atalaye plantation. The skuils exactly resemble three collected in a cave near Port-de-Paix by Dr. W. L. Abbott. In cranial characters this species resembles Erophylla bombifrons of Porto Rico and differs notably from the Cuban E. sesekorni and its relatives E. syops of Jamaica and E. planifrons of the Bahamas. The close correspondence in size between the skulls of E. santacristo- balensis and E. bomifrons is shown by the following measurements of the three best Haitian specimens (a) compared with those of three skulls from Porto Rico (b): greatest length, (a) 24.0, 23.4, 23.4, (b) 24.0, 24.2, 24.4; condylobasal length, (a) 22.2, 22.0, 22.0, (b) 22.2, 22:4, 22.4% breadth: of brainease, (a) 9:6, 10:0, 10:0, (b)) 10:0, 10.4, 10.0; postorbital constriction, (a) 4.6, 4.4, 4.6, (b) 4.4, 4.6, 4.6; breadth of rostrum at base of canines, (a) 5.0, 5.0, 4.8, (b) 5.0, 5.2, 5.0; median depth of braincase, (a) 8.4, 8.0, 8.0, (b) 8.0, 8.2, 8.2. Comparison of specimens shows that the rostrum in the Haitian animal is smaller relatively to the braincase than it is in Erophylla bombifrons, and further material will undoubtedly demonstrate the specific distinctness of the two animals. PHYLLONYCTERIS OBTUSA sp. nov. Type.—tImperfect skull No. 253095, U. S. Nat. Mus. Collected in the crooked cave near the Atalaye plantation, about four miles east of St. Michel, Haiti, March, 1925, by Gerrit S. Miller, Jr. Characters —Like the Cuban Phyllonycteris poeyt but incisive foramina smaller and anterior border of premaxillaries as viewed in palatine aspect less narrowly curved. Skull and teeth—The size of the skull is essentially as in Phyllonycteris poeyi, though the average may prove to be above that in the Cuban animal when it is possible to compare adequate series of specimens. The structure of the anterior part of the palate is alike in the three specimens examined, and is not duplicated by any among the large number of Cuban skulls with which I have compared them. Taking the width of the palate between the incisors and canines as NO. Q MAMMALS FROM CAVES IN HAITI—MILLER ro 100, the length of this region from front of premaxilla to posterior border of foramina averages about 82 in Phyllonycteris poeyi, while in the three specimens of P. obtusa it is only 56.6, 58, and 59.5, respectively. The curve of the anterior premaxillary border of the palate forms part of a circle which, if completed posteriorly, would pass close behind the foramina in P. poevi, but in P. obtusa it would be so much larger that the hinder edge of the foramina would scarcely extend beyond its center. The mandible and the molars, both maxillary and mandibular, do not differ appreciably from those of P. poeyr. Other teeth lost. Measurements—Type and specimen from Diquini (No. 253006) : greatest length, —, 22.2; palatal length, 10.0, 10.2; back of glenoid process to front of premaxillary, 17.2, 16.8; breadth of braincase, —, 10.2; postorbital constriction, 5.6, 5.4, width of palate including molars, 7.2, 7.0; mandible, —, 15.4; maxillary toothrow (alveol1), 7.0, 7.2; mandibular toothrow (alveoli), —, 8.0. Specimens examined.—A skull and mandible from the crooked cave near the Atalaye plantation, a skull from a cave near Port-de-Paix (Dr. W. L. Abbott), and a skull from owl pellets found in the cave at Diquini. Remarks.— Unlike its relative Erophylla the Haitian Phyllonycteris is not particularly like the Porto Rican member of its genus. As Anthony figures (Mem. Amer. Mus. Nat. Hist., n.s., Vol. 2, pl. 60. fig. 12) and describes the Porto Rican P. major it is a larger animal with relatively small teeth ; palatal length ranging from 10.6 to IT.1, but with a toothrow of only 6.7 to 6.8. EPTESICUS HISPANIOL Miller TADARIDA MURINA (Gray) One mandible of each of these small bats was collected in the large cave near St. Michel. RODENTIA 3ones of native rodents representing six genera, only one of which is known to have a living species, form the great mass—probably more than 95 per cent—of all the deposits. Mingled with them are the remains of the large owl, Tyto ostologa, which brought them to the caves. It is easy to realize that the existence of a bird of this type might depend so entirely on an abundant rodent food supply that, with the gradual disappearance of the large indigenous rodents, the owl must also have become extinct, leaving the caves to the small 12 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 81 Tyto glaucops capable of subsisting on introduced rats and mice, and on the native bats, lizards and small birds. Beneath a ledge in one of the caves near St. Michel I found pellets of the small owl on the surface, and, at a depth of from eighteen inches to two feet, compactly moulded masses of extinct rodent bones, evidently parts of the pellets of the extinct bird which once used this same resting place. BROTOMYS VORATUS Miller Plate 2, fig. 1 Two imperfect skulls and 52 mandibles. These specimens represent all the caves worked in with the exception of the deep cave near the Atalaye plantation. The skulls essentially agree with the type, from the Dominican Republic. The mandibles, when compared with specimens of Boromys offela and B. torrei collected in Cuba by William Palmer in 1917 show no striking peculiarities. In both species of Boromys, however, the masseter ridge on the outer side of the mandible is so developed that, in the region beneath m2, its upper surface projects almost at a right angle to the outer surface of the mandible above it, while its extreme edge in some specimens is slightly turned upward. In Brotomys the upper surface of the ridge slopes obliquely downward and the margin is not upturned. The three genera Brotomys, the Cuban Boromys, and the Porto Rican Heteropsomys are at once distinguishable from the other native Antillean rodents by their relatively low crowned, long rooted, subterete cheekteeth. All three are intimately related and it may eventually be found expedient to unite them under one name. For the present, however, it seems preferable to regard them as distinct from each other. The additional material now at hand makes it possible to define their differences as follows: Posterior termination of incisor root visible behind anterior base of zygoma when skull is viewed from below; antorbital foramen rela- tively small, its height much less than length of toothrow...... Hetcropsomys. Posterior termination of incisor root not visible behind anterior base of zygoma when skull is viewed from below; antorbital foramen relatively large, its height nearly equal to length of toothrow. A deep neural channel on floor of antorbital foramen; posterior termination of incisor root marked by an obvious swelling... .Boromys. No definite neural channel on floor of antorbital foramen; pos- terior termination of incisor root not marked by an obvious Swelling i. ccoko cea ndeds oho en ean eee Cerca Brotomys. NO. 9 MAMMALS FROM CAVES IN HAITI—MILLER I tn BROTOMYS (?) CONTRACTUS sp. nov. Plate 2, fig. 2 Type—Anterior portion of skull, lacking zygomata, nasals and teeth, No. 253100, U. S. Nat. Mus. Collected in small cave near St. Michel, Haiti, March, 1925, by Gerrit S. Miller, Jr. Characters ——Resembling Brotomys voratus, but size slightly less, rostrum relatively shorter, interorbital region narrower in proportion to frontopalatal depth and more arched transversely ; teeth broader than in Brotomys voratus, and palate noticeably constricted, its inter- alveolar width at middle conspicuously less than transverse diameter of the adjoining alveoli. Skull—While resembling in a general way that of Brotomys voratus the skull of B. (?) contractus, even in the imperfect condi- tion of the only known specimen, shows well marked differential characters. Most conspicuous among these is the narrowness of the bony palate as compared with the very wide alveoli of the anterior cheekteeth. In three specimens of B. voratus (the type from the Dominican Republic and two from Haiti) the width of the palate between the alveoli of the second cheekteeth is 2.55, 3.0 and 3.0, respectively, and the width of the first alveolus is 2.25, 2.25 and 2.30. In the type of B. (?) contractus the width of the palate at the same level is only 1.65, while that of the first alveolus is 3.60. The narrowing of the skull shown by the palate is also evident when the interor- bital breadth is compared with the fronto-palatal depth. In the type of Brotomys (?) contractus the ratio of this breadth (15.6) to depth (13.0) is only 83.3, while in the three specimens of B. voratus it is 92.5, 90.0 and 92.6. The greater transverse convexity of the interor- bital roof is a character which cannot be expressed by measurements ; it is immediately obvious when specimens are compared in posterior view. Because of the imperfect condition of the skull a comparison of the length of the rostrum with anything but the length of the palate is difficult; hence the apparent shortening of the rostrum may be due in part to an actual lengthening of the palate to accommodate the enlarged teeth. In Brotomys (°?) contractus the length of the palate (9.4) measured from posterior border to level of anterior margin of alveolus of pm* is essentially equal to the distance from the latter level to alveolus of incisor (9.8) ; in B. (?) contractius it is barely more than the distance from alveolar level to front of incisive foramina (that is, about 5 mm. less than the distance to alveolus of incisor). The alveolar length of the toothrow in the type of B. (?) contractus cannot be exactly measured (the alveolus of 7° is entirely | | { ' a} q { 4 (" y i , > es 14 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 8&1 missing on one side and is incomplete on the other) but it must have been essentially equal to the diastema (10.8 mm.). In the three skulls of B. voratus it is 9.8, 10.0 and 9.6, while the diastema in the same specimens is 13.6, 12.6 and 12.8, respectively. Specimens examined.—One, the type. Remarks.—The disproportion between the alveoli and palate in this species as compared with Brotomys voratus is so great as to suggest that, when more completely known, the animal will prove to represent a distinct genus. In all of the other related members of the group from the large Heteropsomys (and Homopsomys if distinct) of Porto Rico to the small Boromys torret of Cuba the proportionate width of palate and alveoli does not greatly vary; the palate, at the m' level is always at least equal to the width of the largest alveolus. The narrowing of the palate to less than half the width of this alveolus in B. (?) contractus may therefore easily be a character of more than specific weight. ISOLOBODON LEVIR (Miller) Plate 2, figs. 3, 3a 1922 Isolobodon portoricensis Miller, Smithsonian Misc. Coll., Vol. 74, No. 3. p. 3. October 16, 1922. 1922 Ithydontia levir Miller, Smithsonian Misc. Coll., Vol. 74, No. 3, p. 5. October 16, 1922. Thirty palates and fragmentary skulls, more than 600 mandibles. This is the most abundantly represented of the vertebrates found in the bone bearing deposits. Its flesh seems to have been the chief article of food of the giant barn owl, Tyto ostolaga; many of the skulls and jaws were found in masses of bones which had the structure characteristic of owl pellets. The original collection from the large cave near St. Michel included two isolated upper premolars of /solobodon. Wrongly determining them as lower teeth I made these specimens the basis of a new genus and species, /thydontia levir, selecting as type what I supposed to be “a right mandibular tooth, probably pm, or m,,” but actually, as the rich material now at hand clearly shows, pm* left. So far as the generic name /thydontia is concerned there can be no doubt—it is a synonym of /solobodon. But the proper disposition of the specific name is less easily determined. For the present it seems necessary to retain [solobodon levir as the designation of the Haitian member of the genus. Although the absence of good skulls from the St. Michel series makes a satisfactory comparison with /solobodon portoricensts NO. Q MAMMALS FROM CAVES IN HAITI MILLER 15 impossible, the smaller size of the Haitian specimens is so constant as compared with material from Porto Rico and the Virgin Islands that the existence of two members of the genus appears to be established. The circumstance must not be overlooked that the Haitian food refuse was accumulated by owls, while that formed elsewhere was chiefly if not entirely deposited by men. It is possible therefore that the difference in size may be partly due to selection of the rodents used as food—the owls tending to capture smaller, more easily devoured individuals, the men preferring the larger ones. That the owls were able to eat animals as large as the largest Porto Rican [solobodon is shown by the frequent presence in the deposits of Aphetreus jaws of equally large size. Whatever bearing the possibility of selection may have, the facts are as follows: Among more than 600 Haitian mandibles the eight largest have toothrows of the following lengths, 16.2, 16.2, 16.2, 16.4, 16.6, 17.0, 17.2 and 17.2 mm., while the extremes of Anthony’s measurements of individuals selected from a series of 200 Porto Rican specimens are 17.6 and 19.2 mm. The three longest maxillary toothrows among the Haitian specimens selected for large size are 16.2, 16.2 and 17.0 mm. ; Anthony gives 17.2 to 19.3 mm. as the range of variation among adults in his series of 17 skulls. The interorbital breadth can be measured accurately or approximately in seven of the Haitian skulls. It ranges from 15 mm. to about 18 mm.; Anthony’s extremes are 19.8 and 23.5 mm. in six skulls from Porto Rico. In two Haitian specimens the length of the frontal bone along the median suture is 18.6 mm. and 20.0 mm.; the extremes of eight from Porto Rico are given as 22 mm. and 30 mm., with only three specimens less than 24.5 mm. The breadth of rostrum at premaxillary suture does not exceed II mm. in any of 15 Haitian specimens (some of them ob- viously immature), while in seven from Porto Rico it ranges from 13 mm. to 14.5 mm. Under these circumstances it seems necessary to recognize the Haitian /solobodon as a distinct form. The status of the /solobodon whose bones have been found in kitchen middens in the Dominican Republic is a matter of special interest now that it becomes impossible to regard the Haitian member of the genus as identical with J. portoricensis. I once said that there appears to be no way to distinguish between Dominican, Porto Rican and Virgin Island specimens ;* and after going over the ground again in the present connection I am of the same opinion. A palate from San Pedro de Macoris, Dominican Republic, is broken in such a Proc. U. S. Nat. Mus., Vol. 54, p. 508, October 15, 1918. 2 16 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 81 manner that the toothrow cannot be measured, but the alveolar length must have been at least 18 mm.; enough of the base of the rostrum is preserved to show that the breadth at premaxillary suture was more than 13 mm. In three mandibles from the same locality the toothrow measures 17.6, 18.6 and 18.6 in contrast to the maximum of 17.2 for the entire series of over 600 jaws from the Haitian caves. Of two mandibles collected by Gabb at San Lorenzo Bay one has a toothrow 18.8 mm. in length, while in the other, an obviously younger individual, it is 16.8, only a little below the maximum for the Haitian specimens. APHZETREUS MONTANUS Miller Plate 2, figs. 4, 4a, 4b Seventeen imperfect skulls and palates, 208 jaws. In both groups of caves the remains of this animal were common, the frequency of their occurrence coming next after that of Jsolobodon levir. The material at hand makes it possible to define the genus more completely than I was able to do in the original paper. It is now evident that the genera Aphetreus, Isolobodon and Plagiodontia form a rather compact group, the members of which are more nearly related to each other than any one of them is to Capromys and its allies. In all three the enamel pattern of the upper molars is tetramerous ; in Plagiodontia the upper premolar has reached the same stage of simplifi- cation, but in Aphetreus and Isolobodon this tooth retains a small fifth element. The maxillary teeth of Capromvs and Geocapromys are all pentamerous. In the /solobodon group the direction of the inner reentrant fold is diagonally forward in the upper teeth, backward in the lower teeth; the reverse is the case in Capromys. The general structure of the crowns in the Capromys group parallels that which has been developed by the voles; this is not true with regard to !solobodon and its allies. The characters of the three genera may be tabulated as follows: Curve of upper incisor short, the root of the tooth lying at anterior margin of zygomatic process of maxillary; lower incisor terminating beneath m1; pm* with one outer reentrant angle, its enamel pattern exactly similar to that of the molars; reentrant folds in upper teeth very oblique, their slant 45° or less as referred to corresponding alveolar line; reentrant folds on inner side of the lower teeth extend- ing less than halfway across crowns; frontal sinus sufficiently inflated to produce an obvious swelling over anterior zygomatic root, to en- croach on area of antorbital foramen, and to a less degree on that of orbit; posterior margin of zygomatic process of maxillary lying about in’) lines wathwanterior alveolar i bondenmnsc. eerie anion eee Plagiodontia. NO. Q MAMMALS FROM CAVES IN HAITI—MILLER 7 Curve of upper incisor long, the root of the tooth lying in antorbital foramen: lower incisor terminating beneath ms; pm‘ with two outer reentrant angles, its enamel pattern obviously different from that of the molars; reentrant folds in upper teeth not very oblique, their slant more than 45° as referred to corresponding alveolar line; reentrant folds on inner side of lower teeth extending more than halfway across crowns: frontal sinus not sufficiently inflated to produce an obvious swelling over anterior zygomatic root or to encroach on area of ant- orbital foramen or of orbit; posterior margin of zygomatic process of maxillary lying at or behind level of middle of alveolus of pm’. Opposed inner and outer reentrant angles of all teeth remaining distinct throughout life, the enamel pattern of each tooth entire; crowns and alveoli of both upper and lower molars Hlearly as*lone as wide... 2. .... 22 cc ee ee eee nen sens Tsolobodon. Opposed inner and outer reentrant angles of all teeth becoming confluent in adults, the enamel pattern of each tooth then di- vided into two sections; crowns and alveoli of both upper and lower molars conspicuously wider than long.....--...--+++ A phetreus. The series of mandibles includes about 30 specimens in which the breaking through of the opposed enamel folds has not yet taken place. Unfortunately there are no sets of upper teeth representing the same stage. In these immature individuals the enamel pattern of the mandib- ular teeth contains exactly the same elements that are present in the corresponding teeth of /solobodon. The characteristic peculiarities of crown outline are, however, evident at a very early stage, and, though less pronounced than in the adults, they are sufficient to be diagnostic. In harmony with the shorter tooth crowns of Aphetreus the enamel folds are narrow as compared with those of /solobodon, and the re- entrants are more completely filled with cement. The crowns conse- quently tend to have a solid, squarish aspect, while in /solobodon they are oblong and always with conspicuous angular emarginations. From the mandibular teeth of Plagiodontia those of Aphetreus are readily distinguished by the less oblique direction of all the enamel folds, and by the greater length of the outer reentrant, which fold invariably extends more than halfway across the crown, while in Plagiodontia it never reaches the middle of the crown. The maxillary teeth have not hitherto been known. Like the man- dibular teeth they contain the same elements that are present in Isolobodon, but these elements are compressed in the axis of the toothrow, and the opposed reentrant folds are confluent in adults, thus splitting the enamel pattern into two sections. The region of breaking through in the maxillary teeth is clearly indicated by irregularities in the enamel outline ; hence it seems probable that in young individuals it will be found that the pattern is not split. 18 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 81 Two toothless mandibles, not improbably pertaining to one indi- vidual, dug from the small available area of original floor material in the caved-in chamber near St. Michel, are unique, among the octodont rodents which I have examined, in the presence of a well developed fifth alveolus behind the normal fourth (pl. 2, fig. 4b). PLAGIODONTIA ADIUM F. Cuvier Seven mandibles (five from the group of caves near St. Michel, the others from the crooked cave near I’Atalaye) are referable to the species represented by the large specimen from San Pedro de Macoris, Dominican Republic, which I have identified (Proc. U. S. Nat. Mus., Vol. 72,.Art. 16, pp. 5-6, September 30, 1927) as an individual of the species originally described by F. Cuvier. Only one of the Haitian specimens is fully adult, and in this the coronoid and angular regions are broken off and all the teeth have been lost. Its size must have been almost exactly the same as that of the Macoris jaw. In each the length of the symphysis menti is 27.6 mm. and the distance from the posterior angle of the symphysis to anterior margin of alveolus of pmy is 20.4. Among 13 jaws of the recently described Dominican Plagiodontia hyleum Miller the maxima for these two measurements are only 25.4 and 19.0, while the usual dimensions in adults are decidedly less, about 24 and 17. The length of the toothrow in the adult Haitian P. edium, 23.4, is only 0.6 mm. less than that in the Macoris specimen; the maximum in the series of P. hyleum is 20.6. In two of the younger Haitian individuals, both of them broken off immediately behind the toothrow, the second molar is not yet fully in place. They are, how- ever, distinctly larger and more robust than in two jaws of immature Dominican P. hyleum, one with m, worn flat but m, not in place, the other with all the crowns worn flat. In the five Haitian specimens with teeth the enamel pattern presents the characters which distinguish Plagiodontia edium from P. hyleum (see Miller, Proc. U. S. Nat. Mus., Vol. 72, art. 16, p. 4, and pl. 1, figs. 1c and 2, September 30, 1Q27): PLAGIODONTIA SPELZEUM sp. nov. Type.—Right mandible of young adult, No. 253160, U. S. Nat. Mus. Collected in the crooked cave near the Atalaye plantation, Haiti, March, 1925, by Gerrit S. Miller, Jr. Characters.—Resembling Plagiodontia hyleum Miller from eastern Dominican Republic but noticeably smaller ; length of mandible mea- sured from articular process probably not much exceeding 40 mm. NO. 9 MAMMALS FROM CAVES IN HAITI—MILLER 19g instead of ranging from about 48 to 54 mm.; mandibular toothrow usually less than 18 mm. instead of ranging from about 18.5 to 20.5mm. Portion of mandible in front of cheekteeth relatively shorter and more abruptly curved than in P. hyleum. Measurements——From five jaws which may be regarded as adult I am able to obtain the following measurements: length of mandible from articular process, 39.6, 39+, —, —, —, length of symphysis, 1o.0,.1O-=; 1725, 7-0, ——+ diastema, 9.0, 9:4, O25, 8:8,,9.2;: depth from alveolar margin to lowermost point of symphysis, 11.2, 11.2, 11.2, 10.8, 11.6; mandibular toothrow (alveoli), 16.2, 16.0, 15.8, 16.0, 15.6; transverse diameter of m, (grinding surface), 4.5, 4.5, 4.5, 4.2, 4.4. The same measurements in a mandible of P. hyleum which appears to be of exactly corresponding age (No. 239886) : length from articu- lar process, 48 ; symphysis, 21.6; diastema, 11.4; depth 13.0; toothrow, 18.6; width of 4, 5.3. Specimens examined.—Fifteen mandibles, all imperfect. Four of these came from the group of caves near St. Michel, the others were found in the crooked cave near the Atalaye plantation. Remarks.—The small Plagiodontia from the St. Michel caves differs conspicuously from the associated large P. @dium in size and in the longitudinally compressed cheekteeth. Its affinities are obviously with P. hyleum of the Samana Bay region, the only member of the genus known to be now living. At first sight the jaws of Plagiodontia speleum might be mistaken for immature specimens of P. hyleum, but when comparison is made between individuals in corresponding stages of development (as indicated in immature individuals by the eruption of the second and third molars, and in young adults by the gradual disappearing of porousness and surface wrinkling of the bone on the lower side of the jaw beneath the roots of these teeth) the differences between the two species become obvious. HEXOLOBODON gen. nov. Plate 3) figs, 1, ta) 1b Type—He.xolobodon phenax sp. nov. Characters.—So far as known most like Geocapromys, but differing as follows: cheekteeth with roots becoming closed at or soon after the stage when the crowns are worn flat ; root of lower incisor passing beneath root of m, and terminating, in fully adult individuals, on outer side of toothrow beneath the floor of the groove which separates the alveolus of m, from the base of the coronoid process; pm, (pl. 3, fig. Ia) with only two reentrant angles on inner side (as in Capromys) ; 20 SMITHSONIAN MISCELLANEOUS COLLECTIONS voL. 81 all of the maxillary teeth with two about equally developed reentrant angles on each side, these imparting to the crowns an evenly six-lobed structure (pli+3: fies 1): Remarks.—In the general structure of the palate and the relation- ship of the incisor roots to those of the premolars this genus is prac- tically identical with Geocapromys. The roots of the premolars come close together in the median line, where they are overgrown by the maxillary exactly as in Geocapromys. The roots of the premolars with their covering of bone fill up the lower part of the narial channel in the region between the incisor roots (pl. 3, fig, 1b). A broken palate without teeth could be distinguished by this character alone from a similar fragment of a Capromys or Plagiodontia skull, in both of which the anterior part of the narial channel is widely open between the roots of the premolars (pl. 3, fig. 2), but might be confused with a similar fragment pertaining to a member of the genera Geocapromys, Tsolobodon, or Aphetreus. In Geocapromys and Capromys (pl. 3, fig. 2) the roots of all four cheekteeth, when exposed by cutting or breaking away their bony covering, are seen to be about evenly spaced in the toothrow—at most the septum between the roots of pin* and m' is slightly thicker than the septa between the molars. In He.volobodon, on the contrary (pl. 3. fig. 1b), the root of the premolar is thrown conspicuously forward away from that of the first molar. The less specialized condition of the roots of the cheekteeth and the extension of the lower incisor root to the outer side of the mandibular toothrow are characters which, like the enamel pattern of the upper teeth, sharply differentiate this genus from its Antillean relatives Capromys, Geocapromys, Plagiodontia, Aphetreus, and [solobodon. HEXOLOBODON PHENAX sp. nov.. Plate 3, figs. I, ta, 1b Type.—Palate with complete dentition of immature individual (im; with only anterior half of crown worn flat), No. 253118, U. S. Nat. Mus. Collected in the small cave near St. Michel, March, 1925, by Gerrit’S;, Miller, Jr: Characters —An animal about the size of Capromys pilorides, but skull probably differing from that of all species of Capromys and Geocapromys in shorter rostrum and generally more robust form. With regard to features which are not obviously generic, such exact comparisons with Capromys pilorides as the fragmentary remains of NO. 9 MAMMALS FROM CAVES IN HAITI—MILLER 21 the extinct animal will permit, are as follows: palate in region between pm* and maxillo-premaxillary suture much smaller relatively to grind- ing area of toothrow (about 10 x14 mm. as compared with 13 x18 mm. in a specimen of C. pilorides with grinding area of toothrow of essen- tially the same length and breadth as that of the type), its upward slope more abrupt ; no obvious pit for attachment of the maxillo-naso- labialis muscle in region between pi, and incisive foramen (these pits are visible in all the living species of Capromys and Geecapromys ; they are not developed in /solobodon, Aphetreus or Plagiodontia) ; posterior emargination of palate extending forward slightly beyond level of posterior border of m? instead of about to middle of m?; narrow inferior maxillary zygomatic root, its width through middle of specialized muscle-insertion area considerably less than width of grinding surface of molars instead of distinctly greater than width of this surface. The upper toothrows are more convergent than in Capromys pilorides, so that the bony palate becomes reduced anteriorly to a width only about one-fifth that of the adjoining alveolus or of its own width posteriorly. In C. pilorides the anterior width of palate is considerably more than half that of alveolus and almost exactly half of its own posterior width. Posterior emargination of palate extend- ing slightly beyond level of septum between alveoli of m*® and m?. All of the mandibles are broken immediately behind the toothrows. In the portion which remains there are several obvious peculiarities as compared with the corresponding region in Capromys pilorides. The diastema is short and more abruptly concave when viewed from the side. The symphysis is conspicuously shorter than in C. pilorides and its long axis is set at a higher angle to the plane of the grinding surface of the molars; about 50° instead of about 35°. The anterior portion of the ridge which extends forward along the outer side of the mandible from the angular process is heavier and more evenly rounded than in the Cuban animal. The enamel pattern of the mandibular teeth appears to be not positively distinguishable from that of Capromys pilorides. Measurements.—Type: distance from posterior surface of m° to anterior border of maxillary directly in front of toothrow, 30.0 (35.0) ;* distance from posterior margin of incisive foramen to poste- rior margin of palate, 24.6 (26.2) ; distance from alveolus of pm* to anterior edge of maxillary, 9.4 (13.2) ; width of bony palate through * Measurements in parenthesis are those of a similarly broken palate of a slightly older individual of Capromys pilorides from Sierra La Guira, Pinar del Rio, Cuba (No. 253232, U. S. Nat. Mus.). SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL. 81 to bo anterior edge of posterior emargination, 5.6 (8.2); least width of palate between toothrows, 1.2 (4.0) ; maxillary toothrow (alveoli) 22.0 (22.4) ; alveolar width of pm*, 5.8 (5.2) ; height of m* from grinding surface to root, 15.0 (14.0). Mandible of an individual with teeth in same stage of wear as those of type: distance in alveolar line from posterior margin of mz to anterior margin of incisor, 36.0; distance from tip of incisor to posterior edge of grinding surface of m;, 38.0; diastema, 10.0; distance from tip of incisor to anterior margin of crown of pm,, 15.8; depth from inner margin of alveolus of pi, to posterior point of symphysis, 14.4; length of symphysis, 22.4; length of toothrow, grinding surface, 22.2, alveoli, 24.0; alveolar width of pm, 5.8. Mandible of an individual with crown of mz entirely worn flat: distance in alveolar line from posterior margin of m, to anterior margin of incisor, 37+ (46.6) ;* diastema, 11+ (18.8) ; depth from inner margin of alveolus of pm, to posterior point of symphysis, 17+ (19.0) ; length of symphysis, 23+ (23.0) ; length of toothrow, grind- ing surface, 24.2 (22.0) ; alveoli, 25.0 (22.4) ; alveolar width of pig, 5-4 (5.0). Specimens examined.—One palate, six mandibles and four isolated cheekteeth. A mandible and two of the isolated teeth were found in the caves near l’Atalaye, the rest of the material came from the large and small caves near St. Michel. QUEMISIA gen. nov. Plate 4, figs. 2, 2a Type.—Quemisia gravis sp. nov. Characters—Size and general features probably as in the Porto Rican Elasmodontomys. Enamel pattern of mandibular cheekteeth (pl. 4, fig. 2a) like that of Elasmodontomys (pl. 4, fig. 1a) but reentrant folds less transverse to the axis of the toothrow, the axis of the folds slanting forward at an angle of only 21° instead of about 50°. Mandibular symphysis extending backward beyond level of middle of m, instead of barely to middle of pm,; shaft of lower incisor not extending behind symphysis, its base lying beneath anterior half of m, (in Elasmodontomys the shaft of the incisor extends far beyond the symphysis to terminate beneath middle of m,); shaft of femur more flattened than in Elasmodontomys. * Measurements in parenthesis are those of an adult Capromys pilorides (No. 143150). NO. Q MAMMALS FROM CAVES IN HAITI—MILLER 23 Remarks.—The genus Quemisia is a member of the group which is represented by Elasmodontomys in Porto Rico and Amblyrhiza in Anguilla. The cheekteeth in all three of these genera are very hypso- dont but not ever-growing. The enamel pattern is pentamerous with the inner reentrant fold of the upper teeth (in Amblyrhiza and Elasmodontomys, at least) and the outer fold of the lower teeth passing behind the posterior outer reentrant. All of the reentrant folds penetrate nearly or quite across to the opposite side of the crown, thus producing a grinding surface which consists of a series of essen- tially parallel transverse enamel ridges. The most striking known peculiarities of Quemusia are the long mandibular symphysis, short lower incisor, and the very unusual for- wardly-directed enamel folds in the lower teeth. I have chosen the name because of my belief that the animal is probably the “ Quemi ” of Oviedo (Hist. Gen. et Nat. de las Indias, Madrid, 1851, p. 389). QUEMISIA GRAVIS sp. nov. Plate 4, figs. 2, 2a Type——Mandible of immature individual (7m, with crown not yet in place), No. 253175, U. S. Nat. Mus. Collected in the crooked cave near the Atalaye plantation, March, 1925, by Gerrit S. Miller, Jr. Characters——As compared with a mandible of Elasmodontomys obliquus in corresponding stage of tooth growth the type specimen of Quemisia gravis shows many peculiarities in addition to those which have already been described. The depth of the horizontal ramus at middle of m, is greater in proportion to the length of the toothrow (21.5: 33 instead of 18:34); the maximum width through the symphysis is greater (17.5 instead of 11) a difference occasioned partly but not wholly by the more posterior point of termination of the symphysis in the Haitian animal. The anterior base of the angular process is laterally compressed in Quemisia, so that it forms about one-third of the transverse diameter of the mandible; in Elasmodon- tomys it is so thick that it forms considerably more than half of the entire transverse diameter. The roots of the third and fourth cheek- teeth extend down into this thickened area in Elasmodontomys. In Quemisia the roots of the three molars form a broadly curved ridge extending backward and upward from the symphysis and separated from the base of the angular process by a shallow groove; this ridge has, at first sight, something the appearance of the ridge which marks the course of the incisor root in Elasmodontomys. 24 SMITHSONIAN MISCELLANEOUS COLLECTIONS voL. 81 The cheekteeth are open at the base, as in Elasmodontomys of the same age; whether or not they eventually become closed as in adult Elasmodontomys cannot now be determined. The enamel pattern is fundamentally the same as in Elasmodontomys, that is, a pentamerous pattern in which all the reentrant folds have been extended nearly or quite across the crown (the outer fold passing behind the second inner fold). The posterior limb of each fold has been thickened to form a conspicuous enamel plate and the anterior limb of each fold except the first has been reduced to the vanishing stage. As compared with that of the Porto Rican animal the pattern in Quemisia shows a mixture of excessive peculiarity and less high specialization. The forward turning of the enamel folds so that the anterior portion of each fold is approximately parallel with the main axis of the toothrow is a specialization of high degree and very peculiar kind. In Elasmodontomys there is an indication of this tendency at the front of the premolar, but the direction of the folds in the molars is normal and not essentially different from that seen in Plagiodontia, Isolobodun or Aphetreus. On the other hand the process of plate specialization has not progressed so far in Quemisia as it has in Elasmodontomys. While the external reentrant fold has extended completely across the crown in all three of the used cheekteeth neither of the two internal folds has quite reached the enamel of the opposite side in pin, and only the first has penetrated so deeply in m, and mz. In each of the molars there is, therefore, one incomplete enamel plate, the second, while in the premolar there are two, the first and second. In Elasmodontomys all the folds have crossed the crown in all the teeth, and there are, consequently, no incomplete plates. The peculiar twist- ing of the enamel pattern almost into the axis of the toothrow in OQuemisia throws the anterior loop of each tooth over on to the inner side of the crown out of contact with the tooth in front of it. The free face of each of these loops carries a fully developed enamel wall. In Elasmodontomys such an enamel wall occurs on the first loop of the premolar only. A fragment of incisor (apparently an upper tooth) 19 mm. in length has a width of 5 mm. and an antero-posterior diameter of 4.2 mm. at level immediately proximal to the terminal worn area. The anterior face is longitudinally fluted by six obscurely developed ridges and the faint intervening concavities. A broken femur which I refer without much doubt to this species differs from the corresponding bone in Elasmodontomys obliquus in the conspicuous flattening of its shaft. The greatest and least diameters NO. Q MAMMALS FROM CAVES IN HAITI—MILLER 25 of the shaft in its narrowest region are 12.2 and 8.2, while in one specimen from Porto Rico they are 10.8 and 8.8, and in another 13.0 and 9.8. Specimens examined. crooked cave near the Atalaye plantation; broken femur from the Mandible and piece of an incisor from the small cave of the same group. XENARTHRA The occurrence of ground sloths in Hispaniola was not known be- fore the discovery of a few bones in the St. Michel caves by Mr. Brown and Mr. Burbank. On the basis of this scanty material—four vertebre, three of them imperfect, a piece of a limb bone of a young animal, and a fragment of a rib—I was unable to refer the species to any genus, and, at Doctor Matthew’s suggestion, | recorded it’ as Megalocnus ? sp. ? On visiting the caves myself I secured teeth and a femur resembling the corresponding parts of the Porto Rican Acratocnus and also a calcaneum so unlike that of Acratocnus as to suggest the existence of two sloths differing generically from each other. The material collected by Mr. Poole now makes the definite separation of these animals possible. One is slender limbed, resembling Acratocnus in size and general features; the other is more heavy, its general build probably somewhat as in Nothrotherium shastense. \ts bulk, however, though considerably exceeding that of Acratocnus, is not likely to have been much more than one-fourth that of the Californian animal. That one or both of these sloths continued to exist on the island until after the advent of man | have no doubt. The facts which have led me to this conclusion are as follows: (a) In the two caves near St. Michel most of the sloth remains were found within two feet of the surface ; and human bones and pottery occurred to the same depth without any indication that they had been dug in. (b) Near the en- trance to the smaller of the two main caves bones of ground sloths (certainly two and perhaps more individuals) were inextricably mixed with bones of man (adult and infant) and domestic pig. The remains were scattered among the small fragments of limestone which made up the greater part of the floor material, and I was unable to deter- mine any definite level-relationship among them. (c) Near the en- trance to the large cave | unearthed with a trowel, in fine, soft, undis- turbed material at the bottom of a trench two feet deep, the femur 1 Smithsonian Misc. Coll., Vol. 74, No. 3, p. 6, October 16, 1922. Sn 20 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 81 of a ground sloth, and, about 18 inches from it, a fragment of coarse dark pottery. There was no evidence of previous digging that I could discover ; and the bone and pottery had every appearance of having been deposited on the former surface of the cave floor and subse- quently covered by the gradual accumulation of detritus. (d) Both of these caves are situated on the side of a high ridge where the material composing their floors is entirely removed from the action of streams. (e) In general the ground sloth bones were. associated with the human remains in exactly the same manner as the bones of /solobodon and Plagiodontia, rodents which are positively known to have been contemporary with man. ACRATOCNUS (?) COMES sp. nov. Plate 5, fig. 2; plate 6, fig. 2; plate 8, fig. 1; plate to, fig. I Type—Right femur (lacking distal extremity) of adult, No. 253178, U. S. Nat. Mus. Collected in large cave near St. Michel, Haiti, March, 1925, by Gerrit S. Miller, Jr. Characters —A small ground sloth agreeing in general size with the Porto Rican Acratocnus odontrigonus Anthony ; its weight probably not exceeding 50 pounds. Femur resembling that of the Porto Rican sloth, and, like it, with a well developed lesser trochanter and without noticeable antero-posterior compression of the shaft, but modified for more directly perpendicular weight-bearing. Femur.—The femur differs from the corresponding bone of Acra- tocnus odontrigonus in at least two features which are important enough to indicate specific or, possibly, generic distinctness. (1) The intertrochanteric ridge is similar in position and development to the corresponding structure in A. odontrigonus, but it is supplemented by a large and conspicuous tubercle situated at the middle of the shaft at a level slightly below that of the lesser trochanter. This tubercle, of which no obvious trace exists in the numerous Porto Rican femora with which I have compared the Haitian specimen, forms the culminating point of a general thickening of the bone which imparts to the upper fourth of the shaft, as viewed from the side, a strongly angular-convex profile very different from the flat or slightly concave profile of the same region in A. odontrigonus (see pl. 6). (2) The neck is shorter than in Acratocnus odontrigonus and is less bent outward and forward from the axis of the upper half of the shaft; as a result, the head is set so as to diverge less noticeably from the general contour of the shaft. The differences in this respect between the Porto Rican and Haitian animals are of the same kind NO. Q MAMMALS FROM CAVES IN HAITI—MILLER 27 as those which exist in greater degree between the femora of Cholepus and Bradypus. The less anterior directing of the neck in the Haitian femur is perhaps most readily made apparent by applying the proxi- mal extremity of the bone to a flat surface in such a way that it is supported by the tripod formed by the posterior surfaces of the head and the two trochanters. The shaft of the bone in Acratocnus (?) comes then takes a position essentially parallel with the flat surface. When the femur of A. odontrigonus is similarly placed the shaft rises above the flat surface at an angle ranging from about 18° to about 23°. The same difference may be observed by tracing the direction of the low but usually evident ridge which crosses the neck from the head to the lesser trochanter. In Acratocnus odontrigonus this ridge extends so obliquely to the inner surface of the femur that its line, when continued downward, passes beyond the contour of the bone at a point situated near the mid portion of the head of the tro- chanter ; in the Haitian specimen it passes out nearly 10 mm. farther down the shaft. The lesser inward bend of the neck is best appreciated by “ sighting ” down the anterior or posterior surface of the shaft of the bone; it then becomes obvious that the head lies nearer to the main axis in the Haitian specimen than in any of those from Porto Rico. Remarks.—The femur on which this species is based resembles in all its general characters the corresponding bone of the Porto Rican ground sloths and of the Miocene South American Hapalops. The peculiarities which I have described as distinguishing it from the femur of Acratocnus odontrigonus separate it equally from the cor- responding bone of Hapalops, at least so far as can be determined from Scott’s figures of three species (Jongiceps, pl. 32, elongatus, pl. 41, and ruetimeyert, pl. 42). Other remains which I refer without much hesitation to Acratoc- nus (?) comes are as follows: (a) the proximal two-thirds of a right tibia (pl. 8, fig. r) not certainly distinguishable from the corresponding part of the tibia of a Porto Rican ground sloth (No. 17711, Amer. Mus. Nat. Hist.) ; (b) an almost perfect atlas (pl. 10, fig. 1) of the proper size to fit a skull of Acratocnus odontrigonus; several canini- form teeth, both upper and lower, agreeing in a general way with those of the same animal; (c) foot bones and ungual phalanges resembling those of the Porto Rican species. On the basis of the femur and of the parts which appear to be almost certainly associated with it I do not now feel justified in separat- ing the small Haitian ground sloth more than specifically from Acratoc- te apace Tp an 28 SMITHSONIAN MISCELLANEOUS COLLECTIONS voL. 81 nus odontrigonus. It would cause no surprise, however, 1f further material should indicate that the animals were generically distinct. The name comes alludes to the circumstance that the type specimen was found so closely associated with fragments of pottery as to lend strong support to the belief that the animal existed in Haiti as a contemporary of man. PAROCNUS gen. nov. Plate 7; plate 8, fig. 2; plate 9; plate 10, figs. 2, 3 Type.—Parocnus serus sp. nov. Characters —Femur differing from that of Acratocnus in the absence of the lesser trochanter; in the conspicuous widening and flattening of the upper half of the shaft; and in the more nearly vertical set of the head (as indicated by the line of the epiphyseal suture in an immature individual), a condition which appears to agree essentially with that present in Nothrotherium as shown on plate 12 of Stock’s Gravigrade Cenozoic Edentates of Western North America. Remarks—The genus Parocnus is readily distinguishable from Acratocnus by the structure of the femur alone. If I have correctly assembled the other parts which I believe to be associated with it there are many important differential characters. These parts are as follows: (a) a right humerus (pl. g), 200 mm. in greatest length, resembling that of Nothrotherium shastense as figured by Stock (Cenozoic Gravigrade Edentates of Western North America, pl. 8, fig. 1, 1925) in general form but less heavily built, with relatively broader proximal extremity and without the entepicondylar foramen present in this sloth and in Acratocnus; (b) the proximal third of a left tibia (pl. 8, fig. 2) and an entire left fibula probably of the same individual; (c) a right astragalus (pl. 9, fig. 3) very different from that of Hapalops as figured by Scott (Rep. Princeton Exped. Patagonia, Pal., Vol. 2, pl. 33, fig. 4) and Acratocnus as figured by Anthony (Mem. Amer. Mus. Nat. Hist., n. s., Vol. 2, pl. 73, fig: 7; 1918) but resembling in a general way, particularly in its calcaneal aspect, the very much larger calcaneum of Mcegalonyx figured by Stock (p. 87, fig. 31, 4, B,C, DD); (d)7 three caleaneay(2alete 1 right) of a form (pl. 9, fig. 2) conspicuously different from that seen in Hapalops and Acratocnus but essentially similar in plantar and astragalar views to the calcaneum of Mylodon as figured by Stock NO. 9 MAMMALS FROM CAVES IN. HAITI—MILLER 29 (p. 175, fig. 96); (e) a fragment of an atlas much larger than the corresponding part in Acratocnus odontrigonus or O. (?) comes. The area of the superior articular process in this atlas is nearly four times as great as that of another specimen from the same cave (the large cave near St. Michel) which I refer without much hesitation to O. (?) comes (pl. 10, fig. 1) ; (£) several foot bones and ungual phalanges of more robust structure than any known in the Porto Rican Sloth. PAROCNUS SERUS sp. nov. Plate 7; plate 8, fig. 2; plate 9; plate 10, figs. 2, 3 Type—Right femur (lacking epiphyses) of immature individual, No. 253228, U. S. Nat. Mus. Collected in large cave near St. Michel, Haiti, January, 1928, by Arthur J. Poole. Characters—An animal considerably larger and more heavily built than Acratocnus odontrigonus or A. (?) comes, its weight as roughly estimated by comparison of limb bones with those of pigs, probably 150 lbs. or more. Femur.—As compared with that of Acratocnus odontrigonus the femur of Parocnus serus (pl. 7) is immediately distinguishable by the absence of the lesser trochanter, as well as by its greater size and the much more noticeable antero-posterior flattening of the upper portion of the shaft. In a large femur of Acratocnus (No. 17716, Amer. Mus. Nat. Hist.) the two diameters of the shaft at middle of its upper half, lateral and antero-posterior, are respectively, 26 mm. and 17 mm.; in the type of Parocnus tardus they are 38 mm. and 14.5 mm. The ratio of antero-posterior to lateral diameters is there- fore about 65 in Acratocnus and only about 38 in Parocnus. At middle of shaft the discrepancy is slightly less: ratio of antero- posterior to lateral diameter about 61 in Acratocnus, about 45 in Parocnus. Below the middle of the shaft the diameters in the two femurs are essentially alike, with ratios of 58 and 59, a difference which is too slight to have any special significance. In addition to this striking peculiarity of general form the femur of Parocnus scrus is further distinguished from that of the known species of Acratocnus by the absence of a lesser trochanter and the presence of a low ridge about 35 mm. in length extending obliquely downward and backward from the middle of the neck across the narrow inner aspect of the bone to its posterior margin ; by the more thickened gluteal ridge; and by the presence of a noticeable con- — a Sg ae a te Bm Sa saa 2 eh deine nataateaiatatan rd —_— ee ee = 30 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 8&1 cavity on the posterior face of the shaft at the base of the great trochanter. Unfortunately no perfect skulls of ground sloths have yet been found in the Haitian caves. One specimen from the small cave near St. Michel includes the interorbital region and anterior part of the braincase. It is about the size of the corresponding part of the skull in a large Acratocnus odontrigonus, but is conspicuously different in form, owing to the absence of the deep postorbital constriction which is such a noticeable feature in the skull of Acratocnus. Whether this fragment pertains to a skull of Parocnus or of Acratocnus (?) comes is a question which cannot be answered. A fragment of a palate from the same cave appears to have come from a skull of much the same size. It indicates a palate twice as wide in proportion to the length of the toothrow as that of Acratocnus odontrigonus, and it further differs from the palate of the Porto Rican sloth in the presence of a median longitudinal ridge supplemented, on each side, by a shallow but well-defined longitudinal furrow. The toothrow in this individual was probably of almost exactly the same length as that of the Porto Rican specimen figured by Anthony on plate 69 (fre ie) SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 815 NOs 9), RES 41 va aes Zi * : ‘ (All figures natural size) 1. Nesophontes paramicrus Miller. Nos. 253062-253076, U. S. Nat. Mus. a, 2 skulls (the type at right) ; b, 6 mandibles; c, three humeri; d, 3 femora; e, innominate. 2. Nesophontes hypomicrus Miller. Nos. 253077-253080, U. S. Nat. Mus. Letters as in fig. 1. Type skull at right. 3. Nesophontes samicrus Miller. Nes. 253090-253094, U. S. Nat. Mus. Letters as in fg. 1. Type skull at left. SMITHSONIAN MISCELLANEOUS COLLECTIONS VOR 815 NO. 95 Ris? Yr" | 4b (All figures natural size) Brotomys voratus Miller. No. 253097, U. S. Nat. Mus. Brotomys (2) contractus Miller. Type. Isolobodon levir (Miller). No. 253117, U. S. Nat. Mus. a. [solobodon levir Miller. No. 253102, U. S. Nat. Mus. Aphetreus montanus Miller. No. 253133, U. Nat. Mus. 4a. Aphetreus montanus Miller. No. 253145, U. Nat. Mus. 4b. Aphetreus montanus Miller. No. 253151, U. Nat. Mus. +HWWN ~ nnn SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 81, NO. 9, PL. 3 1 la ff Ib 2 (All figures natural size) 1. HHexolobodon phenax Miller. Type specimen. Crowns of maxil- iP lary teeth. 1p ta. Hexolobodon phenax Miller. No. 253125, U. S. Nat. Mus. Crowns it of mandibular teeth. th. Hexolobodon phenax Miller. Type specimen, showing alveoius of left incisor, roots of maxillary teeth and intervening floor ot narial passage. 2. Capromys pilorides Desmarest. No. 253232, U. S. Nat. Mus. it Palate broken away from rest of skull in the same manner as hy the type of He.xvolobodon to show corresponding structures. Ai i Fi SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. <8i1),-NO._ 9 Pi 4 2a (All figures natural size) 1. Elasmodontomys obliquus Anthony. Immature, with third molar not yet above level of alveolar rim. No. 17137 h, Amer. Mus. Nat. Hist. 2. Ouemisia gravis Miller. Type. Same stage of growth as fig. I. In both specimens the alveolus of the incisor has been tapped at its base and a black thread passed through the tube. SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 81, NO. 9, PL. 5 (Both figures natural size) Da 1. Right femur of Acratocnus odontrigonus Anthony, anterior aspect. No. 17711, Amer. Mus. Nat. Hist. 2. Right femur of dcratocnus (7) comes Miller, anterior aspect. Type. SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 81, NO. 9, PL. 6 2. (Both figures natural size) | 1. Right femur of Acratocnus odontrigonus Anthony, outer i aspect. No. 17711, Amer. Mus. Nat. Hist. i) 2. Right femur of Acratocnus (?) comes Miller, outer aspect. Type. SMITHSONIAN Right femur of MISCELLANEOUS COLLECTIONS (Natural size) Parocnus serus Miller, anterior VOL. 81, NO. aspect. Type. ORE SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLE. .8'1), (NO" 19) (PES 8 (Both figures natural size) 1. Right tibia of Acratocnus (?) comes. No. 253179, U. S. Nat. Mus. 2. Left tibia of Parocnus serus. No. 253230, U. S. Nat. Mus. SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLES Sil, NO=.9 PEAS (Both figures 34 natural size) 1. Right humerus of Parocnus serus Miller, anterior aspect. No. 252231, U.S. Nat. Mus. ia. Right humerus of Parocnus serus Miller, outer aspect. No. 253231, U. S. Nat. Mus. SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLES, eNOS ReoatO, (All figures natural size) 1. Atlas of 4cratocnus (?) comes Miller, anterior aspect. No. 2353181, U. S. Nat. Mus. 2. Left calceaneum of Parocnus serus ? outer aspect. No. 253226, Wise Nata Mins: 2a. Left calcezneum of Parocinuts serus US. Nat. Mus. 3. Right astragalus of Parocnus serus USS NateMus: 3a. Right astragalus of Parocnus serus ? inferior aspect. No. 253220, Ua SeaNate Nias: % ’ 3 dorsal aspect. No. 253226, > outer aspect. No. 253220, oe “y and e's . $i * ¥ “iy NN ~~ eye see oa Bh aie ) SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLUME 81, NUMBER 10 | | TROPISMS AND SENSE ORGANS OF al LEPIDOPTERA BY N. E. McINDOO ~ Senior Entomologist, Deciduous-Fruit Insect Investigations, Bureau of Entomology, U. S. Department of Agriculture o as (PUBLICATION 3013) ; See, gd) ee Aa \' aU aa INST TRS - i 27 ote j , (APR 5~ 929 OF Fie. tee GITY OF WASHINGTON eS RIN ss ad ~ PUBLISHED BY THE SMITHSONIAN INSTITUTION APRIL 4, 1929 ts Is t : i eA CL gtx rar oe \ > 7 1 —p Le Gaull be "y SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLUME 81, NUMBER 10 TROPISMS AND SENSE ORGANS OF LEPIDOPTERA BY N. E. McINDOO Senior Entomologist, Deciduous-Fruit Insect Investigations, Bureau of Entomology, U. S. Department of Agriculture (PUBLICATION 3013) CITY OF WASHINGTON PUBLISHED BY THE SMITHSONIAN INSTITUTION APRIL 4, 1929 a a x = ye » o = — < i 2 » °o a 2 a » BALTIMORE, MD., U. S. A. ’ TROPISMS AND SENSE ORGANS OF LEPIDOPTERA |: By N. E. McInpoo SENIOR ENTOMOLOGIST, DECIDUOUS-FRUIT INSECT INVESTIGATIONS, BUREAU OF ENTOMOLOGY, U. S. DEPARTMENT OF AGRICULTURE CONTENTS PAGE inne Uc timmy cee teraise ciscrb hey) sf code love’. iste ureteocha tone, Meee te ee Nt arctic me eee 2 PINTS IES OME P IE ERC ee ccc F30 Gok a avaiasw, aichr Masato RRs & orci nanan eke 2 Hmm IHL OR ASSIS HAM te iio) Shea) 4 fala: Salis oa te! ecchayttehear aso EMEP AF ata de RO Pe ee aes 3 PMPINON ICN TO MITEL AGUNG 425.3 ss) clase a aleekaerelete nid erate aes natal stan tote 3 (a) Definitions and problems in the study of light reactions... 3 (b) gre lichtcreactionsadaptive:t..cs'« 2ai.cetiaes eure ste eee 4 (c) Is orientation accomplished by selection of trial movements? 5 (d) How do light rays bring about orientation?............... 5 (e) Do circus movements support Loeb’s theory?............ 5 ({) What wave lengths stimulate insects most?.............-- 6 (g) Light traps are not yet considered successful............. 9 2. Phototactic experiments on codling-moth larvae..............-. 10 HTP Cle OLAESI SH eyeeiroG ees ais iow ovsn 8.509% ayouats hetavdee coercion ioe 14 Tepe eve WIHOtOItCLAtUITEs «i: «srs cree alate Sates een eee 14 2. Chemotactic experiments on codling-moth larvae............... 17 IIE Geo tart Steere See cess creed. ocean eel Ue SIS Rea a Ere 19 Pee OW OTAMILEHALILEK sie, sia(s ss alee aerate ete ere a SL oe eet eae 19 2. Geotactic experiments on codling-moth larvae.................. 20 LRN PemmeInE To Antec Smet Ae otal fb ehcices of 2's aim wi. 38 aie aoe eceleh oravokere oat) Srateemolei a oebaee Brae 21 We RCULG WAM OMEDIIECLATEILE cfs. > chb\s-<,5-alece.v.0 ma sie epee ololely aver creer: creer 21 2. Thigmotactic experiments on codling-moth larvae.............. 22 AM GOPICTECEDUONSW. fice claret ucisrs aia a’ o/u'4 64.4 $0 Sloramndee Oe erlereioe metal ae store 2 TM LAOLORECEDEOUSI Ciara iss aes toh Go Shae Noes ee ee ene 23 Hii rein eCeplLOLS wows thy oeiie -ne-siets 3/228 Side heat a en eee 24 iy So=ceiliigal Gigeieray CiBEMEo gooodtuowacsocododuadoscbadadnnoec 24 (Cay eeAnnitennaltorganse Acs cs kes os cit ars Boreas ee 24 Ch) Ma@liactotyp spores. waxes econ oie oe ee ee ern 27 Pees O-CalledmtastenOLreansio-vn toc, snes OR Mee ee eee 35 SR ANI CIITFECED LOLS) vera o./'5 che arctata cals ashe | Leh aha chien Ae ea re eo es 39 epee NV AITIPANI CHO ANS. |, Wistar secsicdd clio e eee ACT Reta 41 ZeeChOLdotonial ORSANS: wats ics .-2%s.diel reco, Sata ee Ae I ae 41 AMM OL StONMOGS ANS ics ev ka overs cae rare eC ere 42 APP ATICICO TIVE ALES) cious, syacs steies/e detest LOE eT 44 Weebl te 1nOTeceptOnrse (racvaaisc cies cos ao oka loc Ao Oe aaa 45 PeMACHIEMOESANISL {00500 5 ad). ats ctl cis Ae A ean ae 45 Wie GEOLECEDLOTS: iy. @ oede sieit Saw db aan eee ee ed 2 eee 46 PALACE OLZANS) .c: 2 «4+ ta) os nee si aie ee LO 46 RIMM PREEETCCODLOES, vhs ois Ci boas ware Nee DR eo ae ee 47 MeSECHI-NEOUNCING MOFZANS a. 5 iic d2.b4/oieaclnceee weeeiek nee eae 48 UUM EV ge ATI’: CISCUSSIOND: «2:21 ankenee | Aatenna Total | i autenia antenna Total ‘a | l sail 3 A | a n = o no n vu eral ob in bo a | oo BS, a oo é | ob a m0 SOU eee v &, 2 | & ts S ee o a Bie ES > x i 5s meee = 2 * > se lee n a n | a alice a n a a a | | | I} 36| 308 Al) 3100) 77 27 || 1 46 | 331 || 35u| 1345.) St 76 2 5382). 268 030) 5 277; | 74| 545 || 2| 40] 330] 40 360 | 80] 690 3:| 367) 366 | 33 | 373)| 60) 739")|3)| 36) 386) 35.) "368 |o7m e754 4 | 37) 206'|: 34 | 2041) 71) 5Q0)|I"4 ) (35°) 354.) 35.1) S47 5 7e4,, gor 5| 36) 284) 31 | 244 | 67| 528)}5 | 47 | 257 | 43 260) 90) 517 | z A |_ Average for male | Average for female ANfemMaee te ckis erie | 72 | 606 aM{ennaess® (sce) | 781) MOOS * A few styles on each antennae do not bear end pegs. 25 TROPISMS OF LEPIDOPTERA—McINDOO Aueut AIOA Moy Aueut AJOA Aueul ” ” Auewt AIOA MO} ” Aueu oO Oo oO oor Aue AIOA oZ 009 ” uy ” ” Auewt A1dA Oo Aurw AJIOA Mo} Aueut AI9A Auew MJ oo0o°o eee eee Sve je\ai's S616) alee cee eee “snjlod} oyideg SSeder eljuog sfc) or pliorg siisiielvelisisitrieikei site(s edornue essoue \ teeeess gfaqho stuudSIY eee °6 ” ”? Fee ee ee ee ee eee tunid ouy Sivitsiie)s)ie) “oss eoIne eA see 6 *, **P ejauowod esdesodiey se ee we ew ae PIJIstis RIOINIY o- “++ © BSOI]IXO BaplouuIUUPS stwiojoeioawayds xAsojdopiiAy y, . ee eee 3} ee ee eee . } oe . C0 OO O10 ROO 07s = - COO ORO. GO O'6: fo) elieruid eIuOpl yy ° & ” a “/p eliejowod epiydos,y tesa ced e1do.1999 BIWes ” Peseta sojoorun ayoAsg SRST a aS Sac ad a ee heree Tei enbrjue eIASIO oMeks hese maieney 7s 5. = POD OSES KONI xAquiog “** VURIIIOWE BLUOSODRI LIT BUSIJSOONI], edueso1awia FY 899 SHSM a avavah oa SiaaKalisiay sess TOTOOIUN STJOISY ee ee “"-T]eVsOyyuIe eluapolg op “+553 vauno erqueyds Hee dsrcisoiuedy sree sssoijads paymuapiug, oedjejed eruiojyeiay “svyepnoeuonbuinb snijuoyioso[yg Pe)": sepruorideg "Ze "I€}" “aeprpeyduA Ny ‘of ‘6z|" + aepiusesi7 ‘Qzlaepijnowouod x “Lz ‘Qc|* aePIINIIYIIO AAO OO aig HM ay hel" ** * aepiliasoy On cremate “aeplyoIAs g ‘oo “1c ‘0c ‘OI|" * FVPIIJOWIODL) Q1\" + Oeplluinyes vat ‘OI sa “PI “C1 ‘Z1\"*° aeprAquiog *II|'oeprdwesoise Ty ‘orl: tt aepliediy 6 8 is . “*"QepIn}ION so a “bP ie iovene) e “OePIIpIVY a5 iG "IT [cs oeprsuryds ” + Be O O ” ” 0 O ” ” 0 0 juasoid Aueuw Oo oO ehcrs Mo} ozI parse rei a a ozI eratets se | ” yy, 1B} ” a re Aueul “s juasaid ” %” Mo} ” yay | Auew A1dA o _ a | ” ” Mo¥ serene i Auew AoA oO ste Maj Corl eels ie SIC | cc - Lit . neers Aueut AIOA O 3 me AUPUL Aueut AIOA Se quosoid | Aueur AIDA M9J Moy oesioye | O O Oran pak: | Aueuwt AJOA | juasoid = se eae MdJ av . lems ” 08 : 5 | ” a” ” ” = ” ” ” ” eis | Auewt AIOA AuPul e sci Mo Moy - ae +) 99 ”? ” i ae oY ” ” ” ae Auew AIA a kA Z 5 ” ” ” Auew Auruw a = ae ” ” Mot 9? jussoid | Oo ” sae ” 7” 0 ” ee Auew AIdA Mo} Mo} suesi0 (BIpOYsi4y *S)} (BoTaeYO *S) Bonk wojsuyo SIIB aSUIG |SajzSliq asuag 14S (B91U030, Ays *S) sded pug (@91W090]309 "S) sdod 1g Sue310 9Sues JO 1aquUINU puUL pully so1dads j,0 9WieU pue 1oquiny psaidopiday jnpvo {oO apuuajun uo Supb4AO asuas fo AIQuinu a2ijDADquo)—T ATAV J, Ay 1ue 26 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 81 The antennal organs of five male and five female codling moths were carefully examined by the present writer. Little or no sexual differences were observed in the antennae or their organs. The antennae of both male and female are filiform and bear the same kinds and practically the same number of organs (tables 1 and 2). The number of segments in the antennae of males ranges from 55 to 61 with 58 as an average; those in the antennae of females from 59 to 63 with 62 as an average. Each antenna bears one Johnston organ NW i Fic. 3.—Antennal organs of female codling moth, No. 3. A and B, External views, X 125. C to G, Sections; C to F, & 500; G, & 320. A, Second and third antennal segments; B, two segments from middle of antenna; C, pit peg; D, style and end peg; F, sense hair and its innervation; F, non-innervated scalelike hair ; and G, cross section through distal end of segment near middle of antenna. Abbreviations: J, Johnston organ; N, nerve; P, olfactory pore; S, style; Sc, sense cell; Sca, sense bristles (Sensilla chaetica) ; Sco, pit peg (S. coelo- conica) ; Sh, non-innervated scalelike hair; Ss, end pegs (S. styloconica) ; St, sense hairs (S. trichodea) ; Tc, trichogenous cell; and 77, trachea. (fig. 3, A, J), 2 or 3 olfactory pores (P), numerous pit pegs (fig. 3, B, Sco), end pegs (Ss) on styles (S), sense bristles (Sca), sense hairs (St), and scalelike hairs (Sh). Each of these, except the last named, is supposed to be a sense organ, and Freiling (23) has even pictured a slender scalelike hair of another moth as connected with a sense cell. Of these seven structures only the olfactory pores, pit pegs, and end pegs are supposed to be olfactory in function. . Pit pegs may be found on all segments, except the first, second, and the last one or two, of codling-moth antennae. If odors can pass NO. I0 TROPISMS OF LEPIDOPTERA—McINDOO 27 quickly through chitinous structures, pit pegs (fig. 3, C, Sco) would be excellent olfactory organs. Styles, usually terminating in end pegs, may be found on all segments except the first and second. A style (fig. 3, B, S) is nothing more than a prolongation of the distal outer edge of the segment and it is supposed to be innervated, but in codling- moth antennae it (fig. 3, D) has no nerve and consequently cannot be a sense organ. The writer has failed to find a drawing by any author showing a nerve connected with this structure. The antennae of 21 other species (table 2) examined by the writer varied much in respect to barbs, from typical filiform antennae to fully feathered ones. The sense organs, as a rule, were widely dis- tributed on both the shaft and barbs. In Argynnis cybele the pit pegs lie only on the club part of the antenna. Some of them are large and irregular in shape, and perhaps a pit bears more than one peg. In 11 of the specimens pit pegs were totally absent and in 12 no end pegs were observed on the comparatively few styles, and even styles were absent in one moth (No. 23) and in all the butterflies (Nos. 31 to 34) examined. From the preceding it is evident that pore plates (S. placodea), common to three orders of insects (aphids, beetles, and bees and wasps), are totally absent in Lepidoptera, while the pegs (S. basi- conica) are practically wanting. These two types are the ones gen- erally considered as olfactory receptors in most insects. It is said that the end pegs and pit pegs of Lepidoptera replace the pegs and pore plates of other orders, but there is no proof whatever for this assump- tion, and furthermore it is doubtful whether the end pegs are ever innervated. : Granting that the pit pegs and end pegs are the only olfactory organs of Lepidoptera and drawing conclusions from the observations of Schenk and the present writer, eight individuals (table 2, nos. 1 to 4, 17, 18, 23, 24) of the 34 specimens examined cannot smell at all, while four other individuals (nos. 10, 19, 20, 25) have comparatively few end pegs as olfactory receptors. (b) Olfactory pores—At the suggestion of his reviewers the writer (48) in 1914 called the sense organs herein discussed “ olfactory pores.” Guenther (27) in 1901 seems to have been the first to describe the internal structure of these organs in Lepidoptera. He called those in the wings “ Sinneskuppeln ” and found their structure to be similar to that described by the present writer, although he did not see the pore aperture passing to the exterior. Vogel (88) made a more ex- tended study of them in the wings of many Lepidoptera. He con- 28 SMITHSONIAN MISCELLANEOUS COLLECTIONS voL. 81 sidered them as chordotonal organs. The present writer (50, 51) made a thorough study of the disposition and structure of them in many Lepidoptera and their larvae. Priffer (77) has most recently described these pores on the wings of certain moths in connection with the antennal organs. The olfactory pores on five male and five female codling moths were counted. Female no. 3 was examined most carefully and conse- quently the greatest number of pores was found on it. In figures 4 and 5 they are represented by black dots. The groups are numbered from I to 12. The isolated pores which are constant in position are designated by a to e, the others not being thus marked. Groups 1 to 6 and pores a and b lie on the wings (fig. 4); groups 7 to 11 and Fic, 4.—Semidiagrammatic drawings of wings of female codling moth, No. 3, showing location of olfactory pores (1 to 6, a and b, and other dots), sense scales (Ssq), and sense hairs (St), X 5. A, Dorsal side, and B, ventral side of front wing; and C, dorsal side, and D, ventral side of hind wing. Vogel (88) shows “ Sinneskuppeln” or olfactory pores similarly located on wings of Scoria lineata. pores c and d on the legs (fig. 5, A) ; group 12 on the base of the labial palpus (fig. 5, E) ; and pores e (fig. 5, D) on the maxilla (one- half of proboscis). The number of pores in the groups follows: Nos 1;,91; No.-2;:70; No. 3, 525 7Nio.4,712) No. 5, 120 siNOmomn zr No. 7, 4; No. 8; 4; No. 9, 5; Noo10, 4; No. 11,7; and) Nomi: The total number counted on female No. 3 follows: Legs, 191; front wings, 462; hind wings, 417; proboscis, 28; labial palpi, 16; and second segments of antennae, 4; making 1,118 in all. The total number of pores on males range from 933 to 1,049, with 986 as an average ; and on females from 960 to 1,118, with 1,029 as an average. Figures 6 to 8 represent the internal structure and innervation of the pores and sense hairs, and also the internal anatomy of the wings and legs where the pores are found on them. Fic. 5—Legs, maxilla, and labial palpus of female codling moth, No. 3, show- ing location and structure of sense organs on these appendages. A, Inner and outer surfaces of hind leg; B, same of middle leg; and C, same of front leg, showing location of olfactory pores (7 to 11, c and d, and other dots), sense bristles (Sca), and sense hairs (St); X 5. D, Maxilla or one-half of proboscis, and FE, labial palpus, showing location of olfactory pores (e and 12), pegs (Pg), sense bristles (Sca), sense hairs (St), and labial-palpus organ (Bo); X 32. F to M, Structure and comparative sizes of sense organs on proboscis; X 500. F, External view of peg; G, looking down on its tip end; H, cross section of peg; and I, longitudinal section of peg, showing trichogenous cell (Tc), sense cell (Sc), and nerve (N). J and K, External and internal structure of smallest sense hair. L, External view of sense bristle. M, External view of two olfactory pores. 30 SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL. 81 Lepidopterous larvae can smell, but they do not have the so-called olfactory organs such as pegs, pore plates, pit pegs, or end pegs like or even similar to those of adult insects. Therefore, it is only reason- hodus Subcosta ic. 6.—Cross sections of wings and proboscis of Lepidoptera, showing in- ternal anatomy of wings and olfactory pores. A, Semidiagrammatic drawing from an oblique section through front wing of cabbage butterfly, showing groups 2, 3, and 4, of pores, sense cells (Sc), nerve (N), and trachea (77) ; x 100. B, Pores from wing of codling moth; * 500. C, A pore from proboscis of codling moth; X 500. Fic. 7—Schematic drawing of wing of a male Saturnia pyri L., showing innervation of olfactory pores (P) and other sense organs, including scattered pores and various types of sense hairs. The black dots represent those on the dorsal surface, and the circles, those on the ventral side. (Copied from Prutier(77)):) able to suppose that the pores, called olfactory by the writer, act as smelling organs. The olfactory pores of five specimens of each larval instar were counted. Little or no difference in position and number of the pores NO. IO TROPISMS OF LEPIDOPTERA—McINDOO 31 was observed in the six instars. They are found widely distributed (figs. 9 and 10) as isolated pores or “ punctures” on the following parts: Head capsule, 24; maxillae, 16; mandibles, 4; labrum, 2; labium, 6; antennae, 2; legs, 30; first thoracic segment, 4; and anal prolegs, 4; making 92 in all. Some of those on the head capsule were incorrectly named in 1919 by the writer (51), but in figure g they are correctly named according to Heinrich’s (31) first paper and later ones on this subject. In regard to experimental results concerning olfactory receptors, two papers will be briefly reviewed. The first and most important Trochanter a Pd » ig LEE b, SF "Zz ‘y Fic, 8—Semidiagrammatic drawing of an oblique section through femur, tro- chanter, and coxa of a silkworm moth, showing muscles (Jw), trachea (Tr), nerves (N), sense cells (Sc), sense hairs (St), and groups 8, 10, and 11 of olfactory pores, No. ro being shown partially from a superficial view; X 100. experimental work to decide the function of the olfactory pores was done by the writer (48) on honeybees. Of the six sources of odors used three were essential oils. The writer’s critics have apparently overlooked the fact that the results obtained by using the other three odors are reported in such a manner that they can easily be considered alone. Since the odors from the oils might have been irritant, let us consider the other results, which, when expressed in percentages, clearly show how closely the percentage of pores supposed to function corresponds to the percentage of response obtained. On the average, a worker honeybee has about 2,800 olfactory pores. On the bases of the four wings lie 54 per cent of them; the legs possess 23 per cent ; 3 32 SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL. 81 A Itc. 9.—Disposition of olfactory pores or punctures on head and first thoracic segment of a fully grown codling-moth larva, 20; A, dorsal view; and B, ventral view. Abbreviations: Frontal pore (Fa); adfrontal pore (Adfa); ocellar pore (Ob) ; anterior pore (da); posterior pores a (Pa) and b (Pb); lateral pore (La) ; ultraposterior pore (UPa); subocellar pores a (SOa), b (SOb), and c (SOc); genal pore (Ga); mandibular pores a (Mda) and b (Mdb); maxil- lary pores a (Ma), c (Mac), d (Mad), e (Me), and f toi (M-f-i) ; labial pores a to c (Lba-c); labral pore (Lr); antennal pore (Ant); and thoracic pores a (T/a) and b (TIb). 8 ACI Fic. 10.—Disposition of olfactory pores or punctures on legs of fully grown codling-moth larva, * 30, A, Anterior and posterior surfaces of prothoracic leg, and B, dorsal and ventral surfaces of anal proleg. ‘Abbreviations : Femoral pores a (Fea), b (Feb), and c (Fec); tibial pore (Tha) ; tarsal pore (Ta): and anal-proleg pores a (Apa) and b (Apb). NO. IO TROPISMS OF LEPIDOPTERA—McINDOO 33 while the others lie on the sting, head, and head appendages. The indi- viduals were allowed 60 seconds in which to respond. All of the pores on 31 unmutilated bees responded to the odors from honey, pollen, and leaves of pennyroyal in four seconds (48, pp. 283, 284) ; that is, in one-fifteenth of the entire maximum time allowed for the response. Twenty bees with their legs covered with a mixture of beeswax and vaseline, leaving supposedly 77 per cent of the pores elsewhere to Fic. 11.—Diagrams of Minnich’s apparatus used in testing insects to olfactory and gustatory stimuli. A, Section of an odor chamber made of a rectangular museum jar, showing a butterfly, held by a wire (w) and a spring clothes pin (s), responding with extended proboscis (p) to apple juice (a). (After Min- nich.) B, Perspective view of apparatus used to show that butterflies “taste ” with their tarsi; a and b, two small rectangular tin pans, a containing a cheese- cloth pack wet with apple juice and b containing a similar cloth wet with dis- tilled water; and d, a Petri dish nearly full of apple juice in which stand the ‘tin pans just beneath two openings in a wire screen (s). The arrows, 1, 2, and 3, represent the positions in which the butterflies were tested, the position of the walking legs being indicated by the cross-bars. (Redrawn from Minnich’s two figures. ) function, responded 2. 5 times more slowly (p. 336) or gave a response of 83.3 per cent. Twenty-eight bees with their wings pulled off, leaving 46 per cent of the pores elsewhere to function, responded eight times more slowly (p. 335) or gave a response of 46.7 per cent. And finally, 20 bees with their legs covered with the beeswax-vaseline mixture and their wings pulled off, leaving supposedly only 23 per cent of the pores located elsewhere to function, responded 11 times more slowly (p. 337) or gave a response of 26.7 per cent. 34 SMITHSONIAN MISCELLANEOUS COLLECTIONS voL. 81 Cabbage butterflies (Pontia (Picris) rape L.) were confined by Minnich (63) in an odor chamber (fig. 11, A). Since they are fond of apple juice its odor was used to stimulate the smelling organs, and the responses to it were then measured by the extent to which the proboscis was uncoiled for the purpose of partaking of the apple juice, although the insects could not reach it. The antennae were mutilated in three ways: (1) Covered with vaseline; (2) covered with a mixture of paraffin and vaseline; and (3) cut off at the base with fine scissors. When the organs on only one antenna were pre- vented from functioning, the olfactory response was reduced only 6 per cent ; when those on both antennae were eliminated or prevented from functioning, the response was reduced 58 per cent. Thus accord- ing to these results nearly half of the olfactory receptors must be located elsewhere than on the antennae. In his own words Minnich (Pp. 354) says: After the antennae are eliminated the animals were still 42 per cent responsive. Considering the variety of methods employed and the similarity of results ob- tained, this figure is much too large to be attributed to a failure to eliminate the antennal organs completely. It must, therefore, mean that there are olfactory or- gans on other parts of the body as well as on the antennae. . . . I cannot, there- fore, concur with McIndoo in the view that the antennae of adult insects in gen- eral lack olfactory organs. Certainly, such is not the case with Pieris. Nor can I agree with the opposing viewpoint, that the olfactory organs of adult insects in general are confined to the antennae. In this respect the results on Pieris differ from those obtained by v. Frisch in his ingenious experiments on bees. The results of the present experiments show that a viewpoint intermediate be- tween these two is correct for Pieris, and that while the antennae constitute a very important, probably the most important, olfactory region of the body, they do not constitute the sole olfactory region. Regardless of the results obtained by testing insects with mutilated antennae, it has never seemed reasonable to the writer to suppose that odorous air can pass quickly through the hard and dry chitin covering the antennal organs. If it can, why not grant the same privilege to all sense organs covered with thin chitin, including all kinds of sense hairs and even the olfactory pores whose sense fibers, according to other authors, are separated from the outside air by a thin layer of chitin? In the higher animals the olfactory organs (fig. 12) are separated from the outside air by only a thin watery layer of mucus, and the latest results show that the free ends of the olfactory cilia actually come in contact with the air. Eidmann (18) erroneously supposed that the chitinous intima of insect intestines is similar to the coverings of the so-called olfactory and taste organs of insects. He proved chemically that aqueous solutions can pass slowly ee NO. 10 TROPISMS OF LEPIDOPTERA—McINDOO 35 through the intima when the latter is wet on both sides. From this result he concluded that the olfactory organs of insects need no openings through which the nerve endings can come in contact with the odorous air outside. The present writer cannot see any connection between his findings and the chemical sense receptors of insects. J, © AS + Mec Hee ca 9 ‘, : Fic, 12—Olfactory and gustatory organs of higher animals. A, Diagram of a block from olfactory mucous membrane of a kitten, showing in section and per- spective the following: Basal cells (b), olfactory cilia (c), nerve fibers (f), limiting membrane (/), olfactory cells (0), supporting cells (s), olfactory vesicles (v), and walls (w) of the five- and six-sided supporting cells from a surface view. The olfactory vesicles and cilia, which are embedded in and sup- ported by an outer semifluid (not shown in drawing), are the true receptors of smell. (Redrawn from van der Stricht’s (84) photomicrographs and figure 36, the latter in Herrick’s book (33).) B, A single taste-bud from human tongue, showing nerve fibers (f) indirectly innervating the surrounding epithelium (e), supporting cells (s), and taste cells (¢), whose outer ends project into and sometimes beyond the pore (p). (From Herrick (33), after Markel-Henle.) 2. SO-CALLED TASTE ORGANS The so-called taste organs of Lepidoptera, according to Deegener’s review (see Schréder (80) p. 149), consist of two round groups of sense hairs on the under side of the pharynx. The proboscides of Rhopalocera, Noctuidae, Geometridae, and Bombycidae bear at their tips more or less numerous peg-shaped structures of varied lengths and shapes in different species. In Sphingidae and Zygaenidae these pegs are distributed over the entire proboscides. These peculiar 30 SMITHSONIAN MISCELLANEOUS COLLECTIONS voL. 81 structures have been considered both tactile and gustatory in function. Lepidopterous larvae bear on their antennae and mouth parts variously shaped sense hairs, some of which have been called taste organs, some touch organs, and others smelling organs. Each maxilla on the codling moth bears about 50 pegs or about 100 for the entire proboscis. Female No. 3 had 93 of them (fig. 5, D, Pg). They are always found on the distal half of the maxilla and are usually six-sided (fig. 5, F to H) but a few are five-sided. Each one (fig. 5, 1) arises from the proboscis as a fluted column and terminates in five or six sharp pinnacles, which surround the innervated hair. If aqueous liquids, or odors, in order to stimulate the nerves inside the hairs, can pass quickly through the chitinous walls, we can then safely call them taste organs, or smelling organs; if such a condi- tion is not true, they are certainly nothing more than touch receptors. The writer has repeatedly objected to the chemical-sense assumption, but believes that smell and taste in insects are inseparable and that the olfactory pores are their only receptors. Figure 12 shows the similarity of olfactory and gustatory organs in the higher animals and that the stimuli do not pass through any membrane in order to reach the nerves. Let us now consider the chemoreceptors found by Minnich on the tarsi of butterflies and flies. The two species of butterflies used by Minnich (60) may often be seen to alight on injured tree trunks or on decaying fruit in orchards, apparently for the purpose of feeding on the exuding sap of the tree or on the juice of the fallen fruit. In the presence of food it was further observed that the proboscis would uncoil and then coil up again in a definite manner. Minnich called this reaction of the proboscis a proboscis response, and later made use of it solely in measuring or weighing the responses of butterflies to various liquids. In order to determine the responses of the tarsal chemoreceptors, and at the same time to control the olfactory responses, maddy experiments with butterflies in confinement were conducted by using an ingenious and specially constructed apparatus. Briefly stated, the apparatus consisted of a shallow dish (fig. 11, B, d) covered with wire screen (s), in the center of which are two small rectangular openings, which lie just above two small rectangular tin pans (a and b) inside the dish, each containing several layers of cheesecloth. The cheesecloth in one pan (@) was wet with apple juice and that in the other pan (0) with distilled water; and the shallow dish was also full of apple juice. A butterfly to be tested was held by the wings with a spring clothes-pin in position 7; that is, NO. IO TROPISMS OF LEPIDOPTERA—McINDOO 37 with the four feet of the middle and hind legs touching the wire screen and with the antennae extending directly over the cheesecloth wet with apple juice. Since the front legs are rudimentary and not used for walking, they were not considered in these tests. If the insect responded at all in this position, the response was a truly olfactory one. The butterfly was next held in position 2; that is, with the head and antennae just above the cheesecloth wet with distilled water and with the feet of the middle legs resting on this wet cloth. If the insect responded at all in this position, the response was either an olfactory one or one brought about by contact with the feet on the cloth, or the response was a combination of both oifactory and contact stimuli. The butterfly was finally held in position 3; that is, exactly like posi- tion 2 except over the cheesecloth wet with apple juice. In this position the insect always responded, and the responses were of the same kind but differed in degree from those in position 2. As an average for all the responses obtained in the three positions, position 1 gave 29 per cent ; position 2, 17 per cent; and position 3, 100 per cent; clearly showing that these butterflies can distinguish apple juice from distilled water merely by bringing their feet in contact with these liquids. In other series of tests Minnich used solutions of common sugar, table salt, hydrochloric acid, quinine, and distilled water. In order to compare closely the results obtained, the first four substances were used on the basis of their molecular weights. Butterflies were able, by means of their feet alone, to distinguish the sugar solution from those of the hydrochloric acid and quinine, or from distilled water ; and the salt solution from either sugar solution or distilled water. Now the question naturally arises: Are there special sense organs in the tarsi of butterflies, which act as contact chemoreceptors? Minnich gives us definite information about their function, but leaves us in the dark concerning their exact location and structure. Experimentally he located them on the four tarsi of the middle and hind legs. Each tarsus is five-jointed, the first joint being about as long as the other four combined. Minnich believes that these organs lie in the distal end of the first joint, and particularly in the other four joints. He further believes that they are not temperature organs, touch organs, or organs to register the penetrating powers of liquids, but are chemi- cal sense organs, perhaps somewhat similar to taste organs in man. Excepting tactile hairs, there are no other known sense organs in the tarsi of butterflies, although no one apparently has looked for other sense organs at this place. In 1917 the present writer (50) reported finding olfactory pores on the legs of butterflies, but found 38 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 81 none on the tarsi. Recently he has more carefully examined the tarsi of six species of butterflies. No chemoreceptors were seen on the dark and hairy tarsi of three species, but on most of the light colored ones of Pontia rape, Papilio polyxenes, and P. trotlus a row of supposedly olfactory pores were observed on each tarsus. They are very plain on the tarsi of the cabbage butterfly (Pontia). A few pores were also seen on the tarsi of the codling moth (fig. 5). li these pores are the only chemoreceptors on the tarsi, it is not con- ceivable how they can detect differences between liquids except by the odors which might be emitted. If the tarsi of butterflies, which are covered with a thick and hard chitin, contain contact chemo- receptors, the mouth parts of insects in general should be provided with such receptors. In other series of tests Minnich (61) repeated his former ones and obtained similar results. According to his scheme of measurement, the total response of all the butterflies tested was Io0o per cent to the sugar solution used, 84.7 per cent to the quinine solution, and 51.6 per cent to the salt solution. In his third report on this subject Minnich (62) says that the tarsal sensitivity of the butterflies tested to sugar solution may be as much as 256 times that of the human tongue. It is scarcely conceivable, although his carefully planned and admirably controlled experiments firmly convinced him that the feet of butterflies contain sense organs, which, when properly stimulated, are 256 times as sensitive as are the taste organs in our mouths. A fourth paper on this subject by Minnich (65) deals with three species of flies. It was similarly determined that these flies can dis- - tinguish water from paraffin oil, or from sugar solution, by use of the chemoreceptors in the tarsi. Chemical sense organs were also located in parts of the proboscis. These organs are more sensitive than those in the tarsi to sugar solution. Minnich believes that all of these receptors serve as taste organs. Thus, according to these results, taste organs, at last, seem to have.been located on the mouth parts of insects. In regard to the so-called taste organs of insects, the writer has repeatedly stated that no one has demonstrated that they actually receive taste stimuli. Minnich (66) says that the proboscis of a certain blowfly is clothed with hairs, some of which are long and curved, and that these have been proven to be taste organs by the following test: A fly, abundantly supplied with water but otherwise starved, does not extend its proboscis when these hairs are touched with a tiny brush wet with distilled water ; but when they are touched NO. 10 TROPISMS OF LEPIDOPTERA—McINDOO 39 with another brush wet with sugar solution the proboscis is quickly extended. These hairs are so sensitive that a single one, when touched, may produce the response. According to the accepted definition of taste, these hairs are true taste organs provided the sugar solution must actually touch them; if only a close proximity is required, then the sense of smell is involved. When asked this question, Minnich was not sure that he had totally eliminated smell. If these hairs are true taste organs, the present writer cannot understand how an aque- ous solution can pass instantaneously through their walls in order to stimulate the nerves inside. III. AUDIRECEPTORS The common belief that insects can hear is based on three facts: (1) Many of the experimental results obtained indicate that they can perceive sound stimuli, although perhaps they do not hear as we do; (2) many have special sound-producing organs; and (3) many have so-called auditory organs. The first report on the auditory sense of Lepidoptera was probably made in 1876. Since that date much has been published, but critics are still inclined to doubt whether any insect can really hear. Turner (86) and Turner and Schwarz (87) in 1914 produced good experimental evidence to show that Catocala and giant silkworm moths really hear. They used an adjustable organ pipe, an adjustable pitch pipe, and a Galton whistle. Their field experiments demonstrated that most of the moths tested can hear high-pitched notes, but usually low-pitched ones did not produce responses. They believe that re- sponses of moths to sounds are expressions of emotion and that a response depends upon whether the sound has a life significance to the insect tested. For many years it has been known that both adult and larval Lepi- doptera are able to produce sounds and some of the sound-producing organs have been described. For example, the death’s-head moths (Acherontia) make shrill chirping sounds, probably by forcing air through certain parts of the anatomy. Their larvae produce “ crack- ling ” notes. A hissing noise is made by several species of Vanessa and more pronounced sounds are produced by other Lepidoptera. Stridu- lating organs on the wings have been described by several, including Hampson (28) and Jordan (35). In certain Agaristidae and Geo- metridae the sound is made by pressing the tarsi against the ribbed areas on the wings. This subject is reviewed by Schroder (80, pp. 61- 74) and Hering (32, pp. 190-193). 4o SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 8&1 Thorax Fic, 13—So-called auditory organs of Lepidoptera. A, Diagram from longi- tudinal section through portions of thorax and abdomen of a noctuid moth, showing following parts of tympanic organ: Gi, Tympanic pit at whose base is found the drum head (7) and G2, tympanic cavity with drum head (G7). The two tympanic cavities are very deep, dorsolateral invaginations of the in- tegument which touch one another at the median line where they form a com- mon division wall (MWS). In Catocala they do not touch one another. Tb, Tympanic chamber; N, tympanic nerve; Ch, chordotonal bundle; L, ligament of same; Td, tympanic cover; and Lm, chitinous lamella, separating the true drum head (7) from the other drum head (G7) and serving for the insertion of the chordotonal bundle (after Egger (17) ). B, Chordotonal bundle of a notodontid (Phalera bucephala L.), showing drum head (7), ligament (L) of chordotonal bundle, tympanic nerve (N), sense cell (Sc), sense rod or “ Stift”’ (Sr), and cap cell (Cc), (after Egger (15)). C, Portion of drawing from longitudinal section through base of front wing of Lycaena icarus, showing chordotonal organ (Ch) and “Sinneskuppeln” (P) or olfactory pores. D, Single chordotonal element from front wing of Chimabacche fag, showing sense cell (Sc), vacuole (Va), enveloping cell (/c), axial fiber (Aa), sense rod (Sr), and cap cell (Cc). C and D after Vogel (89). NO. 10 TROPISMS OF LEPIDOPTERA—McINDOO 4I I. TYMPANIC ORGANS According to Eltringham’s (19) review, tympanic organs in Lepi- doptera were first recorded in 1889 in Uraniidae. Since that date several other writers have described these sense organs in Lepidoptera, which are similar in structure and probably in function to those in Orthoptera. As Eggers (15) has presented the most comprehensive paper on this subject, his results are here briefly summarized. In all he examined 150 species of moths and 5 species of butterflies, repre- senting over 40 families. No tympanic organs were found in 39 species of moths and in the five species of butterflies. They were found, however, in various stages of development in the thorax of 95 species and in the abdomen of 16 species of the moths. Thus 71.6 per cent of all had tympanic organs. Judging from this study butterflies and many moths, including Sphingidae, Saturniidae, and Bombycidae, apparently have no tympanic organs, and none was found in the codling moth by the present writer. The location and strueture of the organs found by Eggers are represented by figure 13, A and B. Eggers (17) next determined that the tympanic organs in noctuid moths are auditory in function. Noctuids, when in an excited condi- tion, reacted to different sounds by flying or by raising the wings. They were tested under glass funnels to loud, sharp sounds such as those made by hand clapping, and to soft ones, as the twisting of a glass stopper in a bottle. When the drum heads (fig. 13, A, T) of both of the tympanic organs were destroyed the moths no longer reacted to sounds. When the drum head in one organ was destroyed the moths reacted to sounds in seven-tenths of the cases by flying. Moths with intact tympanic organs but with wings removed reacted to sounds in one-half the cases by running; in the other cases, by quick movements of the leg or antennae. Moths with intact tympanic organs but with antennae removed, reacted to sounds by flying. He concluded that these organs are sound receptors, analogous to the ears of mammals. 2. CHORDOTONAL ORGANS The name chordotonal means a chord, or string, which is sensitive to tones. Graber (26) in 1882 presented the first comprehensive paper on the chordotonal organs, and much of our present information on this subject is based solely on his report. He apparently found these organs in a wide range of adult and larval insects, but he evidently included other sense organs too. Excluding the olfactory pores on insect wings, he did not find chordotonal organs in adult Lepidoptera, —— SSS 42 SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL. 81 but found them in the larvae of the codling moth and of Tortrix scrophulariana. According to the review by Turner and Schwarz (87), chordotonal organs are not found in Myriapoda and Arachnida. They are found, however, in some insects which do not need a sense of hearing. They are well developed in caterpillars, even in those of Tortricidae, which spend their entire larval period inside of fruit. Eggers (16) remarks that chordotonal organs have been found in the first antennal segment (scape) of Apterygota, Orthoptera, and Hemiptera; in the second antennal segment (pedicel) of Neuroptera ; and in the third antennal segment (first segment of funiculus) of Orthoptera. Some of these are called the Johnston organs, which are discussed later. In the bases of lepidopterous wings Vogel (89) distinguished two types of sense receptors—chordotonal organs (fig. 13, C, Ch) and “ Sinneskuppeln ” (P) or olfactory pores. The former (fig. 13, D) seem to be true chordotonal organs, but the present writer did not see them in codling-moth wings or in those of other Lepidoptera. Nothing definite is known about the function of the chordotonal organs, but they are usually considered as sound receptors. Since most of the movements of insects result in rhythms, as pointed out by Eggers, Snodgrass (81) suggests that these organs be regarded as rhythmometers. 3. JOHNSTON ORGANS Tympanic organs, chordotonal organs, and Johnston organs are all chordotonal organs, because each sense element is chordlike in shape and has a sense rod, scolopala, or “ Stift’’ according to the Germans. A tympanic organ is quite different from the other two types owing to the presence of a drum head or tympanum. A chordontonal organ and a Johnston organ usually differ little; if found in the pedicel, it is generally considered the latter ; 1f found elsewhere, it is called the former ; but in many insects both occur in the pedicel. A good review on this subject is by Snodgrass (81). The paper by Eggers (16) is the most comprehensive on this subject. He studied the Johnston organs in the pedicels of most of the insect orders and concluded that they are true “ Stift’’ organs and are common to all insects, including Apterygota. In regard to the antennae of larvae he found them in hemimetabolous forms, but absent in holometabolous ones. Therefore, caterpillars do not have the Johnston organs. In both sexes of the codling moth the present writer found the Johnston organs (fig. 14) to be highly developed, and the sense rod NO. 10 TROPISMS OF LEPIDOPTERA—McINDOO 43 or “ Stift”’ (Sr) is only slightly different from that pictured in the Lepidoptera examined by Eggers. The writer also saw external marks of these organs in many other Lepidoptera. Eggers informs us that their structure is not correlated with that of the tympanic organs. Formerly they were assumed to be auditory in function, but more recently they have been called muscular receptors FIST SEGIENP Second SOgnenr a Fic. 14.—Johnston organs of codling moth, 500. A, Semidiagrammatic drawing, showing one olfactory pore (P) and Johnston organ whose distal end is attached to articular membrane (Am). This membrane consists of three con- centric bands of chitin; two thin and flexible ones (represented by lines) and a thick, rigid, and much wider one (solid black) between them. Therefore, it slightly resembles a drum head and apparently may be vibrated by jars or by movements of the flagellum. B, Detailed structure of a single chordotonal ele- ment drawn from two sections. All parts, except the nuclei of the enveloping cell (Ec) and cap cell (Cc), were distinctly seen. Other authors have seen these nuclei in other Lepidoptera. The terminal fiber (Tf) of each element is fastened at the bottom of a pit (p) which usually lies in the rigid and thick band of the articular membrane. The other abbreviations are the same as those in figure 13, D. or statical-dynamic organs to register the movements of the antennae. Eggers believes that they probably perceive the movements of the articular membrane to which they were attached. These movements are caused by the antennae being used as tactile organs, or by the wind vibrating these appendages. In the males of Culicidae and Chirono- midae, however, they may be special auditory organs. The present writer (52) in 1922 studied the Johnston organs in the honey-bee in 44 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 81 which the articular membrane, to which the sense fibers are attached, resembles the head of a drum. It was then suggested that these organs might receive stimuli from gusts of wind, weak air currents, or from jars, but the most reasonable function considered was that they regis- tered the movements of the flagellum. 4. AUDITORY HAIRS Years ago there was a controversy as to whether spiders possessed auditory hairs. When a.dead spider was put under a microscope and certain musical tones were produced, some of the hairs on the spider were seen to vibrate. This observation alone is no more proof for an auditory sense in spiders than to say that one stringed musical instru- ment can hear another if a certain cord of the first vibrates when a cord of the second is struck. Recently Minnich (64) has revived the subject of auditory hairs and shows definitely that certain hairs are the sound receptors in larvae of the mourning-cloak butterfly. When a test such as was given with a dead spider was repeated, no hairs on a freshly killed larva were seen to respond to the same tones to which larvae normally react. Minnich’s review shows that certain caterpillars in all instars react to a variety of sounds, including those made by slamming a door, clapping the hands, the human voice, a violin, and a shrill whistle, but the earlier observers did not locate the sound receptors. Minnich used sounds produced by the human voice, piano, organ, violin, dish pan, Galton whistle, tone modulator, and tuning forks. The larvae re- sponded to all of these, except the whistle and modulator, usually by throwing the anterior third of the body dorsally or dorsolaterally. The extent of the response to sounds varied with the intensity of the tone. For full-grown larvae the upper limit of response was probably not far from C’” (1,024 complete vibrations per second). Responses were obtained from 32 to 1,024 vibrations per second. Responses to sounds increased greatly with age, being least in the first and greatest in the last two instars. The responsiveness was correlated with the number of body hairs, which were fewest on the first instar and most abundant on the last instar. Responses to ordinary mechanical stimulation de- creased with age, being greatest in the first and least in the last instar. Headless larvae and fragments of bodies responded to sounds, but the auditory hairs were found to lie chiefly on the anterior two-thirds of the insect. These hairs are probably some of the ordinary tactile ones (Sensilla trichodea) studied by Hilton (34), who claimed that most of the body hairs of caterpillars are innervated. Minnich believes that the NO. IO TROPISMS OF LEPIDOPTERA—McINDOO 45 body hairs act as sound receptors for three reasons: (1) Singeing the hairs greatly reduced or abolished the responses ; (2) hairs bearing water droplets or flour did not respond; and (3) during the molting periods when the hairs were disconnected with their nerves there was little or no response. Abbott (2) observed that normal Datana larvae gave definite re- sponses to air currents and sudden jars, but to only two notes—C’ (512 vibrations) and F sharp (728 vibrations)—by elevating the anterior and posterior parts of the body. These notes were made by using a closed pipe with a movable plunger, a piano, and a mandolin. He assured us that he believed the normal larvae actually responded to the foregoing musical instruments for four reasons: (1) They were protected from air currents when tested; (2) they were several feet from the instruments; (3) vibrations from the substratum were eliminated ; and (4) no responses were observed when the body hairs were covered with water or shellac, or when the body surface was anaesthetized with a 2 per cent solution of procain. Since these caterpillars responded to only two notes, which are not experienced in nature, Abbott believed that these responses were not adaptive, but perhaps secondary, resulting from an “ adaptation of certain organs to more significant stimuli.” IV. THIGMORECEPTORS I. TACTILE ORGANS It seems that no one has made a thorough study of the tactile organs of Lepidoptera, but those in certain beetles have been carefully studied. The writer (53) found tactile hairs on the cotton bol! weevil as fol- lows: Sense hairs (Sensilla trichodea), on the head capsule, antennae, mouth parts, thorax, legs, wings, and abdomen; sense bristles (S. chaetica), on nearly the same parts; and sense pegs (S. bast- conica), on the head capsule, mouth parts, and genitalia. Besides these three types Lepidoptera have a fourth, the sense scales (S. squami- formia) ; however, it seems that only the small, narrow scales are innervated, while the large, broad ones (fig. 3, B, Sh) have no nerve connection. If the end pegs (S. styloconica) are really innervated, we should add a fifth type of tactile organs. Sense scales on the wings of Lepidoptera have been described by Guenther (27), Freiling (23), Vogel (88), and Priffer (77). Vogel states that innervated scales are found on the wings of all Lepidoptera, occurring on both sides, mostly on the veins and particularly on the marginal ones, but they may be found also on the basal parts of the 46 SMITHSONIAN MISCELLANEOUS COLLECTIONS voL. 81 wings. Quenther believes that the sense scales are wind tactile organs, used in orientation. With the aid of them Freiling believes that night- flying Lepidoptera in their rapid movements are able to avoid obstacles. In regard to tactile hairs on the codling moth, all parts of the integument were not searched for them and in most cases where found they were identified from external appearances. Most of the tactile hairs on the wings seem to be ordinary sense hairs (fig. 4, A, St), but a few sense scales (Ssq) were seen. On the legs, maxillae, and labial palpi sense hairs (fig. 5, A and D, St) and sense bristles (Sca) are more or less numerous. On the antennae are found numerous sense hairs (fig. 3, St), sense bristles (Sca), and end pegs (Ss). The large non-innervated scales (fig. 3, G, Sh) overlap one another like shingles on a roof and on some segments they cover nearly all the sense organs. The peculiarly shaped pegs found on the maxillae (fig. 5, D, Pg and I) are also to be classified as tactile organs. Some of the tactile hairs on the antennae and mouth parts of codling-moth larvae are shown in figures 9 and Io. V. GEORECEPTORS I. BALANCING ORGANS When an animal responds to gravity a special static or balancing organ is not necessarily involved, but such organs are known in four Phyla—Ccelenterata, Mollusca, Arthropoda, and Vertebrata. Semi- circular canals occur in the vertebrates, while otocysts or statocysts are found in certain medusae, molluscs, and crustaceans. A statocyst may be an open or closed cavity, lined with sense hairs. In the center of the cavity may be one or more concretions of carbonate or phosphate of lime, called otoliths or statoliths. In the shrimp a statocyst is found in a segment of the claw. It is an open sac in which the shrimp places grains of sand. As the animal moves about in all directions, the grains of sand fall against the sense hairs thus enabling the shrimp to keep its equilibrium. A statocyst, therefore, is nothing more than a special touch organ, and the same may be said about the semicircular canals in which the liquid in them takes the place of the statoliths. A good review on this subject is by Dahlgren and Kepner (8, pp. 207- 2rGe), Insects so far as we know do not have organs similar in function to the semicircular canals and statocysts ; nevertheless, they certainly have great balancing powers. The only case in which such organs have been surmised is in the Diptera. The so-called balancers or halteres were formerly considered organs of equilibrium, but flies can fly just as well without them. NO. IO TROPISMS OF LEPIDOPTERA—McINDOO 47 Vom Rath (78) first described a flask-shaped structure in the distal segment of the labial palpus of the cabbage butterfly. The structure is lined with innervated hairs which he considered olfactory in func- tion. He imagined this structure to be a special olfactory organ for detecting the presence of food. This structure, whose shape varies considerably, seems to be common to all Lepidoptera. It was seen in practically all of the specimens examined by the present writer. It is present in the labial palpi (fig. 5, E, Bo) of both sexes of the codling moth, in which it is sac-shaped, opening to the exterior by a wide mouth (fig. 15, A). The innervated hairs (fig. 15, B, Hr), instead of being narrow and hollow as figured by vom Rath, are wide, heavy, and club- Fic. 15.—Sense organ in labial palpus of codling moth. It is probably a static or balancing receptor. A, Diagram of a longitudinal section through terminal segment, showing organ made up of sense hairs (Hr), sense cells (Sc), and a large nerve (N); B, drawing from an oblique section, showing same parts, X 750. shaped. They certainly cannot be olfactory in function. Since their slender bases arise from very delicate chitin, their clubbed ends prob- ably swing in various directions as the insect moves about. This organ reminds the writer of the statocysts, especially those of the shrimp and crayfish, and it probably has a similar function. If it does not contain statoliths, the hairs may operate sufficiently without the use of them. VI. OTHER RECEPTORS Among the general sensations of Lepidoptera might be mentioned those of temperature, humidity, direction, hunger, fear, and pain, but they are probably not connected with special sense receptors. 4 48 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 81 Much experimental work on various temperatures, particularly as control measures, has been done on Lepidoptera, but little or none of it can be discussed from the tropic point of view. The sense of temperature is probably well developed, although in insects, as in our- selves, there are probably no special thermoreceptors. The subhypo- dermal nerve plexus, if present in adult Lepidoptera as found in caterpillars by Hilton (34), could easily perform this function. Humidity, which is closely related to temperature, also has much to do with the behavior of Lepidoptera. Hering (32, p. 201) remarked that butterflies have a barometric sense, because sultriness and low barometric pressure have a characteristic effect on both the adults and larvae. He imagined that some of the antennal organs are the recep- tors. Guenther (27) hazarded the opinion that the “ Sinneskuppeln ” (olfactory pores) were barometric receptors. Pruffer (77) states that his results and those of Patijaud demon- strate that female moths cannot lure the males from long distances, in spite of evidence shown years ago by Forel, Fabre, and others. He says that the females of Saturnia pyri L., as an example, can attract the males from a distance of not over 50 meters. Noel (68) concluded that neither sight nor smell is sufficient to explain the attrac- tion from long distances. As a hypothesis, he suggested that certain insects emit special waves or rays, resembling X-rays, or the Hertzian waves, or even the N-rays of Dr. Blondlot. He firmly believed that these rays, which have not yet been isolated or verified, really exist and that they are used in distant communication. It has also been suggested that the bushy antennae of certain moths support this theory. C. ScENT-PRODUCING ORGANS The study of scent-producing organs follows as a corollary to that of tropisms and tropic receptors. Since chemotaxis is such an impor- tant means of communication among insects, it is probably true that all insects have structures for producing odors. In fact these structures have already been described in numerous species belonging to most of the insect orders. Several years ago the writer (49) reviewed the literature on this subject. A brief summary of that review concerning Lepidoptera follows: Scent scales on the wings constitute the almost universal type of scent-producing organs in male butterflies. Clark (4) has recently reviewed this subject and added much new information. A pair of invaginated sacs located at the ventro-posterior end of the abdomen has been found in certain male butterflies. These sacs are NO. 10 TROPISMS OF LEPIDOPTERA—McINDOO 49 partially lined with scent hairs at the bases of which lie unicellular glands. In a certain female butterfly the same organ is present, but there is also a circle of scalelike scent hairs around the anus. In another female butterfly there is a single invaginated sac, similarly located. In the females of the maracuja butterflies, a pair of styled knobs at the posterior end of the abdomen serves as a scent-producing organ. The most common type of scent organ in male moths is a tuft of scent hairs on the tibiae of the third pair of legs. Occasionally there are also tufts of hair on the tibiae of the first and second pairs of legs. Another common type in certain male moths is a pair of tufts of scalelike scent hairs at the base of the abdomen. In the males of other moths a pair of invaginated sacs, lined with scent hairs, lies in the ventro-posterior end of the abdomen. In the females of certain moths a paired tuft of scent hairs lies near the anus. The scent-producing organ of the female silkworm moth (Bombyx mort) is the most highly developed of any found in a female lepidopteron. This organ is a pair of invaginated and greatly folded sacs in the posterior end of the abdomen. The female attracts the male by evaginating and turning these sacs inside out, thus fully exposing the inside which is moist with an aromatic substance. In all cases where scent hairs are present, each hair is connected with a unicellular gland. The only scent-producing organ found by the writer in codling moths is a pair of invaginated sacs (fig. 16, A) in the ventro-posterior end of the abdomens of males. The mouth of the sac seems to be a long slit along the ventral median line. Muscles (uw), which nearly surround the sac, apparently change the slit into a wide opening, forcing the go scent hairs (H) to the exterior between two abdominal segments. Each hair (fig. 16, B) is long and its base is connected with a single gland cell (Gc) at the anterior end of the sac. In cross section (fig. 16, C) the hairs are round or oblong, are transparent, and have a spongy texture. The outer wall is rough and a pore (~) can occasionally be seen in it. When greatly magnified the gland cells (fig. 16, D) are large and typical for scent-producing organs. Judg- ing from this structure alone male codling moths attract the females by means of emitting odors from an aromatic substance which passes through pores in the scent hairs to the exterior. No one seems to have described a scent-producing organ like the one in the codling moth, but Freiling (23) has described a similar one in a male butterfly (Danais septentrionalis). In this case the mouths of the paired sacs lie on either side of the anus. Most of the oe ee Sas 50 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 81 scent hairs are attached to the anterior portion of the sac. When the sac is evaginated and the tuft of hair is expanded, this organ resembles a cylindrical fan whose contents are turned inside out to form the circular part of the fan. The scent hairs are filled with a matrix substance and the secretion passes through tiny pores in the walls of the hairs to the exterior. Jordan (36) discovered in a number of Notodontidae a flap, which he called a cteniophore. It is movable and partly covers a cavity in lic. 16.—Scent-producing organ of a male codling moth. A, Cross section through tip end of abdomen, showing location of a pair of invaginated sacs, which are evaginated by muscles (Mu) thereby exposing the hairs (H) to the ex- terior, 53; B, longitudinal section through tip end of abdomen, showing muscles (Mu) attached to invaginated sac filled with scent hairs (H) to which are attached unicellular scent glands (Gc), * 53; C, cross section of scent hairs, showing their spongy texture and pores (p) in outer wall, & 500; D, longitudinal section through bases of two scent hairs (H) and their gland cells (Gc), X 500. the pleurum of the fourth abdominal segment. It is a special male apparatus developed in connection with scent organs. He believed that the hind tibia and hind wing were rubbed across the cteniophore to receive an odorous substance, probably from glands in the cavity. A remarkable combination of tympanic organ and cteniophore was earlier discovered by Jordan and more recently pictured by Hering (32, p. 195). In codling moths, a projection, probably a cteniophore, lies on either side of the abdomen of the males, but no cavity is present. ai NO. 10 TROPISMS OF LEPIDOPTERA—McINDOO 51 SUMMARY AND DISCUSSION In order to throw light on the biology of the codling moth, a thorough investigation of the tropisms of this insect was begun in the spring of 1927. Definite results were obtained only by using the larvae. In all 154 larvae, belonging to the two broods at Silver Spring, Md., were tested individually in the laboratory under various conditions. In bright light, although not direct sunshine, larvae of the first instar were weakly photopositive. Certain tests indicated that objects are perceived and located by the senses of smell and sight, and by mere ‘chance. Chance alone seemed to be only 30 per cent efficient ; sight and chance combined, 40 per cent efficient ; whereas smell, sight, and chance combined were 65 per cent effective. Therefore, since larvae of the first instar have photopositive eyes, they remain in the open on apple-tree foliage and search freely for food, apparently not being aided by their senses until within a few millimeters of the food. The larvae were found to be easily repelled by odorous substances, but attracted with difficulty. Larvae of the second, third, and fourth instars were weakly photo- positive to weak light, but indifferent to strong light. Larvae of the fifth instar sometimes acted indifferently to light but generally were weakly photonegative. Larvae of the sixth instar were either weakly or strongly photonegative, the degree depending on their age; and those with blackened ocelli did not respond to light. At cocooning time the larvae were strongly photonegative, geopositive, and thigmoposi- tive, whereas during their earlier instars they either behaved indiffer- ently to light, gravity, and touch, or were photopositive, geonegative, and thigmonegative. Consequently, when the larvae are ready to spin cocoons they avoid bright light as much as possible, usually move toward the ground and hunt for dark and tight places in which to pupate. When bands are placed around the trunks of apple trees to serve as a supplementary control method, we are merely taking ad- vantage of nature’s laws. It therefore seems that so far as tropic responses are concerned the vulnerable period in the life history of codling-moth larvae is brought about by a change in tropisms. It is well known that certain varieties of apples are more susceptible to codling moth injury than are other varieties; why, no one knows, but several factors, including thickness, toughness, and waxiness of apple peel, and the odorousness and acidity of apples, might be con- sidered. Owing to one or more of these factors the larvae probably gain entrance to the more susceptible varieties with less difficulty ; or the female moths perhaps distinguish differences between apple trees, 52 SMITHSONIAN MISCELLANEOUS COLLECTIONS voL. 81 and if so, they probably lay more eggs on the preferred varieties. No attempt was made in the present investigation to determine which was true, but it is certain that the larvae can distinguish apples by smell and touch, and the moths are certainly guided by tropic stimuli to the proper places for depositing eggs. A study of this kind raises more questions than it answers, yet there is no other way to make progress. Not being able to throw light on this question, a thorough study of the morphology of the sense organs of the codling moth and its larvae was made, hoping that a little light might finally be had. Since the moths are nocturnal fliers, their eyes cannot be their chief sensory receptors for locating the proper host plant. As already stated, the eyes of the larvae change slowly from photopositive ones in the first instar to strongly photonegative eyes in the last instar. This change may be caused by a migration of pigment, as found in certain other larvae, and it seems to be in harmony with the habits of these larvae, which spend most of their time inside of fruit. Before entering apples, photopositive eyes are needed ; but after emerging for the purpose of pupating, photonegative eyes are required. Two kinds of smelling organs—certain hairs on the antennae, and the pores, called olfactory by the writer—are fully described. It seems doubtful whether these hairs, called pit pegs and end pegs, can serve as olfactory organs owing to their hard covering of chitin. Granting that these hairs are the only olfactory receptors of Lepidoptera, eight of the 34 individuals discussed in table 2 cannot smell at all, while four others can smell only slightly. The codling moth, however, has a good supply of them. Larvae do not have these so-called olfactory organs, yet they can smell. The olfactory pores are common to both adult Lepidoptera and their larvae. In the adult they are found on the wings, legs, mouth parts, and second segment of the antennae. In the larvae they occur on the head, mouth parts, antennae, legs, first thoracic segment, and anal prolegs. There are supposedly two types of taste organs. The first type consists of certain hairs on the mouth parts, but since these are covered with hard chitin the writer does not believe that aqueous liquids can pass quickly through them in order to stimulate the nerves inside. The second type is Minnich’s tarsal chemoreceptors, which, when properly stimulated, are 256 times as sensitive as are the taste organs in the human mouth. We know nothing about the structure of these recep- tors, and the present writer so far has found only two kinds of sense organs—sense hairs and olfactory pores—in the tarsi of insects. NO. I0 TROPISMS OF LEPIDOPTERA—McINDOO 53 We now have good evidence that both adult Lepidoptera and their larvae can hear, although probably not as we do. Four kinds of so- called auditory organs have been described. They are tympanic organs, chordotonal organs, Johnston organs, and auditory hairs. The first three have been found in adult Lepidoptera, while the second and fourth occur in caterpillars. Of these four the writer found only the Johnston organs in the adult codling moth, but Graber in 1882 saw chordotonal organs in the codling-moth larva. It has been shown experimentally that tympanic organs and auditory hairs are affected by sound waves, but we know nothing definite about the functions of the chordotonal and Johnston organs. Other special sensory receptors of the codling moth include certain innervated hairs serviceable as tactile organs and a well-developed structure in the labial palpus. The latter might function as a balancing organ. The general senses to temperature, humidity, etc., are not supposedly connected with special sense organs, although these senses seem to be well developed in Lepidoptera. In connection with the olfactory organs the scent-producing organs were studied. The only one found in the codling moth is a pair of invaginated sacs in the abdomen of males; thus it seems that the males attract the females and not the reverse. In conclusion it has been shown that considerable information is now available on the tropisms and sense organs of Lepidoptera, but there is much yet to be learned, and the problem should be attacked from all angles, using the best equipment obtainable. A recent review by Kennedy (38) helps to clarify certain phases of insect behavior. He remarks that while sensitivity is a function of the nervous system, it is conditioned by other structural features, such as small size and chitinous exoskeleton. 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The Behavior Mono- graphs 1 (2): 1V +90, 13 figs. (92) YETTER, W. P., Jr. 1925. Codling moth work in Mesa County. 16th Ann. Rep. Coll. Sta. for 1924, Circ. 47:32-40. (93) Yotuers, M. A. 1927. Summary of three years’ tests of trap baits for capturing the codling moth. Journ. Econ. Ent. 20:567-575, 1 fig. HSONIAN MISCELLANEOUS COLLECTIONS a ~ VOLUME 81, NUMBER 11 Hodgkins Fund "PHENOMENA BY FREDERICK E. FOWLE Oreos. .(PpBiicatioy (3014) CITY OF WASHINGTON _ PUBLISHED BY THE SMITHSONIAN INSTITUTION MARGH 18, 1929 ‘ ~~ re le ‘ io . 4 { } ’ 4 ay aay : l v ~ t J 7 ¢ 7 4, 4 i bate > 4 4 ‘ 7 7 > ‘ ) ean A thu f > 7 , } Coy a (78 } ‘ ” “ ’ ‘ J | ‘ 4 f itis . 2 fo Re t on ; : ) t ‘ 3 ‘ ee oat « 1 "= 7h 4 . Z “ ~ vy : A 4 wi ty, * r¥ 2 ‘ - } . ¢ my 4 \ a | yw . Tie ae . ‘ . ~ » " ‘ t “4 ‘ ea wey ' 1 . j ve te 4 } C/ yh } f ' ¥ S wlPy ; ' 7 A, x \ 1 i¢ . j mF “ ' / , \ : ‘ a4 t ‘ f h i ' 1 ‘ 4 i a / {4 ‘ iy \ ° \ 7 ye. ‘ ‘ ‘ i : ‘ . ' r ed } j \ { { y ' * { , ‘ ’ fi \ y ‘ ’ 1 - * 1 . - > 9 ; Lif { ‘ , ‘ y Sal a ly . a ft "Ta, ‘-e std T SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLUME 81, NUMBER 11 Hodgkins Fund ATMOSPHERIC OZONE: ITS RELATION TO SOME SOLAR AND TERRESTRIAL PHENOMENA BY FREDERICK E. FOWLE (PUBLICATION 3014) CITY OF WASHINGTON PUBLISHED BY THE SMITHSONIAN INSTITUTION MARCH 18, 1929 BALTIMORE, MD., U. &. A, § » & 2 : é @ ae g a 3 ee ee ee er Hodgkins Fund ATMOSPHERIC OZONE: ITS RELATION TO SOME SOLAR AND TERRESTRIAL PHENOMENA By FREDERICK E. FOWLE* The reduction of the measurements of the output of radiation from the sun obtained at the Smithsonian station on Table Mountain, California (altitude 2,300 m.), encountered some difficulty which did not seem to be present at the station at Montezuma, Chile (altitude 2,900 m.), in the southern hemisphere. Preliminary reductions showed the presence of a direct relationship between the values obtained at Table Mountain for the radiation from the sun and the amount of ozone above that station. A yearly march present in the Table Moun- tain solar results, together with other irregularities, were eliminated when proper allowance was made for the amount of ozone above that station. That ozone plays an important part in the interception of radiation coming to us from the sun, especially at the violet end of the spectrum, has been known for some time. It exerts absorption in the following places in the spectrum: * (1) A very strong band in the ultra-violet, 0.2300 to 0.3100p, with its maximum at 0.2550» (the Hartley band). (2) A complicated group, extending roughly from 0.3100 to 0.3500 (the Huggins band). (3) A group in the yellow and red, 0.4500 to 0.6500 (the Chappuis band). (4) A band in the infra-red between 9 and IIp. *A preliminary report of this research was read at the oth annual meeting of the American Geophysical Union, April, 1928 (Ozone in the Northern and Southern Hemispheres, Journ. Terr. Magn. and Atm. Electr. 33, 151, 1928). ? Adapted, with alterations in the wave-lengths of the infra-red band, from “The absorption of radiation in the upper atmosphere,” C. Fabry, Proc. Phys. Soc. 39, I, 1920. SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 81, No. 11 2 SMITHSONIAN MISCELLANEOUS COLLECTIONS voL. 81 The longer wave-length portion of the Hartley band (1) has been used by Fabry and Buisson" and others to measure the amount of ozone in the atmosphere. On June 7, 1920, they found an equivalent layer of a little more than 3 mm. at normal temperature and pressure (ntp). They estimated that at 0.2800n the ozone absorption would reduce the incident solar energy to 107° of its entering value. Ozone, therefore, by its absorption in this band, limits the solar spectrum at its violet end as observable at the surface of the earth. Dr. Dobson * uses this band for measures both of the amount and the Mount WiItS0N IQIO-191\ Zz S rr o c ° w oO xt = c uw = a vn ° = &E < 0.5623, A129 427134 SITE. Fic. 1.—Atmospheric absorption coefficients showing ozone band (Fowle). height of atmospheric ozone. He found a height of 30 to 4o km. above sea-level. The Huggins band (2) was used by Cabannes and Dufay ®* for measures of the altitude of the ozone layer by light reflected from the zenith at the time of the setting sun. They found an altitude of 40 to 50 km. above the earth’s surface. The Chappuis band (3) is used in the present research. The band in the infra-red (4) is of importance because of its location at a wave-length where otherwise the atmosphere would be nearly trans- “Journ: de Phys. 2, 107, 1021. “Proc. Roy. Soc. 110A, 660, 1926; 420A, 251, 1928. *Journ. de Phys. et le Rad. 8, 125, 1927. NO. II ATMOSPHERIC OZONE—FOWLE Ge parent to radiation out-going from the earth. It was observed in the laboratory by Ladenburg and Lehman,' and by the writer in the solar spectrum.” A set of atmospheric transmission coefficients, freed as carefully as was possible from the effects of non-selective absorptions due to water vapor, dry dust, and particles associated with water vapor and called wet dust, was published by the writer in earlier papers.’ The observations are shown in figure 1, redrawn from Fabry’s article (loc. cit.). Cabannes and Dufay* used this data to show that the Xx I- 1,10 X* OZONE in cm. stp. Fic. 2—Atmospheric absorption in Chappuis yellow ozone band (Colange). departures from the straight line of the points at wave-lengths greater than 0.472Qu were caused by ozone present in the atmosphere. Mak- ing the assumption that the atmospheric ozone amounts to 0.32 cm. utp., they used the differences of ordinates between the observed points and the straight line, in the region of figure 1, just indicated, to calculate values of the absorption coefficients of ozone for a standard depth of 1 cm. ntp. Figure 2 shows the 7 resulting values plotted as circles and also a curve showing transmission coefficients * Ann. d. Phys. 21, 305, 1900. “Smithsonian Misc. Coll. 68, I, 1917. * Astrophys. Journ. 38, 392, 1913; 40, 435, 1014. * Journ. de Phys. et le Rad. Sept. 1926. 4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 8&1 for 1 cm. ozone as obtained in the laboratory by Colange.’ The agreement is remarkable. The layer of ozone used by Colange was 18 cm. ntp., and from this the above curve was computed for I cm. ntp., by Bouguer’s formula. The same authors, using a somewhat similar process, later utilized published observations, made by Smithsonian observers at their various stations, for further determinations of the ozone above these stations. These data had not been corrected for water vapor; also the values were taken from somewhat smoothed curves drawn through the plotted observed points. Further, because of gradually progressive changes in the transparency of the sky, comparatively few days furnish observations which are good enough for the above treatment. On these several accounts the investigation just cited is not fully satis- factory. In the following discussion only the original observations are used and they are treated by a method probably nearly independent of sky changes. The results presently to be considered are to a considerable extent a by-product of spectro-bolometric observations originally made tor the determination of the radiation emitted from the sun. Values from about 1,000 days have been utilized. In the ordinary reductions of this work, the ordinates of the solar energy curves (generally 6 curves per day) obtained with a 60° u. v. glass prism had already been read for about half of the days used. It has been the custom to read them on our plates at abscissae, among others, of 18, 20, 22, 24, 26, 28, and 30 cm. towards the violet from the infra-red band, o,, at 2u. These places correspond to wave-lengths of 0.764, 0.686, 0.624, 0.574, 0.535, 0.503, and 0.475, respectively. This spectrum region includes the yellow Chappuis band due to ozone. A preliminary futile attempt was made to use these ordinates to determine directly the depth of the ozone band. The band is masked by the numerous solar lines in that part of the spectrum. Indeed Fabry says: ** The Chappuis bands have never been observed directly in the solar spectrum. I have often looked for them in the spectrum of the setting sun, but have never found them.” However, if the several observations of any day, made at each place in the spectrum at different zenith distances, are used to deter- mine atmospheric transmission coefficients, and the resulting values are plotted against the corresponding deviations, the band is strongly * Journ. de Phys. et le Rad. 8, 257, 1927. * Journ. de Phys. et le Rad. 7, 257, 1926; 8, 353, 1927. as NO. II ATMOSPHERIC OZONE—FOWLE cn brought out as may be noted in figure 3. This figure shows results for days of great, medium, and negligible absorption in this band. The abscissae are prismatic deviations, the ordinates, atmospheric transmission coefficients for zenith sun. As the quantity of atmos- pheric ozone may be correlated to the amount of energy cut out by this band from the radiation coming to us from the sun, the area of 30 28 26 24 22 20 30°28 26 24 22 20 DEVIATIONS PRISMATIC so So > co No) w ATMOSPHERIC TRANSMISSION ZENITH SUN. \o Oo Mean Yearuy 88 SunSpot No 2 7; MariS(TableMt) —'Fes28 (Montezuma) // May 15 (Harqua Hala) J 1928 i 1925 / 1921 * 301-28 26024 22 Fic. 3—The Chappuis yellow ozone band. this band, reduced to the proper energy units, has been utilized as a measure of the amount of ozone in the atmosphere. A smooth curve is first drawn over the top of the band as indicated in figure 3. At any particular abscissa, let a, represent the ordinate on the smooth curve drawn across the band, or, in other words, the transmission of the air for zenith sun with no ozone present; ad, is the corresponding transmission coefficient for ozone, and a the ob- served transmission. Then @=G20o, Of Oo =O) de: 6 SMITHSONIAN MISCELLANEOUS COLLECTIONS voL. 81 Calling e the corresponding energy at the selected place in the sun’s spectrum, it may be assumed that approximately the amount of energy absorbed from the sun’s rays by ozone is a 2 (2 e) summed for spectrum places 22, 24, and 26. a The accuracy of these measurements, depending, at the greatest, on differences of the order of (0.890—0.860), cannot exceed 1 part in 30, assuming no accidental errors. Further, the measurements extend over times of from one to three hours. It is presumptuous to assume always a negligible change in the amount of ozone during such considerable times. Any change in the general transparency of the sky is probably negligible, since it would affect both the numerator and the denominator of the above expression. It takes only 30 seconds for the run through the part of the spectrum used, so that the time is short to produce differential errors within this band. Because the results presently to be given differ so considerably in magnitude and range from the values of Dr. Dobson and those asso- ciated with him, it has been thought advisable to devote considerable time and study to the indications of the Chappuis band. Is the discrepancy due to the presence of other atmospheric lines within the Chappuis ozone band? A count of the number of atmos- pheric lines, designated as such in St. John’s recent revision of Rowland’s Solar Spectrum Table,’ leads to the following table: \F Wave-length | Z pane Of lines Spectrum | | | | range range atm¢ | HO | Os | 27-209 0.490—. 520m 0 O 0. 25-27 0.520-.555 16 I 0 23-25 0.555—-.600 it 244 43 21-23 0.600—.653 81 104 42 In figure 4 the area of that part of the ozone band under trial corresponding to the region of the first three lines of the above table is plotted against the corresponding precipitable water vapor in the atmosphere ; in figure 5, is similarly plotted that corresponding to the lower line. No connection with water vapor can be certainly inferred from these two plots. What little dependence there seems to be is in the wrong direction; that is, the greater the water vapor, the smaller, on the average, seems to be the area of the band. This apparently inverse effect probably results because the season of greatest water * Carnegie Institution Publications, 396, 19028. Si NWOT TE ATMOSPHERIC OZONE—FOWLE Fic. 4.—Abscissae, ppt. H.O; ordinates Ox; 0.47 to 0.60%. Fic. 5.—Abscissae, ppt. H2O; ordinates O:;-0.60 to 0.661. 8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL. 8&1 vapor comes considerably later in the year than that for the area- maximum of the band, yet before the time for its maximum. A far more detailed study of the transmission coefficients in the region of this band has been made than was possible with the some- what separated measurements in the spectrum made for the solar- radiation work. Plates for two days were reread and coefficients determined for each maximum and each minimum of the solar lines visible in the observed energy curves (fig. 6, curve a). Unfor- tunately, between deviations 20 and 22, and 27 and 28, such a process was impossible because of instrumental contingencies. The resulting coefficients determined independently for the two days of observations are plotted in curves b and c. This is a useful transformation, result- ing, as it does, in a spectrum, b or c, showing only atmospheric lines, from an energy curve like a where the solar lines are domi- nant practically to the exclusion of any indication of atmospheric absorptions. Assuming for the time being the validity of Bouguer’s formula, a further step was taken. Entering figure 2 for the corresponding wave- length with the transmission coefficient determined at place 24 from the curve c of figure 6, the amount of ozone was determined. With this amount of ozone, and the transmission coefficients at all the maxima and minima of the curve in figure 2, an ozone band was computed, using the line across the top of the band in curve c of figure 6 as the basis. The result is plotted in curve d of figure 6. The agreement between c and d is better than could be expected and is indeed remarkable. Apparently because the writer is using a purer spectrum than Colange, the deflections in curves b and c are more marked than in curve d, but the agreement in position is satisfactory. Between deviations 26 and 30, the coefficients are too small to expect any accuracy. It seems therefore highly probable that practically all of this band as observed is due to ozone. The writer, as already stated, prefers to express the results which follow in terms of a quantity fairly directly coming from the observa- tions, namely, the amount of energy cut out from the incoming solar energy by this yellow Chappuis band. These results may be approximately reduced to amounts of ozone (ntp.) by using Bouguer’s formula with the constant determined by Colange (Joc. cit.) as indicated by the following table : lan j | | | | | | | Bandearedemerne sie 30 40 | so | 60 | 70| 80! aa 100 | cal. & 10“ Ozotlet Gene oe: |0.90| .160| .200 .230/ .260| .290 .320| .350/ cm. ntp. ATMOSPHERIC OZONE—FOWLE SHUTTER INSERTED 26 24 ae: PRISMATIC DEVIATIONS Fic. 6. 1O SMITHSONIAN MISCELLANEOUS. COLLECTIONS voL. 81 The use of Bouguer’s formula is unsafe for banded absorptions, except possibly for a very pure spectrum, and as an interpolation formula. Langley* long ago showed its inapplicability in a region where quite different coefficients of absorption occur, and his logic is even more applicable in the present case where these occur in close juxtaposition, and in banded spectra where the resolving power is comparatively poor. Safer substitutes for Bouguer’s formula may be employed. For instance, in estimating atmospheric precipitable water the writer always uses an absorption curve calibrated as far as possible in the laboratory. A curve approximately of the shape Abusorbent thickness Fic. 7. indicated in figure 7 would be expected. Where lines of strong absorption occur alternately with those of high transmission, the curve of figure 7 does not tend to approach a zero value of J with increasing absorbent, but to become horizontal for a finite value of J. Assuming Bouguer’s formula to hold we should have a straight line, tangent to some portion of this curve. In view of the state of affairs indicated in figure 7, we should hesitate to use Bouguer’s formula for computing the amounts of ozone, unless for data requiring very little extrapolation from the amounts of ozone used in the laboratory to determine the constant of the formula. It may be that these con- siderations explain certain discrepancies between Dr. Dobson’s results "Ann. Astrophys. Obsery. Smithsonian Inst. 2, 16, 1908. Se —; NO. II ATMOSPHERIC OZONE—FOWLE it and mine at the same stations. He is working at a spectrum place where the coefficient a in the formula, (= ioc is very large, ranging from about I to 4. He is therefore probably working far down on the nearly horizontal portion of a curve such as is indicated in figure 7 where a large change in ozone makes a comparatively small change in the observed spectrum intensity values. On the other hand, in the Chappuis band used by the writer, the coefficient a is so small, about 0.04, that the band is very difficult to observe visually. Therefore we may assume that the writer is measur- ing in a band where a small change in ozone produces a great change in the observed quantity. In other words, for the amount of ozone present in the atmosphere, the Chappuis band is a more sensitive indicator of changes in atmospheric ozone than that employed by Dr. Dobson. ' With these preliminary remarks, attention may be drawn to figure 8, in which recent observations made at Table Mountain with Dobson's apparatus, and reduced by him to cm. ozone ntp. are compared with the writer’s results as expressed in areas of the Chappuis band. The average amount of ozone for this interval of time as computed by the preceding table from the writer’s results is about 0.23 cm. ntp., while Dobson finds about 0.22 cm. The range of the variation found by the writer much exceeds that found by Dobson, but nevertheless a marked correlation exists between the two series. The writer cannot leave Dr. Dobson’s work without one further remark about his method. He states,’ “ It has been shown that there is a close connection between the amount of ozone in the upper atmosphere and the pressure conditions in the upper part of the troposphere and the lower part of the atmosphere,” and states that. “it is remarkable that the ozone situated at so great a height ” (40 to 50 km., as indicated by the results of Cabannes and Dufay, 30 to 40 km. by Dobson himself) should be so closely connected with variations of pressure much lower down.” Dr. Dobson” uses two methods in his evaluation of the amount of atmospheric ozone. In the first he takes as the general atmospheric transmission coefficient 1 Proc. Roy. Soc. 120A, 251, 1928. * Mo: Not. R: A. S. 86, 259, 1926. Proc. Roy. Soc. r10A, 660, 1926. VOL. 8&1 LLANEOUS COLLECTIONS SMITHSONIAN MISCE ‘ULeJUNOIY 9qVy OF s dnfeBA S to}tUM 94} “9AIND JIMOT “UleyUNOPY eGR L 1OF Sonypea Ss uosqod ‘qd ‘SAINI paqyjop taddq— ‘OLY XRD _olx|e2g Yer ATMOSPHERIC OZONE—FOWLE I XE Arnica A 1928 1924 ' i ' ' ' 1 1 1 ' ' 1 1 t ' ’ ' ' an Fely Mar Apr May Jun Jul Aug, Sep Oct Nov Dec—O-Jan Fels Mar Apr May Jun Jul Aug Sep Oct Nov Dec 5 Herqua Hala, Arizona. © Montezuma.Chile. 2 Table Mt.California. %& Mt Brukkaros, Africa Fic. 9. 14 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 81 where K=B+8+ar B is the absorption coefficient due to small particles, 8 is the absorption coefficient due to large particles, a is the absorption coefficient due to I cm. ozone ntp., wv is the thickness of ozone atmospheric in cm. ntp. Now it seems to the writer that the very variations with atmos- pheric pressure which Dr. Dobson throws into #, belong fully as OZONE: N. HEMISPHERE S15 SUN SPOTS 7 ”Y ft oO ae YY z =) WY a— 1921 922 923: 1924" 1925, 1926: 1927 ai9Zzs EIgs te: legitimately, and very probably, to both B and 6. All this relates to what Dr. Dobson calls his “long method,” dependent upon several observations during the day. In his second or * short method” he uses the expression _ (log I,—log I’) — og J —log I’) — (B—B’) sec 2 al (a—a’) sec 2 In the determinations by this “short method” he assumes that 6 does not vary with A and uses a value for 8 obtained from the formula NO, 21 ATMOSPHERIC OZONE—-FOWLE 15 of Rayleigh. Although both these assumptions may be allowable up to a certain accuracy it seems likely that from either of them a variation dependent upon the atmospheric pressure or water vapor may have been introduced. Let us now turn to the results of observations made at Harqua Hala (altitude 1,770 m.) and Table Mountain (2,300 m.) in the United States of America, Montezuma (2,900 m.) in Chile, and Mt. Brukkaros (1,600 m.) in Africa, embodied in the following table and figures 9, 10, and 11. The table gives only the monthly and Qua =o SUN SPOTS 3 O | 3 4 ErGyats yearly means; hence the plotted points, especially in the plots of yearly means, figures Io and 11, depend upon a considerable number of day’s observations but not always every successive day. The Wolfer spot numbers and the magnetic character values here given are computed employing only the days of radiation observations. In my preliminary paper, already referred to, the plots related to daily values, and even with the few values there utilized from the 1926 and 1927 observa- tions at Table Mountain, showed a distinct correlation between the ozone, the spot numbers, the magnetic character and the flocculi for the corresponding days. 16 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 81 THE OBSERVATIONS “Harqua Hala, 1921 | Harqua Hala, 1922 end | Se eee ae Month | Ne | Ozone | Walier | saan, | ey ee Pe 5 ed Jan, 1 © | 420) 627 0.6 Jan. 8 | 34 | II 0.7 i Feb. 14 ATS \ln3t aA Feb. 5 47 23 T.0 ; Mar. ) a | 64 5I 8 Mar. 5 5On esa | -Y ; Apr. 7 58 | 16 1.0 Apr. 5 56 ES) 0 : May _ 3 | 34 | 45 1.t || May 7 55 Sah paz . June G 52 27 4. | June | 7 Fo 1 aS ) July T5400) 100 5 | July 4 | 52 18) A Aug. A> | 242 20 9 | Aug. 2 49-1 irr 2 Sept. 13 4418 -3 || Sept. 5 AG a eG 8 , Oct. It {| 542 17 oa Oct 4 34 | 4 8 } Noy. ese Ge 18 6 Nov. 3 27,5)\ een aE Dec. Ae FG 14 23. Nl Dec. 4 31) [ato 12 Year | 89 | 48 27° || 0:6 Year| 57 44 17 0.7 | Harqua Hala, 1923 | Montezuma, 1923 em eter) ate | es | ee eae ee Taree | ue | ae | | | | | Jan. 6 21 | 5 | 0.4 | Jan. | 2 24 10 || O40 Feb, 3 23 i2ea | esa Bebe | |. 934: 2 6 Mar. 4 1 8) 7 Mare llearG wel eee Bail) ae Apr. 4 | 48 } 7 -6 | Apr. | 12 | 28 Tih) eae May 5.2 S2Hg me ‘Sf May | 7 | 3 37 Es June Br ail eas TOMM ees June) ra | 36 13 4 July Le wor 0 4 daly) 60 | ae 2 | ..4 Aug. 3 ar Sant es Aug. Apa ine 5 2h lesen Sept. 2 37 To’ | 38 Sept. 5 le an 10 | 4 Oct. 3 27 12 -4 | Oct. | 4 | 36 Saad ike Nov. J SEZ ie ah al Owes tag) gill aee 9 8 Dec. ZR i Ba ts | «I | Dec. | eB ) 79 o |. 6 Year | 37 | 34 ! 6 | 0.5 || Year | 73 | 32 7 | 0.6 | | NO. II ATMOSPHERIC OZONE—FOWLE 17 Harqua Hala, 1924 ; Montara: 1924 Bec cee ee | ae | meen [Re eee ate ee | ah Va | | Jan. 4 | 48 2 0.9 Jan. 4 35 oO | Ons Feb. 3 49 7 .2 || Feb. fs - | Peay ilo ee Mar. 2 28 0 a Mar. 4 3021 ie | 8 Apr. I 61 0 s2 Apr. 4 32 10 ag May 3 26 14 8 May Ae 2 22 7 June | 7 39 18 a7 June 4 34 22 4 way | 4 | 2.30 9 8 | July 4 40 25 6 Aug. I | 22 II sit Aug. 5 40 II ar Sept. Be eects 19 6 Sept. | 4 33 22 a5 Oct. Sas 25 26, WeOcts | 4 40 26 6 Nov. Saws 31 a7 Nove 4 Se ee 2O 22 6 Dec. | Dec. | 14 25 2I | 4 Year 36 | ATs eats i On6 Year 56 33 v7, 0.5 eee | Montezuma, 1925 : Table Mountain, 1926 7 Breer ee | atte | Mae accany (NE [Cee ilies ae Pe en ee Jan. | 10 | 20 6 | Os. wang: leueo | 88 | 69 | 0.9 Rep | 3) |) 28 TS) eons Feb. II 84 | 68 | Rag Mar} 017° | 42 22 .4 || Mar. 15 84 50 fe Apres | ors 44 29 -5 || Apr. 14 90 38 8 Mayr |) -13 | 44 38 4 May | 24 90 68 .6 June 14 | 49 24 .8 || June | 18 Sra ea 5 July ees | 37 3, | Jalyc olmease emia ie ae 5 Aug. 2 | 42 19 8 Aug. 22 88 | 62 a5 sept |) 4 48 63 9 Sept 28 67 | 62 LG Oct. His | 38 43 a7 Oct. | 24 62 | 66 6 Nov. | 2 hes 54 8) Nove on enGs | 50 ig Dec. Zan a36 76 4 Deere] 7 Zn 80 4 4 Sr ee ere aes, ol emcee Year 90 42 20 | os || Year | 211 8 | 62 | 07 | | 18 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 81 Montezuma, 1926 Table Mountain, 1927 Month | Ne | eens | Waltes | Bien, | sfocn | de | zane | Magn. | wotter | | | | | Jan. I 34 | 84 | 0.7 Jan 2 |, 468 72 0.4 Debs! eat To daloatgot Me eou Feb 4 | 72 80 8 Mar, | 2 BOPP esa) 14 cg ean Nie! 4 78 | 89 af Apr. 4 28 | 43:9) (ao | Apr. 5 76 87 8 May 2 27°) 38 6 May 5 80 | 79 4 June oe all -. | June 4 |} 79 65 a3 July Sy 87 | 34 6 July 3. 463 40 :1 Aug. 2 | 42 61 4 | Aug 12 W107 a ex 8 Sept. a ca | ++ || Sept 6 | 66 | 60 8 Oct. BF alien ar | 82 |) 1.2 Oct. 5. | "66 | 60 a7, Nov. 2 | 40 73 5 Noy 6 50 | 66 4 Dec. Sept cere i | | Dec 2 62 aio T:0 Mone eee ee Year 20 33: | “so 4! 028 Wear | 58 69 ~—s 65 0.6 es | Montezuma, 1927 Ail Table Mountain, 1928 ee vl] ES | } Wolfer | Magn. Mom obs | Site® [ERT MY" atom | Mo | Ozone | Wait tte Jan. I 39° | 77 ae Jan. | 16 76 70 Feb. I 43 82 sO “eb. II 87° | Fo Mar. I 46 131 oe Mar. 19 92" | 703 Apr. 4 35 86 1-2. |) Apr. I2 72 83 May 3 45 03 «2 May 20 LOIN 738 June 3 36 86 -I |} June 25 98 =o July 3 36 52 3 | July 20 77 | 102 Aug. “3 44 44 8 | Aug. 31 67 | 83 Sept. 2 38 90 6 || Sept. 18 60 | 79 Oct. 2 44 68 2.0 | Oct. 18 OOM Bie rts Nov. 8 45 61 = 19 Noy. 0). ora 67 Dec. 2 24 65 a Dec. | Year 28 38 75 0.7 || Year | 204! 81 85 | Ee NOT LT ATMOSPHERIC OZONE—FOWLE 19 From the data of the preceding table and the corresponding figures several phenomena are notable: (1) There is a very decided yearly march, as has been noted by other observers. (a) In the northern hemisphere we may take the maximum and minimum of this march as follows: Maximum Minimum 1921 March’ Sept. 1922 March Nov. 1923 April-May Aug. »? 1924 April Aug. ? 1925 (April, Dobson) (Oct., Dobson) 1926 April-May Oci: 1927 April-May (April, Buisson) Nov. ( Nov., Buisson)* 1928 May Sept. (b) and in the southern hemisphere as follows: Maximum Minimum 1923 Sept. March 1924 Aug.-Sept. March 1925 ? Feb. 1926 Aug. April ? 1927 not definite not definite. whence: (2) In the yearly march the maxima and minima occur at nearly the same seasons of the year in the northern and southern hemi- spheres, though of course not in months of the same name. The maxima occur between April and May, the minima between August and November in the northern hemisphere and vice versa approxi- mately in the southern. (3) A marked correlation exists between the ozone and the Wolfer sun-spot numbers for the observations of the northern-hemisphere stations, as indicated in figures 10 and 11. The range of the yearly means for the area of the yellow band is from 20 to 100 (see fig. 9). ‘The writer is inclined to discount the appearance of the low value in May, 1921, as abnormal, possibly due to erroneous observing, and to consider the general march of the curve as indicating the minimum in September. Somewhat similar judgments occur later in the table. Gh ene O.0220, 1OLS: 20 SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL. 8&1 (4) In the southern hemisphere no such strong correlation is appar- ent between the spot numbers and the ozone. The corresponding range is only from 20 to 30. However the errors of the readings of the area when this is small are comparatively great; indeed the observations do hint a slight relationship. The writer suggests that the following considerations point to a fifth deduction from the observations. The ozone present in the upper air has been generally considered as formed from the oxygen there present by the action of ultra-violet light from the sun. Radiation of very short wave-length (less than 0.1850) acts upon oxygen, transforming it into ozone. It is not improbable that radiation of this wave-length reaches the earth from the sun. If so, it must produce ozone in the earth’s atmosphere, but only in the highest levels, because it cannot reach the lower strata. Radiation of wave-length 0.1850p is completely absorbed by 10 m. of air at ntp., and could scarcely penetrate lower than a stratum 4o km. above the earth. On the other hand, radiation lying between 0.2000 and 0.3000%2 decomposes ozone, and between these two opposite actions a state of equilibrium would be established. Since the ozone- destroying wave lengths penetrate deeper into the atmosphere, this naturally limits the ozone layer to a high altitude. It is possible though that another agency than ultra-violet light works to produce ozone. The investigations of Milne’ and Pike’ indicate the great probability that electrified particles gain such velocities on the sun that they are projected outwards into space from that body. Mme. Curie * has shown that the a-particles emitted from radium salts ozonize oxygen. Electrons with a velocity of 1.80 x 10% cm./sec.’ are capable of producing ozone from oxygen. It is also produced by the silent electrical discharge. Suppose then that there are two causes at work producing the ozone of the earth’s atmosphere: One portion may then be due to the ultra- violet light from the sun, and present over both hemispheres; the other, caused by particles emitted from the sun of such a polarity that, when they reach the earth’s field, they drift towards the northern hemisphere, above which alone would the ozone due to this last cause be abundantly present. The particles would then necessarily have a positive charge, ¢.g., a-particles. *Mo. Not. R. A. S. 86, 259, 1926. * Mo. Not. R. A. S. 88, 3, 1927. 7 Gapke ens se “Franck and Hertz, Verh. Deutsch. Phys. Ges. 15, 34, 1913. | INO. TI ATMOSPHERIC OZONE—FOWLE 21 With the assumption of these two sources for the origin of the atmospheric ozone, several of the phenomena shown by the observa- tions of this paper fall in line, and our fifth conclusion will be: (5) (a) Due to the ultra-violet light from the sun, there is a layer of ozone, varying apparently very little with the sun-spot period, and situated over both the northern and southern hemispheres and showing an annual march having its maxi- mum in the spring of both hemispheres and its corresponding minimum in the autumn. (b) There is another layer formed under the bombardment of electrical particles (probably positive ions) emitted from the sun and showing strongly a dependence upon solar activity as indicated by Wolfer’s sun-spot numbers. At the only minimum of spots observed this layer appeared practically absent, the measurements indicating the presence of the (a) layer alone. Though the corresponding marches during the year of the ozone (which the writer proposes to attribute to the first of the above causes) occur in different months in the two hemispheres, the seasons of maximum and minimum are the same, namely, spring and autumn. One might be led to suppose that these variations are due to some dependence upon the annual and reciprocal marches in the two hemispheres of the air-masses through which the sun’s rays could penetrate for the formation of ozone. Further at the tropical station at Montezuma the sun is more nearly overhead and the air-mass change smaller, which might perhaps account for the smaller annual range there. However the maxima and minima do not occur at times when the sun is farthest from or nearest to the zenith, when there would be the greatest and least air-masses. Another circumstance might lead to an explanation of the annual march and its reciprocal effect in the two hemispheres so far as concerns the times of occurrence of the maxima and minima. An- nually, as viewed from the earth, the sun’s equator reaches its greatest southern displacement (7.25°) about March 7, and its greatest northern displacement about September 8. The aspect of the sun’s disk as seen from the earth at these epochs is shown in figure 12. Since the earth subtends only about 30’, as seen from the sun, under either circumstance, the sun would have practically the same’ aspect as viewed by ultra-violet light from either the northern or southern hemispheres of the earth. However, the ultra-violet light would probably be strongly scattered by the particles of the solar corona, and this annual shift of the far more extensive and considerably 22 SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL. 81 more oblate corona might have a differential effect on the intensity of the ultra-violet light reaching the separate hemispheres. Returning again to Dr. Dobson’s results, he finds much the same values in both hemispheres. He now has an observer in New Zealand (1928). He states* that he finds very little connection of his observations with the sun-spot cycle, and that little apparently in a reverse sense from that clearly indicated by the writer’s results. He obtains an altitude for his ozone layer from the Hartley band at 30 to 40 km., whereas Cabannes and Dufay get an altitude from the N point N point Mar./7 Sept.8 Pig! 12: Huggin’s band of 40 to 50 km. One might hazard the suggestion that it is not beyond possibility that the band used by Dr. Dobson corresponds to such a state of the molecule that only ozone formed by ultra-violet light is in the proper molecular state to effect absorp- tion; whereas the band in the yellow is due to a molecular state which measures absorption due to ozone formed by either process. It will presently be seen how such supposition as to two layers of ozone is 1n line with the conclusions drawn from magnetic data relating to two separate strata, the lower of which is assumed to be due to ultra-violet light. ‘ Observatory, 51, 381, 1928. * Proc. Roy. Soc. 114A, 532, 1927. NO.. IT ATMOSPHERIC OZONE—FOWLE 23 Turning to the literature of Terrestrial Magnetism, the writer was both surprised and pleased to find decided support lent by the phenomena of terrestrial magnetism to this hypothesis of two quan- tities of ozone formed by two separate agencies. Furthermore, these two layers were not only ascribed to the same two agencies as already stated, but assumed to be probably separate layers. If we consider magnetically quiet days, we find a similar yearly march in the magnetic elements, the maxima and minima, however, occurring somewhat later, namely, in June and December at Green- wich; and further a regular march with the sun-spot period. This march is so regular as to lead to the inference that it is due to a general change in the whole solar disk accompanying the sun-spot period. Further there occur disturbed days which seem to be connected with specially disturbed conditions localized on the sun’s disk, for they show a definite tendency to recur at successive rotations of the sun’s disk." . “ There are few facts of greater significance,” writes Dr. Chapman, ‘with respect to the relation between magnetic changes and the sun, than the tendency shown by the earth’s magnetic activity to return to its condition at any particular time, aiter the lapse of one or more ‘ periods of synodic rotation of the sun.” There are discordances between the succession of events with the ozone phenomena and those that are magnetic, so that the events may be confused with complications not due to the same cause, but the following discussion by Dr. Chapman (loc. cit.) seemed of special interest : “These conclusions regarding the * disturbance ’ solar agent have a direct bearing on the ‘ general’ solar agent which affects the regular diurnal magnetic variations over the sunlit hemisphere. If the former consists of electrical corpuscles, the latter cannot do so—no mere difference of mass or sign of charge would account for the complete difference of distribution of the two agents reaching the earth. On the other hand, the apparently sole alternative among possible ionizing agents, viz., ultra-violet light, seems to accord with all the properties which the ‘ general’ solar agent has been shown to possess : for the latter affects the sun-lit hemisphere almost exclusively, it arises from the sun’s surface as a whole, and its intensity varies only gradually, from time to time, in correspondence with the general activity of the sun.” Dr. Chapman, Trans. Cambridge Phil. Soc. 22, 341, 1919. 24 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 81 And he later continues, “The facts hitherto reviewed may next be considered in their bearing upon atmospheric questions. One such question is, Are the layers affected by the two kinds of solar emissions the same or different, and if different, what is their relative situation ? “Tven, a priori, it would be expected that two such different emissions as corpuscles and ether waves will have different powers of penetration into the atmosphere, though it would not be possible, on such grounds alone, to decide whether the ‘ absorbing’ layers were wholly distinct or not. The magnetic phenomena, however, give a fairly clear indication that they are practically distinct without over- lapping * * *,” and he reaches the conclusion “that the magnetic disturbance layer is situated at a higher level than the diurnal variation layer.”” He infers from this that the magnetic disturbance layer (due to ions from the sun) is situated between go and 120 km. and the diurnal variation layer (due to ultra-violet light), between Ito and go km. Dr. Chapman has added a note dated July, 1919: “In a paper read (on May 22, 1919) before the Institution of Electrical Engi- neers, and shortly to be published, I have suggested that the ultra- violet radiation * * * may be some type of gamma-radiation, and that the corpuscles are (as Vegard has urged) alpha-particles. If both these processes originate from radio-active processes on the sun, the gamma-rays would be expected to penetrate more deeply into our atmosphere than the alpha-particles.” All of which falls in with the observations and suggestions of the present paper. Lord Rayleigh’ has recently published observations which relate to a phenomenon possibly allied to that of ozone. These observations are measurements of the intensity of the auroral green line in the light of the night sky together with similar measurements of the intensity in the spectrum of the night sky on each side of this line. McLennan~* has shown that this green line owes its existence to a metastable state of the oxygen atom. Whereas the green line is always present in the light of the night sky, the negative bands of nitrogen * Proc. Roy. Soc. r1gA, 11, 1928. * Nature, 122, 38, 1928. NO. II ATMOSPHERIC OZONE—FOWLE 25 which are an important feature of the auroral spectrum, are not usually present. Rayleigh’s observations (fig. 13) apparently indicate an annual march in the intensity of this green line with two maxima—the smaller maximum occurring nearly contemporaneously with the single maximum in the ozone march, the larger with the ozone minimum. In the southern hemisphere, as with ozone, the months of the occur- rence of these maxima are reversed but, of course, not the season. Omitting observations made at Claremont which Rayleigh considers faulty, together with those for some stations with only few observa- Fic. 13.—Lord Rayleigh’s observation on Aurora green line. The gap in the northern hemisphere results is due to the impossibility of observations during these months in England because of twilight. tions the following table was formed. The values given relate to a comparison of night sky observations with a standard source of light. For details the reader is referred to Lord Rayleigh’s article. The scale units are such that in passing up one unit the intensity is multiplied by the anti-logarithm of 0.1 or 1.259; three steps on the scale are equivalent to a factor (1.259)° or approximately a doubled intensity. The presentation is different from that of Rayleigh so as to separate the stations of the northern and southern hemispheres. It indicates that the auroral line averages of greater intensity in the northern hemisphere while the parts of the spectrum on either side show no such difference. 20 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 8&1 Mean intensity Place, aaa ae Time range Cae i eaeenlre me ey Red green Blue Lerwick +60° 1° W 86 Sept. ’25-Apr. ’27|| —28 +06, +5.7 england 52 eee 81 | Nov. ’25-Oct. 26) —2.5 +05 +6.4 Kingston 44. 77 W 6 | Apr. ’26-May ’26) +01) +23; +9.5 Victoria 40 123 Wi 108 || Sept.’25-Mar. ’27]| —3.2| +08! +6.3 Mt. Wilson 24 Tr SW 83 Sept.’25-Feb. ‘27, —29 +11 +68 Hawaii 1OMISON Wel <116 Oct. ’25-Nov. ’26') —3.2] -+0.3)| ---6:5 Kodaikanal NOM 77a 54 Oct. ’25=Jan: °27'| —2:8) —o.1| -56.2 Gilgil O° 37.8 5 Sept. 25 —2.7' +1.5 +6.6 Northern mean . | —2.5! +09! +6.7 Cape —34° 18° E | 199 | Nov. ’25-Nov. ’26 } —2.7, +07 +7.1 Arequipa 16 71 W 31 || Apr. ’26-Nov. 726!) —3.1, —o.3) +6.5 Canberra 35 149 E | 149 | Mar. ‘'25-Oct. ’26| —3.2) +04| +6.5 Christ ‘Church. 44 73° E 56 Feb. ’26-Feb. °27, —1.0;' +0.4|) +7.0 Southern mean —2.5| +0.3| +6.8 Before summarizing the results of this paper, the writer wishes to express his appreciation of the criticisms of Dr. Abbot, and the aid furnished by Miss Margaret Marsden and Mr. Hugh Freeman in the many computations, as well as his indebtedness to the workers in the field whose observations made possible this discussion. SUMMARY The amount of energy absorbed from the incoming solar radiation by the yellow ozone band has been used to measure the variations in the amount of atmospheric ozone during the years from 1921 to 1928. These observations have been made in both the northern and the southern hemispheres. The resulting values show a distinct yearly march in both hemi- spheres. In the northern hemisphere the maxima of this march occur between April and May, the minima between August and November ; in the southern hemisphere the maxima occur between August and September, the minima between April and May. In other words in both hemispheres the maxima occur in the spring, the minima in the autumn. In the northern hemisphere a marked relationship exists between the ozone and the Wolfer sun-spot numbers. The range in the monthly mean values for the ozone numbers is great and between 20x 10% and 100 Xx 10~! calories absorbed per cm.? per minute from the incom- ing solar energy. NO. II ATMOSPHERIC OZONE—FOWLE 27 In the southern hemisphere no such marked relationship is noted, although one may be masked by the small range and corresponding inaccuracy in the values. The range is only from 20XIo0~* to 50 X 10° calories. It is suggested—and such a suggestion is strengthened by magnetic data—that we are dealing with two layers of ozone. The first is due to ultra-violet light coming from the sun and hence existing over all the stations. The second is assumed to be due to positively elec- trified particles emitted from definitely disturbed areas of the sun. This second effect reasonably shows a strong correlation with the Wolfer sun-spot numbers. Probably because these positive particles are deflected towards the earth’s north pole this layer of ozone is found over the northern hemisphere stations only. At sun-spot mini- mum it is negligible so far as the present measurements indicate. All the results of the present paper are based on monthly and yearly means. A consideration of the daily values would be another story. The plot published in the preliminary paper was based on daily values for only two years at Table Mountain. The short study then made of the daily values would indicate that what may be said of the connection between many magnetic values and solar disturbances may be said of ozone; that although with monthly and yearly aver- ages, solar spottedness, for example, goes hand in hand with the amount of ozone, yet a day of many spots may pass with no increase of ozone and vice versa. aie — SS _ . *. n 4 » Sf 4 < « ‘ z ‘ ; ae { ‘ / SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLUME 81, NUMBER 12 ARCHEOLOGICAL INVESTIGATIONS IN THE TAOS VALLEY, NEW MEXICO, DURING 1920 (WitTH 15 PLATES) BY J. A. JEANGON f ‘ ALIN tx 1999 a OFFIC FE CITY OF WASHINGTON PUBLISHED BY THE SMITHSONIAN INSTITUTION JUNE 11, 1929 IBRAY a s: Sete keds wer J 4 7 Pi = ot ee en 7 Seen ie * - ee; 5 See e = 4 1! 116” ee Oe ae hie Itc. 2.—Map of large ruin at Llano. NO. I2 ARCHEOLOGY OF TAOS VALLEY JEANCON II Judging from present appearances there were no openings in the outside lower walls, not even ventilators, such as have been observed in other places on the Jemez Plateau. It is more than probable that the lower outside rooms were supplied with air and light entirely through hatches in the roof. As far as could be determined by the excavation, the old walls did not serve as foundations for the secondary ones. There is practically no difference in the width of the walls of the buildings of the first and second occupations. All of them, old and new, are very irregular, and average from eight to twelve inches in thickness. Where the first walls were established, they ran under and on different lines from the newer ones. This was especially evident on the west side of the kiva plaza, where the original wall extended across the whole west side of the plaza and new ones were built on either side of the old one, the later ones rising from the present ground level while the older one extended for some distance below it. When these walls were Fic. 3.—a, new wall; b, old wall; c, new wall; d, cache between walls; e, showing how spaces between walls were filled to make banquette. completely excavated and swept it was found that the eastern one formed a banquette which had been plastered over the top and outer side (pl. 3, 4). There was an open space between it and the central wall, which was built against the western one. The newer walls were not as well mixed and firm as the central one, which extended south below the room next to the plaza (fig. 3). With the exception of room 7 all of the corners were square. In room 7 the southeast and northwest corners had been rounded off by the building of short walls across the corners. The northwest corner appeared to have been used as a fireplace, as the wall was smoke-stained from the floor to the top of the standing wall. There was no evidence of a chimney or even a hood such as the Zuni and Hopi use. The southeast corner was not filled in solid, and the curv- ing front wall, with the rectangular corner behind it, formed a sort of cupboard which was divided into an upper and lower shelf by a huge, flat, river boulder. The cupboard opening was circular and about one foot above the floor. There were no objects of any kind inside it, nor any smoke stains to suggest its use as a fireplace. I2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 81 The walls of the large mound were built in the same manner as in the smaller mound, the only difference being their marked variation in thickness. ROOMS The rooms varied widely in size (pl. 2, B, C). No complete out- lines of the first occupation were determined, but all of the second occupation rooms were uncovered and their dimensions are given in the following list: No. 1.—16 ft. x 6 ft. 6 ins. No. 10.—13 ft. 6 ins. x 11 ft. 2.—14 ft. 9 ins. x 6 ft. 6 ins. 12.—13 ft. x 7 ft. 3.—I5 ft. 6 ins. x 6 ft. 13.—14 ft. x 8 ft. 4.—I15 ft. 3 ins. x 6 ft. 14.—I5 ft. x 9 ft. 5.—I15 ft. x 6 ft. 4 ins. 15.—II ft. x 7 ft. x 11 ft. g ins. 6.— oft. x 14 ft. 16.—I0 ft. x 9 ft. 9 ins. x II ft. 3 7,—I0 ft. 9 ins. x 9 ft. ins. x 8 ft. 6 ins. 8.—14 ft. x 6 ft. 6 ins. 17.—10 ft. 6 ins. x 9 ft. 9.—1I3 ft..x 5 ft. 6 ins. 18.—11 ft. 6 ins. x Io ft. The kiva plaza was roughly 30 ft. square. The following measurements made in 1864 by Mr. John Ward at the present pueblo of Taos are interesting as showing the increase in size of the rooms more recently constructed : Several rooms on the ground floor were measured by Mr. Ward and found to be in feet: 14 x 18; 20 x 22; 24 x 27; with a high ceiling averaging 7 to 8 feet. In the second story they measured in feet: I4 x 23; 12 x 20; and I5 x 20; with a height of ceiling varying from 7 to 7} feet. The rooms in the third, fourth and fifth stories were found to diminish in size with each story.’ It would be extremely interesting to have an opportunity of exca- vating and measuring rooms in the ruin of old Taos Pueblo, which lies a short distance east of the present village. Perhaps the gradual growth in size of the rooms over the period between the Llano houses and the modern Pueblo, could be obtained in these rooms. However, it would be next to impossible to obtain permission to exca- vate in the Old Pueblo as the Indians will not even permit the mounds to be measured, much less excavated. STORAGE ROOM One of the most interesting rooms in the group was one which had every appearance of having been used for storage purposes. It is situated just south of the kiva, and is called room 12 (pl. 3, B). A raised bench occupied almost three-fourths of the floor space. In ? Morgan, L. H., Houses and House Life of the American Aborigines. Contri- butions to North American Ethnology, Vol. 4, p. 145, Washington, 1881. NO. 12 ARCHEOLOGY OF TAOS VALLEY—JEANCON 13 the south and east ends were buried seven pots, four on the south side and three on the east side (fig. 4). Fic. 4——Ground plan of room 12, showing arrangement of storage jars below the bench level. The tops of these pots came to within two or three inches of the floor ; they may have been used for storing meal. At the western end was a sort of cupboard running under the wall, with an entrance about one foot square and one foot deep projecting into the room. In Nie o (71 12 Fic. 5.—a, showing position of cupboard; b, cupboard with jar in place. this cupboard was found a very handsome black and white heart- shaped jar (pl. 13, d), accompanied by a lid of micaceous schist and two small manos with flattened ends; also a fine buckhorn chisel, a large piece of buckhorn, a good bone awl, and several black and white sherds. A sketch of the cupboard is given in figure 5. Standing erect 14 SMITHSONIAN MISCELLANEOUS COLLECTIONS voL. 81 in about the center of the room were the remains of a post approxi- mately ten inches in diameter which was probably intended to support the roof. KITCHEN Another room in the eastern part of the group was probably used as a kitchen or cooking room. There were several poorly defined fireplaces in the floor, and a shallow trench ran along the entire south- Toor 18 well plastered Z! ooy. Fic. 6.—a, firepit containing ashes; b, pot rest with flat stone in bottom; c, metate; d, ash pile; e, f, storage boxes, Io inches deep; g, shallow trench well plastered. ern wall. This room was in such bad condition that it was almost impossible to get a definite idea of its original equipment of pots, etc. The few indications were enough, however, to establish its character (fig. 6). KIVA One of the most remarkable features of the ruin is the location of the kiva. Instead of being in an open plaza, detached from the main ruin, it is located almost in the center of a compact mass of rooms and is surrounded on all sides, thus completely cutting it off from the outside. The northeastern corner is enclosed by a wall which was in such bad condition that it was impossible to establish whether there had been rooms there or only an enclosing wall. The mound at this point is badly washed out, and although a thorough exploration was made, nothing definite could be established. When the plaza, in which the kiva is located, was first cleaned off, it appeared that the area was used for dance purposes only. There NO. 12 ARCHEOLOGY OF TAOS VALLEY—JEANCON I5 was a good hard stamped floor, with only a large river boulder in about the center to break it. After the floor was cleared, the writer decided to raise the boulder to see if there might be something below it, as it seemed out of place in the plaza. When the stone was raised a section of curved wall was disclosed, which led to the excavation of the kiva. The one prominent problem of the kiva is the double wall on the west side (pl. 4). It was impossible from the excavation to determine why the two walls were used. At times it seemed to suggest two occupa- tions ; again this impression was destroyed by other indications which seemed to point to a single occupation. Both walls are alike in ma- terial and construction. The outside wall runs around about one half of the inner one. Starting at a point almost due north it runs around the western side and terminates almost due south. There is no outer wall on the eastern side. The ventilators are on the inner walls only, one east and one west (pl. 5, 4). Both of these are barrel-shaped. The fill between the walls was composed of soft dirt, débris of roof material, river boulders, bits of pottery, and a few artifacts in bad condition, although a very handsome pipe was found in the trash. A curious feature occurring only in the kiva is the hori- zontal lines running around the inside of the walls. At first it would appear that these walls were laid up in regular courses, but an examination of shattered fragments shows that the wall was built in the same manner as all the others in the group, and that the horizontal lines do not indicate the use of moulds as would be suggested by their appearance. The walls still rise seven feet in height from the floor. They are about nine inches thick, and are very well made and hard. The floor was formed of packed adobe, probably mixed with blood and ash as was the custom formerly. Three feet above the floor, in the débris which filled the chamber, was a deposit of two and one- half feet of drift sand, and between the sand and the floor were the remains of the roof beams, but these were in such bad condition that nothing could be learned of the roof construction. There were two erect cedar posts at points shown on the map of the kiva, and in related positions were two holes in the floor, where additional supports had stood. These four uprights served to support the roof and show an interesting method of roof construction, one which was continued in later structures in the region. 2 10 SMITHSONIAN MISCELLANEOUS COLLECTIONS voL. 81 The Castafieda Report refers to the supporting pillars as follows: The young men live in the estufas, which are in the yards of the village. They are underground, square or round, with pine pillars. Some were seen with twelve pillars and with four in the center as large as two men could stretch around. They usually had three or four pillars.’ And again: At this village they saw the largest and finest hot rooms or estufas that there were in the entire country, for they had a dozen pillars, each one of which was twice as large around as one could reach and twice as tall as a man.? The feature has been found in other prehistoric ruins, however, for to quote from Judd: fragments of curved adobe walls remained on the eastern side and these, if continued, would have circled a central fireplace about which four large pillars [posts?] formerly stood..... kivas with roofs supported by uprights were noted, also, during preceding expeditions.® The fallen roof masses on the floor were not in condition to give further information concerning the construction. Scattered all through the debris were fragments of human and animal bones, as well as a few pottery sherds. There were no indications of burials in the kiva. The fireplace (pl. 5, B), in about the center of the floor, was un- usually fine. When first found it was full to the top with wood ashes. Its inside diameter was two feet three inches. To the east of the fireplace stood a flat river boulder about one foot wide and one and one-half feet high; next to this (east) was a small pit, oval in form and measuring one foot nine inches by one foot six inches, with a depth of nine inches. The use of this pit is not known. Four feet east of this, in the wall, was a doorway measuring two feet in height and one foot in width from which a passage, roofed over with poles four inches in diameter, led through the wall for a distance of four feet. Here it ended abruptly against another wall. The east ven- tilator was directly above this, but was not connected with the passage. The only entrance to it appeared to be from the kiva. No ceremonial objects of any kind were found in the kiva proper. * Winship, Translation of the Castafieda Report. 14th Ann. Rep. Bur. Amer. Ethnol., page 518. *Idem, page SII. * Judd, Neil M., Archeological Investigations at Paragonah, Utah. Smith- sonian Misc. Coll., Vol. 70, No. 3, p. 15. } NO. 12 ARCHEOLOGY OF TAOS VALLEY—JEANCON 7, Minor ANTIQUITIES STONE ARTIFACTS Most of the stone artifacts were of the usual character. Manos and metates were of several forms, the unusual depth of the latter indicating a long period of use (pl. 6, b). Maul heads of the usual type presented no new features (pl. 7, d@). No axe heads were found. Several small cylindrical stones of unknown use were found. While they were not of any definite character, they showed plainly the marks of having been used for some purpose, probably polishing (pl. 7, b). Pot lids and other articles of micaceous schist were rather plentiful (pl. 6). The writer has never before heard of the use of this material for stone artifacts. Several larger slabs of it had evidently been used as baking stones, as one surface was smoked and the other covered with a heavy deposit of grease. Eccentric forms.—This group includes a troughlike stone which was possibly used for smoothing arrow shafts (pl. 7, @). It measured two inches long by one inch wide, the trough being 5/16 inch deep and 11/16 inch wide at the top. It showed marks of rubbing in the trough and on the top, bottom, and sides. Another was a triangular shaped stone, with indentation, which also was probably an arrow or javelin shaft smoother (pl. 7, c). It was found with a fine red pipe, a banded stone, and a fossil (Turwillana). Chipped implements—Rather ordinary forms and workmanship were presented in the chipped implements (pls. 8 and 9), the only exception being the object shown in the upper left corner of plate 8, an unusual knife blade of especially fine chalcedony. The cutting edge is well chipped, and the top forms an excellent handle. The material used for chipped and crude cutting edges, arrow heads, javelin heads, drill points, etc., included chalcedony, agatized wood, slate, moss agate, obsidian, and a sort of hard shale. One especially fine javelin head measured 2} inches across the broadest part by 33 inches in length (pl. 8, lower left-hand figure). BONE ARTIFACTS Among the bone artifacts, also, only a few examples were out of the ordinary. Elk or deer horn was represented by flakers and chisels (pl. 10, b). Three pieces of rib bones (pl. 10, ¢) were inter- esting; the longest of these is 11 inches, the shortest 2} inches in length, although the latter was probably longer originally, as one end of it is broken. All three of these specimens are notched, two with 18 SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL. 81 teeth or notches on both sides and one with the teeth only on one side. Their use is unknown, although they suggest the notched stick which is used by many of the southwestern Indians to accentuate the rhythm of a song. The notched stick is placed with one end resting on a drum head or the bottom of an inverted basket, and another stick is rubbed across the notches ; the noise thus produced falling upon the regular beat of the measure serves to accentuate the rhythm. Only a few of the best bone articles are shown in the illustrations. These artifacts were in great abundance. PIPES Although only one whole pipe was found, fragments from several others are sufficient to indicate that the ordinary form was that of the tubular pipe or cloudblower. All were made of pottery clay. The best specimen (pl. 11, a), found between the double walls of the kiva and about two feet from the floor, is 25 inches long, 3 inch in diameter at the large end and 2 inch at the mouth piece; it is red in color and was decorated by a series of striations beginning 4 inch from the mouth and running to the edge of the bowl. The clay of which it was made is so hard that it was thought at first to be made of stone. The fragment shown on plate 11, b, is from a cloudblower made of the black cooking-pot paste. It is 2 inches long, $ inch wide at the top, and 58; inch at the mouth, and is decorated with an incised snake design. The indented fragment (pl. 11, c) has a grayish black soft paste, and measures 2;°; inches long, ? inch wide at the top of the fragment, and 2 inch at the mouth. Plate 11, d, shows a fragment from a pipe made of soft black paste, with line and dot incised decoration. It is of the cloudblower shape, and is 14 inches long and 3 inch in greatest diameter. The last fragment shown (pl. 11, ¢) has a gray paste and is dec- orated with 7 incised horizontal lines which do not show in the illus- tration. It measured } inch in greatest diameter and 1} inches in length. FETISH OBJECTS The prehistoric Indians in the Southwest had a tendency to collect curious fossils, concretions, brightly colored stones, and other natural objects, and it seems quite probable that they were used as fetishes or medicine stones. The people at Llano were no exceptions to the rule, and a number of fossils and odd-shaped stones suggesting such usage were found. Among the fossils are two which are interesting because NO, 12 ARCHEOLOGY OF TAOS VALLEY—JEANCON IQ they bear the same name, in the Taos dialect, as a clan which formerly existed at Taos but is now extinct, the Turwillana clan.’ The fossils are of the cylindrical variety marked with rings. It is possible that their presence in the ruin may indicate that such a clan had lived there. Another of these objects was a handsomely polished banded piece of ribbon agate (pl. 11, 2) probably used as a medicine stone, found in room 8. The colors are white, cream, gray, light brown, and black in irregular bands. It measures 24 inches in length and 4 inch in greatest diameter. INDICATIONS OF BASKETRY While no baskets or fragments of basketry were found, many potsherds with basket impressions on them (pl. 11, 7) were picked up during the summer, and these may be considered as a good indication that the people did have baskets. Judging from the impressions on the pottery fragments, the sticks of all the baskets used in the village were about the same size. In one case only does it appear that a larger bundle or rod had been used. On one fragment of pottery, there are what appear to be textile impressions or smears; these can be seen only under a magnifying glass; how they occurred it is at present impossible to tell. The presence of basket impressions on pottery, which are more common in this region than at any other place where the writer has worked, has been explained by Dr. A. V. Kidder in a discussion of similar markings found on vessels from the Jemez Plateau. Dr. Kidder writes: An extraordinarily high percentage of basket marked sherds is found at the small house ruins. Such sherds occur, it is true, in most other Black-and-White groups, but they are of greatest rarity. Here, however, they can be picked up at almost any site. The impressions show that bowls and lower parts of ollas were often formed in baskets. In these cases the clay was apparently coated on the inside of the basket and pressed down hard enough to render the marks of the weave sharp and clear. The upper parts of the ollas were probably constructed by the regular coiling method. Some bowls, however, seem to have been molded or cast entire in basket forms, as the impression of the weave runs to the rim. The baskets themselves were of the coiled variety, tray or bowl shaped; the coils measure 4 to 5 mm. in breadth and there are about six sticks to the centimeter.’ *Handbook of the American Indians, Bull. 30, Bur. Amer. Ethnol., Pt. 2, p. 690. * Kidder, A. V., Pottery of the Pajarito Plateau and some of the Adjacent Regions of New Mexico. Mem. Amer. Anthrop. Assoc., Vol. II, pt. 6, p. 414. 20 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 81 POTTERY The pottery from Llano may be classed in three groups, the black cooking vessels; the black on white bowls, ollas, and jars; and the black on red pottery. There were no examples of the indented or corrugated pottery in the ruin at Llano, but on the surface there was an abundance of such sherds. The large jars from the ruin appear to have been made in the usual coil technique but the coils were obliterated, by wiping the surface with a corncob or some other object with a rough surface, as work on the vessel progressed. After the completion of the pot, decorations were incised around its upper part. On many the lower portions show traces of the coils which were not completely obliterated by the rubbing process. The following quotation from Kidder gives a good impression of the general appearance of the ware: The latest black ware so far identified is the striated; there is, of course, no sign of the coil, but the surface is scored with a series of fine and more or less parallel scratchings, criss-crossing over each other in all directions. These were produced evidently, by some finishing tool used while the clay was moist. Experiment shows that a corncob with the kernels removed most nearly duplicates these marks.” Although the coiling is still slightly apparent in places on the Llano vessels, the striations described by Kidder are also in evidence. As a matter of fact, the Llano forms seem to represent a combination of Kidder’s black corrugated and striated forms. The incised designs are of the simplest character consisting usually of horizontal lines, although on one pot (pl. 12, @) there has been an attempt to establish three zones of decoration, the middle zone dif- ferent from the upper and lower ones. The short dashes of the mid- dle zone appear to have been made with a long thumb nail. The writer encountered a somewhat similar ware in an investigation of La Jara Cafion, New Mexico, situated at the southern end of the Jicarilla Apache reservation. However, there was considerable dif- ference in the pastes of the two areas, that of Llano being much softer and without a great amount of temper, while that of La Jara Cafion was composed of materials that burned almost to a vitrified brick. The vessel marked C on plate 12 shows unobliterated coils on the neck beginning just above the shoulder. Below the coils are incised lines. The bottom has the usual obscured coils observed on practically all of the pots. 1 Kidder, A. V., Notes on the Pottery of Pecos. Amer. Anthrop. n. s., Vol. 19, p. 339, Lancaster, 1917. NO. 12 ARCHEOLOGY OF TAOS VALLEY—JEANCON 21 As will be seen from the illustrations the incisions on the various vessels were not all made with the same sort of an implement. Most of them are sharply cut, but in those shown on plate 12, d, e, f, a blunt broad implement was used. All of the pots are more or less asymmetric, but so slight is the ir- regularity that it is rather pleasing than otherwise. The balance in form is also uneven; taking the vessel as a whole and noting the line of greatest diameter, we find that there is no set rule as to where it occurred in the vessel. Sometimes the greater part of the jar was above this line, and sometimes below it. Plate 13, 0, illustrates a water jar of beautiful form, grayish in color and showing only partially obscured coils in places. The paste of this vessel is much finer than that of the ordinary cooking or storage pot; it seems to fall between the black ware and the black on white decorated ware. All of the forms are strikingly reminiscent of the earlier Apache forms, especially of the Apache water bottle. Basket markings on the bottom of pottery vessels are so common as to give the impression that a large majority of the pots were started in the manner described by Dr. Kidder on page 19. Some of the crude black pots have excellent prints all over the bottom and often rising as much as two inches up the sides. In plate 13, c, a black on white piece, there may still be seen basket imprints just below the shoulder where the white slip has more or less disappeared. The prints on the bottom of this vessel are very plain. A number of interesting sherds are shown in plate 11, 7. Basket impressions have been recorded from many places. Dr Kidder’ speaks of them in the San Juan drainage as follows: “Pottery vessels, on the other hand, were scarce and crude, and usually bore on their bottoms the imprint of the baskets in which they had been formed.” Black on white ware——Black on white ware was well represented at Llano and appears to be related to the Rio Grande black on white ware as well as to the southwestern Colorado wares. In the Taos region fully one-third of the sherds are very good white with excellent jet black decorations. One feature is very noticeable, namely, that the crackling or crazing of the white slip appears to have been done more by exposure to the elements than in the original baking of the ware. 2 Prehistoric cultures of the San Juan drainage. Proc. roth Internat. Cong. Americanists, p. 108. 1917. SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 81 to to On the sherds that are buried the white slip is not crazed, but where the sherds have been lying exposed on the ground, there appears to have been a deterioration and crazing has taken place in varying degree, depending probably on the length of time they have been exposed and on weather conditions. No hard and fast rule seems to have been followed as to the zones of decoration. In the bowls the percentage of decoration on the ex- terior is practically the same as on the interior. This of course applies only to bowls; ollas are decorated only on the outside. As far as could be determined from the rim fragments found, the rims were so thin that they could not be decorated. No dots, ticks, or zigzag lines were found on the rims; in many cases the exterior as well as the interior decorations extended to the rim. No out-turned rims were found. Decorations of banded lines occur, but are not common. The usual arrangement of such designs is a broad band at the top and bottom, Fic. 7—Animal figure on potiery. with three or four thinner lines between these. The paste of the black on white ware is usually hard and homogeneous. Little variation in color can be noted in the cross section of a sherd. The following quotation from Dr. Kidder’s notes on the black on white ware of Pecos is interesting for comparison: Slip-color ranges from light to dark gray, very rarely purely white [good white about 333 per cent at Taos]; it is applied to the interior oi the bowls in a relatively heavy coat; to the exterior usually as a thin wash, occasionally as a heavy coat, rarely omitted altogether. Slip often cracked, particularly when it is applied heavily on the exterior. Finish of the interior even but never glossy; of exterior much rougher. No specimen with corrugated or basket marked exterior observed. [Note difference in Taos basket marks.] Ornamentation-pigment dull black, usually of slaty cast. Zones, interior of bowls exceedingly rare. [Note difference from Taos pottery.] Straight bowl rims usually plain, occasionally dotted; out-curved rims bear ticks or more commonly zigzag lines. The main ornament appears to have been in the form oi a broad band encircling the interior oi the bowl and leaving small blank spaces in the bottom. Bands framed above and below by single heavy lines (“N™ Plate VII, Figure 7); less commonly by one heavy line with a series i NO. I2 ARCHEOJ.OGY OF TAOS VALLEY—JEANCON 23 of lighter ones between it and the band (‘““N” Plate VII, Figure 8). Line breakers in framers rare. “All over” decorations apparently fairly common, particularly on small bowls. Design preponderatingly geometric and rectilinear, life and curvilinear forms practically absent. [At Taos, a few sherds with curves, one life figure.] Elements of design most commonly observed; coarse hatching and cross-hatching; plain and dotted checker-board series of plain triangular figures ; dotted lines and edges; large stepped figures in opposed pairs. Brushwork normally crude and uncertain, lines coarse. Paste composition—Paste, fine, very hard and homogeneous; color ranges from pure white (rare) through light gray to dark gray (rare). Many sherds of even color from surface to surface, but in the majority the center is darker than the edges. Gross tempering (sand or pounded rock) apparently absent.’ No true biscuit ware was found, unless some of the vessels might be considered to come under the classification given by Nelson in the following quotation: There seem to be two kinds of biscuit ware, the most common being of a dull white or light gray color, the other of a yellowish tone. This latter has its probable forerunner in a more or less distinguishable variety of black on white ware, but the prototype of the former has not been found so far.” Although biscuit ware is found in the region, the writer is inclined to believe that the Llano black on white ware does not represent a form of biscuit ware, but a true black on white. To quote again from Nelson: The pottery actually figuring in the table is a local variety of the black on white ceramic identified with the general sub-stratum of Southwestern Pueblo culture. Bandelier generally associated the ware with “small houses,” 7. e., with what might be called a pre-pueblo stage of sedentary life; but the data now at hand enable us to state that the large quadrangular form of village typical of the Rio Grande valley in later times was fully developed before the black on white pottery went out of style. The ware as a whole is perhaps not quite so fine [not the case at Taos] as that of Mesa Verde and Chaco regions on the one hand or of the Upper Gila and Mimbres regions on the other. It is particularly lacking in variety of form. In decorative symbolism it approaches the abandoned northwestern Pueblo area rather more than the southwestern, and is little, if at all inferior to it” As far as could be determined there were no pre-pueblo houses in the Taos region. The small ruins were apparently of the same period as the larger ones. No differences were noted between the pottery and other artifacts from the large and small sites. * Kidder, A. V., loc. cit., pp. 327-328. ? Nelson, N. C., Chronology of the Taos Ruins, New Mexico. Amer. Anthrop. n. s., Vol. 18, p. 169, Lancaster, 1916. * Idem, page 171. SMITHSONIAN MISCELLANEOUS COLLECTIONS fi) ‘QU QTL 3 LLL aN Fic. 10.—Design on black on white ware. VOL. 81 a RL rr a Pate to ut NO. I2 ARCHEOLOGY OF TAOS VALLEY—JEANCON VEX. 7® NSS h Fic. 11.—a, b, c, d, interlocking fret, Taos. e, f, g, h, interlocking fret, Aztec. 26 SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL. 81 As may be seen from the figures drawn from sherds, no curvilinear designs at all were found, all being strictly rectilinear ; only one life form was noted (fig. 7). The black on white ware was mostly in the form of ollas and bowls. The small heart-shaped jars are particularly handsome (pl. 13, c, d). Two of these were found, one whole and the other in such condi- tion that it could be restored. The designs are shown in plate 13, c, d, and figures 8, 9. Figure 10 is a reconstructed design from the top of a water jar. Enough fragments of this were found to recon- struct about five-eighths of it. The interlocking fret shown in a number of cases on the Taos black on white ware is almost identical with that found in the Aztec ruin Fic. 12.—Interlocking fret design. From Oak Tree House, Mesa Verde National Park, Southwestern Colorado. (fig. 11), and figured by Earl Morris in the American Anthropologist, Vol. 17, 1915, p. 676, and also on sherds found by the writer when acting as assistant to Dr. J. W. Fewkes, at Oak Tree House, on the Mesa Verde, in 1921 (fig. 12). The dividing of the top of an olla as shown in figures 8 and 9g is common to most parts of the southwestern prehistoric Pueblo culture area. Figure 13 was taken from the paper cited above on the Aztec Ruin by Earl Morris. The varied designs of the Taos region include zigzags, triangles, checkerboard and many other forms (fig. 14, and pl. 14). Black on red ware—Only a few sherds from vessels belonging to this group were found and these are not enough to give any definite opinion as to the general characteristics of the type in the Taos region. a6 WO; 12 ARCHEOLOGY OF TAOS VALLEY—JEANCON , water jan; Gyros black VOL. 81, NO. 12, PL. 13 on white ware. SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 81, NO. 12, PL. 14: Types of pottery designs. a a A tt al nlp ip lig ay - 1 a ee Mtn ee 15 2s. NO. VOL. 81, ottery handles. pes of p Ty SMITHSONIAN MISCELLANEOUS COLLECTIONS Pe SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLUME 81, NUMBER 13 ‘| DESCRIPTIONS OF FOUR NEW FORMS OF BIRDS FROM HISPANIOLA AS IBY ALEXANDER WETMORE A +h Ba A &NX J, ( List om i \ joss ’ “ Se DEF), Fh PE CITY OF WASHINGTON PUBLISHED BY THE SMITHSONIAN INSTITUTION MAY 15, 1929 v - 4 Sa . . ie < ‘s ef ~s s . : ‘ ‘4 5 suse a i , . : $ ‘ ‘ ite | = y ‘ 5 i pas S I ; : = = ~ ¢ u 5 ‘Ss o ‘i x } 4 = x is ¥ . 7 ¥ ¢ - ; Aa 1 bh 2 \ a 7 2 A 7 ~ “ nt \ 7 : , z ' } s } ; : e s ‘ 2 i L eee < > j , ios - / s ee ~ ; ‘ - ¥ 2 ; i ‘ - i a S ; . * rst Chet ete eS x 5 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLUME 81, NUMBER 13 DESCRIPTIONS OF FOUR NEW FORMS OF BIRDS FROM HISPANIOLA BY ALEXANDER WETMORE (PUBLICATION 3021) CITY OF WASHINGTON PUBLISHED BY THE SMITHSONIAN INSTITUTION MAY 15, 1929 The Lord Battimore Press BALTIMORE, MD., U. 8 A. DESCRIPTIONS OF FOUR NEW FORMS OF BIRDS FROM HISPANIOLA By ALEXANDER WETMORE Continued studies of birds from Haiti and the Dominican Republic in the National Museum have brought to attention three geographic races found on small islands off the coast that differ sufficiently from the groups inhabiting the main island to merit subspecific distinction. In addition there has been found in the American Museum of Natural History a very distinct species of the peculiar genus Calyptophilus which is described here through the courtesy of Dr. Frank M. Chapman. Family MNIOTILTIDAE DENDROICA PETECHIA SOLARIS, subsp. nov. Characters —Similar to Dendroica petechia albicollis (Gmelin)’ but lighter in color ; averaging slightly larger. Description—Type, U. S. Nat. Mus. No. 278,738, male, collected at Z’Etroits, Gonave Island, Haiti, March 18, 1920, by Dr. W. L. Abbott. Upper surface slightly brighter than pyrite yellow, rump brighter yellow; anterior portion of crown somewhat brighter than sulphine yellow, with concealed portions of feathers sudan brown; wings dusky, the feathers margined externally with sulphine yellow ; wing-coverts pyrite yellow, edged with lemon chrome; rectrices dusky, lightly edged externally with sulphine yellow, with inner webs exten- sively lemon yellow ; under surface between light cadmium and lemon chrome, with breast and sides streaked with sudan brown. Mazxilla dusky neutral gray, mandible deep neutral gray, tarsus and toes dull brown (from dried skin). Measurements (in millimeters) ——Ten males, wing 64.2-68.0 (65.9), tail 50.0-57.0 (52.1), culmen from base 12.3-13.1 (12.8), tarsus 20.5-21.5 (21.3). Seven females, wing 61.3-63.2 (62.2), tail 48.1-51.0 (48.9), culmen from base I1.5-12.9 (12.2), tarsus 21.2-22.0 (21.4). Type, male, wing 67.0, tail 57.0, culmen from base 12.5, tarsus 25. 2 Motacilla albicollis Gmelin, Syst. Nat., vol. 1, pt. 2, 1780, p. 983. (“S. Dominici” = Hispaniola. ) * Average of nine. SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 81, No. 13 SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL. 81 bo Range.—Confined to Gonave Island, Haiti. Remarks.—tThe present form shows approach to Dendroica petechia eoa of Jamaica in size and in brighter coloration, but is less golden yellow, with the dark coloration of the anterior part of the crown rufescent rather than orange brown. Its differences from albicollis, easily apparent in series, are more plainly evident in females than in males. Following are measurements of Dendroica p. albicollis for com- parison : Eighteen males, wing 59.3-64.5 (62.5), tail 47.3-53.0 (50.5), culmen from base 11.0-13.3 (12.6), tarsus 19.9-22.0 (21.0) mm. Five females, wing 57.8-63.0 (60.3), tail 47.8-50.2 (49.2), culmen from base I1.2-13.2 (12.4), tarsus 19.8-21.0 (20.6) mm. Family THRAUPIDAE CALYPTOPHILUS TERTIUS, subsp. nov. Characters.—Similar to Calyptophilus frugivorus frugivorus Cory * but decidedly larger, with heavier bill; much darker in color ; feathers encircling eyelids dark, instead of yellow; wings and tail rufescent. Description—American Museum of Natural History No. 166,421, male, from higher slopes of MorneLa Hotte, Haiti, taken June 22, 1917, by R. H. Beck. Crown feathers centrally chaetura black, margined with grayish olive; sides of head chaetura drab; a spot of primuline yellow on lores, separated from eye by an area of dull black; hindneck, back and scapulars between deep and dark olive, shading back across rump to the bister of the upper tail-coverts ; rectrices somewhat darker and more rufescent than warm sepia; remiges dusky brown, sec- ondaries and inner primaries with external webs bister, outer pri- maries with external webs olive brown, and wing-coverts clove brown ; edge of wing, with a wash of reed yellow; under wing-coverts mixed whitish and deep mouse gray ; throat and center of breast white (more or less soiled) ; abdomen smoke gray; sides washed with deep mouse gray; flanks brownish olive; under tail-coverts bister. “ Iris brown, bill black, horn below ” (from collector’s label) ; tarsi and feet bister (from dried skin). Measurements (in millimeters).—Five males, wing 92.5-104.0 (98.9), tail 96.5-108.0 (101.2), culmen from base 23.7-27.4 (24.9), tarsus 32.5-35.0 (33.4). *Phoenicophilus frugivorus Cory, Quart. Journ. Boston Zodl. Soc., October, 1883, p. 45. (‘‘ Santo Domingo” = Rivas, D. R.) NO. 13 NEW BIRDS FROM HISPANIOLA—-WET MORE 3 Two females, wing 84.4-91.5 (88.0), tail 83.9-89.0 (86.5), culmen from base 22.8-23.6 (23.2), tarsus 30.8-31.1 (31.0). Type, male, wing 100.0, tail 100.2, culmen from base 24.5, tarsus | 32.8. Range-——Known only from the higher slopes of Morne La Hotte, Haiti. Remarks—The series of seven Calyptophilus in the American j Museum of Natural History, collected by R. H. Beck from June 20 to | july 4, 1917, back of Les Anglais on the higher ridges that lead up to | the peak of Morne La Hotte, are so distinct in larger size and darker | coloration from Calyptophilus frugivorus of the rest of the island that | there is no question in assigning them specific rank. The new form | seemingly is confined to the higher slopes of the mountain range of ’ La Hotte, since during extensive collecting elsewhere it has not been found. It is another of the peculiar mountain forms of Hispaniola whose presence has been wholly unexpected. This new species is described here through the kind permission of i Dr. Frank M. Chapman of the American Museum of Natural History. | PHAENICOPHILUS PALMARUM EUROUS, subsp. nov. Characters—Similar to Phaenicophilus palmarum palmarum (Lin- i naeus)* but lighter in color; above brighter green, with gray of hind- i neck lighter, becoming nearly white on sides of neck anteriorly ; below with white more extended. | Description—Type, U. S. Nat. Mus. No. 252,843, male, taken on Saona Island, Dominican Republic, September 13, 1g19, by Dr. W. L. Abbott (in somewhat worn plumage). Crown and entire sides of head jet black, except for a spot on either side of the forehead, one above i either eye, and the lower eyelid, which are white ; hindneck, and sides , a little paler than gray no. 7, the former becoming white at posterior i border of black covering the ear-coverts; upper surface, including exposed portions of wing and tail feathers, between pyrite yellow and warbler green; concealed portions of remiges dusky ; median under- | parts including the under tail-coverts extensively white ; under wing- coverts smoke gray edged with white. Bill black, base of lower mandible dark dull grayish; tarsus dusky neutral gray (from dried skin). Measurements (in millimeters ).—Type specimen, male, wing, 90.0, tail, 67.5, culmen from base, 20.7, tarsus, 22.5. * Turdus palmarum Linnaeus, Syst. Nat., ed. 12, vol. 1, 1766, p. 295. (‘‘ Habitat in Cayennae Palmis ’’ = Hispaniola.) 4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL. 81 Range.—Restricted to Saona Island, Dominican Republic. Remarks.—Though occasional birds in a considerable series of the palm tanager from the main island of Hispaniola approach in colora- tion the single skin at hand from Saona, none is quite so extensively white below, or so bright and light a green above, so that it seems permissible to name a Saona form on the basis of this scanty material. Family FRINGILLIDAE LOXIGILLA VIOLACEA MAURELLA, subsp. nov. Characters —Generally similar to Lowvigilla violacea affinis (Ridg- way )* but larger, with heavier bill. Description—Type, U. S. Nat. Mus. No. 250,456, Tortue Island, Haiti, collected February 2, 1917, by Dr. W. L. Abbott.” Throat, short line above eye, and under tail-coverts somewhat darker than burnt sienna ; axillars and under wing-coverts partly burnt sienna and white mixed ; plumage otherwise deep black. Bill, feet, and tarsi black (from dried skin). Measurements (in millimeters) —Three males, wing 82.4-84.3 (83.4), tail 70:2-71.9 (71.2),,culmen from: base/16.2-16:8 (1614), depth of bill at base 12.9-13.8 (13.3), tarsus 22.1-22.8 (22.4). One female, wing 77.7, tail 65.8, culmen from base 15.5, depth of bill at base 11.9, tarsus 22.9. Type, male, wing 82.4, tail 71.9, culmen from base 16.8, depth of bill at base 13.8, tarsus 22.8. Range.—Known only from Tortue Island, Haiti. Remarks.—tThe greater size of the present form is more plainly evi- dent on direct comparison of skins from Tortue with those from Hispaniola proper than from examination of the measurement tables, maurella being very appreciably larger and more robust. The only female seen is blacker than the majority of affinis. The.new form is represented by three males and one female, all collected by Dr. W. L. Abbott, February 1 and 2, 1917. Following are measurements of L. v. affinis for comparison : Sixteen males, wing 71.9-79.2 (76.7), tail 61.7-69.3 (65.3), culmen from base 14.2-16.5 (15.2), depth of bill at base 11.0-12.9 (12.3), tarsus 19.2-23.4 (21.1) mm. Nine females, wing 67.2-75.8 (71.2), tail 59.8-67.0 (63.5), culmen from base 12.6-14.3 (13.6)* depth of bill at base 10.4-11.8 (10.9), tarsus 19.7-22.3 (21.0) mm. * Pyhrrulagra affinis “(Baird)” Ridgway, Auk, 1808, p. 322. (Port-au-Prince, Haiti.) * Average of eight. | MISCELLANEOUS COLLECTIONS ‘ ae _ VOLUME 81, NUMBER 14 : PREHISTORIC ART OF THE ALASKAN con ESKIMO” RIAN ASTI | 2 , Vay = : (WitH 24 PLATES ie NOV 15 1929 3 | ! OFFice LBRANY 4 BY. HENRY B. GOLLINS, JR. a Assistant Curator, Division of Ethnology, U. S. National Museum | ERMEINCRE Le a ole CASO SGE'A e: : ss Sah ee Pe Oy ate ee ae Nba Tusoxt | ms Se eee qrry OF WASHINGTON. ap ae | PUBLISHED BY THE SMITHSONIAN INSTITUTION Reig eae® ~ NOVEMBER 14, 1929 Wakeen A SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLUME 81, NUMBER 14 PREHISTORIC ART OF THE ALASKAN ESKIMO (WITH 24 PLATEs) BY HENRY B. GOLLINS, JR: Assistant Curator, Division of Ethnology, U. S. National Museum (PUBLICATION 3023) CITY OF WASHINGTON PUBLISHED BY THE SMITHSONIAN INSTITUTION NOVEMBER 14, 1929 © 2 vs bad = 2 o £ & 3 @ 2 oa 6 ay 2 a » BALTIMORE, MD., U. S. A. PREHISTORIC ART OF THE ALASKAN ESKIMO By HENRY B. COLEINS; JR, ASSISTANT CURATOR, DIVISION OF ETHNOLOGY, U. S. NATIONAL MUSEUM (With 24 PLATES) INTRODUCTION Until very recently information on the archeology of the American Arctic was limited practically to the descriptions by Wissler* and Boas* of a relatively small number of specimens collected by Stefansson, Comer, and others from northern Alaska, the Hudson Bay region, and northwest Greenland. The first systematic excava- tions in the eastern regions were those made in Baffin Land, Melville Peninsula, and northwest Greenland by Therkel Mathiassen for the Fifth Thule Expedition from 1922 to 1924. The publication in 1927 of the results of these important investigations afforded for the first time an adequate view of the archeology of a large section of Arctic America.” This work verified the conclusions of Boas and Wissler that in earlier times there had been a closer similarity between the Eskimo cultures of Alaska and the eastern regions than exists at present. Mathiassen, however, with a much larger mass of material system- atically excavated at a number of widely scattered sites, was able to go further and show that the similarities were so numerous and strik- ing that the Thule culture, the name he gave to the ancient eastern culture, must have had its origin in Alaska. In 1926 Dr. Ales Hrdli¢ka made an anthropological survey of the Alaskan Coast from Norton Sound to Point Barrow * and in the same year Mr. Diamond Jenness inaugurated archeological work in the Bering Sea region by excavating at Cape Prince of Wales and on the *Wissler, Clark, Harpoons and Darts in the Stefansson Collection. Anthrop. Papers Amer. Mus. Nat. Hist., Vol. XIV, Part II, 1916. Wissler, Clark, Archeology of the Polar Eskimo. Anthrop. Papers Amer. Mus. Nat. Hist., Vol. XXII, Part III, 1018. ? Boas, Franz, The Eskimo of Baffin Land and Hudson Bay. Bull. Amer. Mus. Nat Elist Viol, ove, Part I; toor; Part i, 1907: ® Archeology of the Central Eskimos. Report of the Fifth Thule Expedition, 1921-24. The Danish Expedition to Arctic North America in Charge of Knud Rasmussen, Ph. D., Vol. TV, Parts 1 and 2, Copenhagen, 1927. * Anthropological Work in Alaska. Smithsonian Misc. Coll., Vol. 78, No. 7, PP. 137-158, 1927. SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 81, No. 14 ee Se EO FT St OT ETE - es <= Or Sa <2 A pa RI a as 2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 8&1 Little Diomede Island.* As a result of these investigations it was shown that underlying the existing Eskimo culture of northern and northwestern Alaska there had been an earlier and in general more advanced culture which was marked especially by elaborately carved and ornamented objects of old ivory. The ancient style of art re- vealed by Hrdli¢ka and Jenness was distinct and new, although some of the harpoon heads and foreshafts corresponded with the Thule types of the east. In 1927 I conducted anthropological work in southwest and west Alaska with Mr. T. Dale Stewart of the United States National Museum, examining sections of the coast and islands from the Alaska Peninsula northward to the mouth of the Yukon.’ This work con- sisted mainly of measuring the Eskimos and making collections of skeletal and cultural material. Although many old graves, village sites, and some few refuse piles were examined, no trace was found of the newly discovered ancient culture above referred to, which seems ac- cording to the present evidence not to have extended as a type south of St. Lawrence Island. In 1928 I returned for a second season’s work, and excavated on St. Lawrence and the small nearby Punuk Island, and later at Metlatavik on the Arctic coast just above Bering Strait.* This work resulted in the collection of a large number of specimens that appear to be of particular interest as showing successive stages of art de- velopment in the newly revealed ancient Bering Sea culture. The detailed description of all this material will necessarily be somewhat delayed, but in order that the more important results may be made available as soon as possible it seems desirable to present in advance a brief description dealing with the art of St. Lawrence and Punuk Islands, together with a description of such additional examples of the old art from other Bering Sea and northern Alaskan sites as I have been able to obtain or have photographed. * Archeological Investigations in Bering Strait. Bull. 50, Ann. Rep. for 1926, Nat. Mus. Canada, pp. 71-81, Ottawa, 1928. Ethnological Problems of Arctic America. Amer. Geogr. Soc., Special Publ. No. 7, Problems of Polar Research, pp. 167-175, New York, 10928. * The Eskimo of Western Alaska, Explorations and Field-Work of the Smith- sonian Institution in 1927, pp. 149-156, 1928. *The Ancient Eskimo Culture of Northwestern Alaska. Explorations and Field-Work of the Smithsonian Institution in 1928, pp. 141-150, 1920. The expense of the expedition was borne by Mrs. Mary Vaux Walcott, the Bureau of American Ethnology, and the American Association for the Advance- ment of Science. NO. I4 PREHISTORIC ART OF ALASKAN ESKIMO—COLLINS 3 Jenness has described some of the archeological material which he collected at Cape Prince of Wales and on the Little Diomede as rep- resentative of what he has called the Bering Sea culture, and this designation will be followed in the present paper. Hrdlitka, in a publication now in press, has also described the specimens he obtained in 1926. In addition, Mathiassen has recently published a description of several examples of the Bering Sea art which were in the collections of the Museum of the American Indian and in the National Museum at Copenhagen.” To Dr. George G. MacCurdy, however, belongs the credit for first calling attention to this unique style of Eskimo art some years before any archeological work had been done around Bering Strait and when there was no material with which to compare the single specimen that he found in the American Museum of Natu- ral History.” This was an ivory object, identified as a whip handle,” and bearing a decoration so different from anything known to the Eskimo that Dr. MacCurdy published a brief description of it in the American Anthropologist. In all, probably 30 objects showing the old Bering Sea decoration have been illustrated and described. Other examples of it are in various museums and private collections, and some of these will be described in the following pages. OBJECTS REPRESENTATIVE OF THE OLD BERING SEA CULTURE It will be understood that all of the objects to be described are of walrus ivory unless otherwise stated. In color they range from cream, through buff and brown, to a dark green or even black. This dis- coloration has resulted from the ivory having remained buried in the frozen ground for many years. An occasional artifact was shaped from a piece of mammoth ivory or old walrus tusk that had been washed up by the waves, but most of them were carved from the fresh walrus ivory and have since taken on their rich coloring. On plate 1 are shown four harpoon heads embodying the features which may be regarded as typical of the most highly developed and apparently the oldest phase of the anctent Bering Sea art. Plate 1, a-b, is an exceptionally fine harpoon head owned by Messrs. Wilfred and Albert Berry of Seattle. Its provenience is not known except that it came from northern Alaska. * Mathiassen, Therkel, Some Specimens from the Bering Sea Culture. Indian Notes, Museum of the American Indian, Heye Foundation, Vol. 6, No. 1, pp. 33- 56, January, 1920. * An Example of Eskimo Art, Amer. Anthrop., Vol. 23, No. 3, pp. 384-385, 1921. * This appears, however, to be an adz handle. Cf. Mathiassen, 1920, p. 4I. 4. SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 81 The two heads, c and e-f, are also from northern Alaska, the exact locality unknown. They are in the Washington State Museum and I am indebted to Mr. F. S. Hall, Director of the Museum, and to the Messrs. Berry for the kind permission to figure their specimens. The heads are all of the closed socket type, with the line hole at right angles to the blade slit. The basis of the design consists of pairs of circles, at times somewhat elliptical, drawn free hand and surmounting low rounded elevations in such a manner as to suggest eyes on the head of a bird or mammal. There are two of these “ heads,” one at the terminal barb and another just above and in part overlapping it. In c there is a suggestion of a third “ head ” above the line hole. Plate 1, e-f, differs from the others in having a groove for a lateral blade on each side above the line hole. These harpoon heads are of the same form and style of ornamentation as one described by Mathiassen.’ Besides the circles, which form the central motive, there is a graceful arrangement of lines, some deeply and some lightly incised, straight and curved, solid and broken or dotted. Small spurs are also attached to some of the lines and circles. In plate 1, d, is shown an unusual harpoon head which I bought from an Eskimo on St. Lawrence Island. It was excavated from the old village at Sevuokok (Gambell) on the northwestern end of the island. Its ornamentation is of the same nature as the other three, although the careless scratching and unfinished appearance of the upper end distinguish it from most of the objects similarly decorated. There are also deep dots at the centers of the circles. In form, how- ever, this harpoon head presents a number of anomalous features. The line hole, instead of being at the center, is placed at one side. The open socket is rounded instead of rectangular, and in position is more like what would be expected in a closed-socket type. Finally, the grooved band opposite the socket was cut after the decoration had been applied. These facts make it appear that the harpoon head was intended to have, and may originally have had, a closed socket, but that either in the drilling or subsequently while in use, a part broke away, leaving the socket exposed. In order that it might still be utilized the groove was then cut around the side and the foreshaft lashed on. Figure I is a very interesting closed socket harpoon head from Plover Bay, Northeastern Siberia. It is of fossil mammoth ivory and is creamy gray in color in contrast to the usual brown of the old walrus ivory ; it is badly pitted on the opposite side but enough of the ornamentation remains to show that the design was identical on both 1TIndian Notes, Vol. 6, No. 1, fig. 13, a. NO. I4 PREHISTORIC ART OF ALASKAN ESKIMO—COLLINS 5 sides. In ornamentation it differs from those shown on plate 1 (except d) in having a dot within the elevated circles and in having two small cross-hatched areas on the terminal barb. The lines con- nected with the circles are also somewhat more finely incised and are applied with more precision. It differs from the first four specimens in having three barbs at the base instead of one, in this respect being similar to another type of the old Bering Sea harpoon head, examples of which are shown on plate 2. Fic. 1—Closed socket harpoon head of fossil mammoth ivory from Plover Bay, Northeastern Siberia. Plate 2, e, was collected by Mr. T. S. Scupholm from the deserted village of Kukuliak on the north side of St. Lawrence Island. It has an open socket for the foreshaft and rectangular slots for lashing. The line hole is parallel with the blade slit. On the terminal barb has been left a rectangular projection which gives it an appearance some- what similar to figure 1. The central barb and two smaller ornamental remnants of barbs below it are features around which the decoration centers, and are to be compared with similar slight projections along the sides of a, c, and e, plate 1. The decoration consists of straight and curved lines, including very lightly incised broken lines. There 6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 81 are no circles, but instead two (originally four) small eye-shaped designs at the center. Plate 2, a-b, and f, are from the Washington State Museum; the exact localities from which they came are not known. The first is very similar in shape to e, the only difference being the more irregular outline of the terminal barb. The ornamentation is reduced to only a few simple lines. Plate 2, f, is a small harpoon head of a peculiar type, of which four examples have been described previously: one by Wissler,’ from Cape Smythe near Point Barrow; two by Jenness* from the Diomede Islands, and one by Mathiassen® from Banks Island. The features common to harpoon heads of this type are three terminal barbs, the central one being the longest ; a long open socket that reaches to about the center ; small side blades of stone, either parallel or at right angles to the shaft socket ; and most striking of all, where the slots for the lashing come to the surface on the upper side, a deep sunken area at either end of which is a circular perforation also for lashing. In the present specimen the upper perforation has been started from both sides but has not been drilled through. The decoration, which is very similar to the other specimens of the type described, consists of nucleated concentric circles and finely incised lines. The decoration on c-d, also from the Washington State Museum, consists of straight and curved lines and concentric circles, but no dotted lines. It differs from the other two in having a rectangular sunken area around one side to hold the lashing in place, instead of a second slot through which it usually passed. There is also a smaller barb, undecorated, opposite the larger one. In g and h are shown two undecorated heads from St. Lawrence Island owned by Mr. C. L. Andrews of Seattle. They have open sock- ets, no slit for an end blade, two and three terminal barbs, and on each side just above the line hole a deep groove for holding a side blade. Plate 3, b, is a box handle, Washington State Museum collection, from northern Alaska. It is ornamented with four pairs of raised elliptical “ eyes’ with small holes sunk deep into the centers. They are separated and encircled by flowing lines. Plate 3, a, is a similarly decorated box handle that I bought at Gambell, St. Lawrence Island, where it had been excavated. This * Anthrop. Papers Amer. Mus. Nat. Hist. Vol. XIV, Pt. II, p. 410, 1916. * Ann. Rep. for 1926, Nat. Mus. Canada, pl. XIII, a; and Amer. Geogr. Soc., Special Pub. No. 7, fig. 3, c. * Indian Notes, Vol. 6, No. 1, fig. 13, b. Probably not from Banks Island. See footnote, p. 36. NO. 14 PREHISTORIC ART OF ALASKAN ESKIMO—COLLINS i specimen is of bone and the five pairs of circular and elliptical “ eyes” are not raised to the same extent as those on the smaller ivory handle but the elevations can be plainly felt by running the fingers over the surface. Plate 4, a, is an object of unknown use from the Washington State Museum collection, reported to have come from Nelson Island. A similar specimen, from the Diomedes, is figured by Jenness.* The Washington specimen is of beautifully stained, creamy brown ivory and bears a comparatively simple ornamentation of circles, dots and straight lines, with a small unconnected figure made up of curved lines toward the end. The centers of the circles are small cylinders of ivory surrounded by rings of baleen. Two baleen discs were inserted above these on the upper side of the object. The two pairs of large circles and appended lines, together with the contour of the surface to which they are applied, produce an effect strongly suggestive of seal heads. There are two perforations on the lower side, and between these a deep, rounded notch, indicating that the object was intended to be lashed to something. The two heavy lines extending upward from the basal notch are deeply cut, leaving the lower edges of the seal heads slightly overhanging. The decoration on the opposite side is identical except that instead of the detached curved figure on the wing there are three concentric arcs. Plate 4, b, is a similar, though cruder, object from St. Lawrence Island. It is owned by Messrs. Albert and Wilfred Berry of Seattle. _ It has no features suggestive of life forms like the preceding specimen, but like it has a deeply cut groove beginning at the notch. A single large circle near the center on each side, and a few small circles and slightly curved lines are the only ornamentation. The specimen shown on plate 5 is the property of Rev. C. K. Malmin of Ketchikan, Alaska. It was found at Imaruk Basin, east of Teller, Seward Peninsula. The front end is carved to represent an animal’s head, the long sharp canines indicating a carnivore. The ivory is mottled in rich shades of brown, cream, and gray. The principal design is placed at the center between the two large holes. This is divided into two parts by a deep, curving groove. The front design, ranging about the forward perforation, consists of rather deeply incised lines with occasional spurs and a curved petaloid figure within which is a small circle from which descend two short converg- ing lines. This figure resembles the flat ivory hat ornaments, some- times representing gulls’ heads, used to decorate the wooden hunting *Ann. Rep. for 1926, Nat. Mus. Canada, pl. XIII, b. 8 SMITHSONIAN MISCELLANEOUS COLLECTIONS voL. 81 hats in Southwest Alaska. On the other side of the deep groove the design is continued by more lightly incised lines and a central ovoid figure outlined by small spurs and enclosing a small circle set between two pairs of short converging lines. The decoration is the same on the opposite side. On the upper surface are two drilled holes 5 mm. deep representing the nostrils and further back two representing the eyes. There is a fifth hole near the end and two very shallow ones just back of the eyes. Beneath the nostrils and on the edge of the lower lip are a few slanting lines and spurs. At the rear the upper part of the head is set off by a rounded curv- ing ridge or lip, immediately above which is a deep groove. Above this are a few short lines and spurs, and below it on the curving handle-like rear portion is a simple pattern consisting of straight and curved lines, spurs and small circles at the inner angles of converging lines. Of all the objects that have been described from the old Bering Sea culture this one is the most suggestive of the Northwest Pacific coast. It is also similar to the animal heads found so frequently in the Kuskokwim region of southwest Alaska (see pl. 21). On plate 6 are shown two views of a remarkable object of unknown use, from Point Hope, collected in 1880 by Capt. E. P. Herendeen. This belongs with the class of objects described by Gordon* and Mathiassen.” It has two symmetrical, beautifully carved wings and a central section, in the base of which is a square excavation 15 mm. deep, probably for receiving a handle. The front or flat surface is the more elaborately decorated. Both wings are divided into three sections by deeply cut oblique lines, within which are placed nucleated con- centric circles, four to each wing. The circles are slightly elevated with small round holes about 3 mm. deep at the centers. Tangent to each circle are two pairs of finely incised parallel or converging lines, while between these, on the outer arc of each circle, are three equi- distant spurs. The design is completed by additional straight and curved lines, so placed as to accentuate and utilize to best advantage the angles and curves that give to the object its peculiarly graceful outline. The designs on the two wings are as nearly identical as is possible. The artist has achieved a pleasing effect by applying, even to the most minute detail, a perfect bilateral symmetry of form and Gordon, G. B., The Double Axe and Some Other Symbols. The Museum Journal, Univ. of Penn., Vol. VII, No. 1, 1916, figs. 99, 100, 105, 100. * Indian Notes, Vol. 6, No. 1, pp. 43-46. a ee ee ee ean o NO. I4 PREHISTORIC ART OF ALASKAN ESKIMO—COLLINS 9 design. In the top of the central wedge-shaped section between the wings is a narrow triangular incision 5 mm. deep. The circle at the center is the only one that is not raised. Some of the more finely incised lines have been almost obliterated by wear, and at several places, especially at the end of one wing, the grain was defective and has chipped off, marring to some extent the design. On the reverse the central portion is raised, or rather, the wings are cut down from it but at a sufficient distance so as to leave two flanges or shorter wings; through both of these are drilled oblique circular perforations 7 mm. in diameter. The ornamentation consists of lines and circles of the same general character as those on the op- posite side, but they are much more worn down. This masterpiece of Eskimo art could hardly have had a practical use ; it was no doubt employed in some ceremony, probably connected with whaling, as Gordon suggested, or perhaps as a charm used by the boat captain to bring success in the hunt. I do not think it likely, however, that there is any genetic connection between this class of objects, found only in Alaska, and the well known prehistoric banner stones from the eastern part of the United States. It is true that this particular specimen and those figured by Gordon are somewhat similar to certain of the banner: stones, but, as will be seen later, this is only one of several forms that occur in Alaska, the others assuming shapes quite unlike anything known from the United States. Furthermore, the enormous area between Northwestern Alaska and the Great Lakes where no such objects are found makes it seem extremely improbable that the two classes of objects are related in origin. On plate 7, a-b, are shown two views of a broken object of the type just described, with both wings missing. This was purchased in Seattle and the locality in Alaska from which it came is not known. It is very similar to the specimens from Point Barrow and East Cape described by Mathiassen.’ The centers of the circles are 4 mm. in diameter and from 6 to 9 mm. deep, and may originally have had insets of some other material as in the object shown on plate 4, a. There are also five small nucleated circles, each set at the inner angle of two straight lines, to some of which are attached small spurs. The lines are closely applied, covering practically the entire surface. The decoration as a whole, with its combination of straight and curved lines, spurs, and circles, is of the same type as that shown on the three harpoon heads on plate 2, c-d, e, and f. *Indian Notes, Vol. 6, No. 1, pp. 43. SR Rae a = rs SSS. IO SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 81 In the base is an oval slot 21 mm. long and 15 mm. deep, which appears to have been cut rather than drilled. Slots of about the same diameter extend up through each flange and are connected near the center by a small circular perforation. At the top of the central projection is a long shallow notch which is a characteristic feature of these objects. The broken harpoon socket piece shown on plate 7, c, was excavated at Gambell by Mr. Otto W. Geist, for the Alaska School of Mines. The decoration consists of nucleated circles connected by lines, the pattern being enclosed within a field set off by a deeply cut line. There, was evidently a somewhat similar design on the other side of the shallow sunken band which encircles the object. The space between the two design elements at the bottom has been cut away, leaving the rounded ends in low relief, as was seen also on the harpoon heads shown on plate 1. Plate 8, a-b, is an object of unknown use, collected at Point Hope by Henry D. Woolfe in 1885. It is of dark greenish-brown ivory and is carved to represent a seal. There is a slot at the back end 20 mm. long which continues on the underside for an additional 25 mm. This suggests a slot for a blade, but the central perforation, with grooves leading down to a flattened base, is evidently intended for attachment, in which case it is difficult to see how the object could have served as a knife or other cutting implement. The object is of particular interest, however, for the reason that, despite the absence of any etched designs, certain features of the outline are sufficient, quite apart from the patination, to show that it is a product of the old Bering Sea culture. Extending from below the neck to the middle of the back are two wide and deep grooves, the edges of which are bor- dered by lightly incised lines. The incision through the head is bor- dered in a similar manner. Around the ends of the grooves on the back is a somewhat deeply incised curving line bordered on one side by a ridge or lip. Such ridges, although more pronounced, may be seen on plate 5 and on plate 8, c. The eye is formed by a circular excavation 5 mm. in diameter set at the center of a large slightly sunken circu- lar area. Freshly cut discs of wood and small black glass beads have been recently inserted in these cavities. The broken object shown on plate 8, c, was purchased in Seattle; its provenience is not known. It is evidently part of a harpoon socket piece of the type illustrated by Mathiassen* from Point Hope and Kotzebue Sound. A round hole 6 mm. in diameter is drilled trans- * Indian Notes, Vol. 6, No. 1, p. 39. NO. I4 PREHISTORIC ART OF ALASKAN ESKIMO—COLLINS iE versely through the object, emerging at both sides immediately below the rounded projections at the somewhat constricted center. Cut into the sides and leading down from this perforation are two very narrow grooves similar to the groove in one of the specimens from Kotzebue Sound figured by Mathiassen. The rounded lip-like projection at the center may be compared with the raised border on the preceding specimen and the others first mentioned. The lightly incised lines forming the decoration have been almost entirely effaced on the two flat surfaces but still are faintly visible along the sides, though not in the photograph. The pattern consists. of concentric circles with small cylindrical holes at the center 5 mm. deep. About these are con- tinuous and broken lines and small spurs. The objects shown on the preceding plates illustrate the various known forms of the ancient Bering Sea art. The features character- istic of this art may be outlined as follows: (1) nucleated circles and ellipses, usually concentric and often surmounting low rounded elevations ; (2) the arrangement of these on certain objects so as to suggest the eyes of an animal; (3) deeply excavated centers to some of the circles, sometimes inset with discs of baleen or other material ; (4) small circles at the inner angle of two converging lines; (5) spurs attached to the circles and lines; (6) straight and curved lines, singly or in bands, serving usually to accentuate the circles and fill in the vacant spaces; (7) finely incised broken or dotted lines; (8) an occasional checked or hachured area; (9) raised borders to grooves and rounded lip-like projections. When we turn to consider the shapes and the surface elevations and depressions of the objects it is seen that they also form an im- portant part of the decorative scheme. The notches in the wings of the specimen on plate 6, for example, are the beginning of the sections or panels into which the wings are divided and within which the lines and circles are so gracefully arranged. Similarly, the entire rounded end of the wing is set off by a curving line enclosing and unifying the design. The circles on the terminal barbs of the harpoon heads, such as those shown on plate I, are placed in relation to the shape of the barb itself to suggest eyes, and the effect is repeated in the pair im- mediately above when a small triangular area is cut away, leaving a rounded and slightly elevated “nose.’’ The same principle may be seen in the specimens on plate 4, a, and plate 7, c. In short, it may be said that the old Bering Sea art is marked by a profuse but ex- tremely graceful application of lines, curves, and circles in such a way as to realize to the utmost, within the limits of the accepted patterns, the artistic possibilities of the surfaces to be covered. I2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 81 The ivory specimens bearing ornamentation typical of the old Bering Sea culture are quite uniformly patinated and discolored. It is natural to inquire as to the affinities of this new and distinctive type or phase of Eskimo art, which is apparently so different from that of the modern Eskimo. It has been suggested that it may be related to the art of the Northwest Coast Indians or the Amur tribes of north- eastern Asia. These are the two areas of highly developed art nearest the Bering Sea region and might naturally, therefore, be looked to as bearing some possible relationship to it. As Jenness has pointed out, however, the closest resemblance seems to be to Melanesia, with which there is no reason whatever to assume any relationship. I would suggest further that the resemblances here are more apparent than real. The old Alaskan pieces that most closely approach the Melanesian are those in which a series of circles or ellipses are sepa- rated and bordered by curving lines, giving to them a semblance of scroll work. It is important to note, however, that the circles are al- ways complete and that the lines likewise are completely attached and never left with a free curving end. With the two other regions in question, namely, the Amur and the Northwest Coast of America, I can again see no real resemblances. The art of the Amur tribes, with its ornate and highly conven- tionalized animal and floral forms, abounding in intricate spirals and panels of continuous curving figures in maze-like patterns, appears to me to be totally unlike the ancient art of the Bering Sea. In the same way we will search in vain for any real resemblance to the peculiar art of the Northwest Coast. The Bering Sea art consists essentially of circles and lines, and the designs show no internal evi- dence of ever having been associated with realistic patterns, although carved representations of animals are not lacking (see pls. 5 and 8). The Northwest Coast art, on the other hand, is more solid and compact and, inextricably linked with totemic and other cultural concepts, is marked by symbolic and conventionalized representations of animal forms. The Bering Sea art is graceful and comparatively simple; the Northwest Coast art is heavy and complex. The only characteristic elements of Northwest Coast decoration that might be compared di- rectly are the eye motives and the cross-hatched surfaces, for occasional oval figures with spurs attached, suggestive of eyes do occur in the old Eskimo art, and, infrequently, very small cross-hatched areas. The eye, however, is only a natural variant of the prevailing simple circle or ellipse, having resulted from the combination of this with another typical feature, the spur ; while in the Northwest Coast the eye effect is produced by a continuous line enclosing a solid figure. NO. I4 PREHISTORIC ART OF ALASKAN ESKIMO—COLLINS 13 Boas regards the Northwest Coast eye as a development of the circle, and here arises an important theoretical point; for if it should be true that this highly important element in Northwest Indian art arose from a design which forms an even more important part of the old Eskimo art, we would have valid ground for considering that the art of the two regions was historically connected. But, assuming such a connection to have existed, there would still be no more reason for considering that the cultural impetus had been exerted from south to north than from the opposite direction. It must be remem- bered that the Eskimo culture under consideration is ancient; just how ancient, it is yet too early to say, but at any rate it is the oldest culture that has come to light in the extreme Northwest. It is, further- more, closely related, apparently ancestral, to the very wide-spread Thule culture which, in early times, extended across north central Canada into Greenland, and which has been so thoroughly revealed through the recent researches of Mathiassen. On the other hand, there is no good reason for assigning an equal antiquity to the peculiarly local and highly developed art of the present tribes of the Northwest Pacific Coast. It seems fairly evident, viewed solely from the ethnological stand- point, that certain culture traits now in the possession of the Alaskan Eskimo were derived from the Alaskan Indians. But this, along with a possibly similar condition in regard to physical type, may well have been of a secondary nature. At any rate, it does not seem possible at the present stage of our knowledge to point to any particular aspect of the earlier Eskimo culture that might be said, with any degree of certainty, to have been similarly derived. In spite of the absence of any important specific resemblances between the art of the two areas, there still remains a vague, general similarity which may lead to the expectation that future archeological work may reveal an earlier stage of Northwest Coast culture closer to the ancient Eskimo culture of Bering Sea, or even, somewhere, a culture that may have been ancestral to both the Northwest Coast and Eskimo cultures. In this connection there is also the possibility that very important results might come from investigation of ancient sites in Southwest Alaska, or around Prince William Sound where Eskimo territory impinged on that of the Tlingit. While it is difficult to trace true relationships of the newly found ancient Bering Sea culture with Indian tribes of America or tribes of northeastern Asia, it is plainly evident that there is an unbroken line of succession where such would be most expected, namely, in the Eskimo matte I4 SMITHSONIAN MISCELLANEOUS COLLECTIONS voL. 8&1 territory of Alaska. With all its relative richness and undoubted superiority over the present Eskimo art there can be no doubt that the old art was the direct forerunner of the modern. At first glance there may appear to be little resemblance between the artistic and flowing ornamentation of the old Bering Sea culture and the familiar and much simpler designs employed by the modern Eskimo. But there are sufficient links to bridge the gap. It would be possible, I believe, to trace the development of the old Bering Sea art into the modern if we had no more than the objects shown on plates 1 to 8, and those described by Jenness, Mathiassen, and Hrdli¢ka. The transition would be somewhat sudden, it is true, and some of the designs might appear to be but little related. Fortu- nately, however, we now have additional data that throw light on a secondary stage of art development within the old Bering Sea culture. This new evidence makes it appear that the old Bering Sea art did not come to a sudden end, to be succeeded immediately by that of the modern Alaskan Eskimo, but that, on St. Lawrence Island at least, it entered upon a period of transition during which the designs became simpler, definitely foreshadowing the later Alaskan Eskimo art. Some of this material will be described in the following pages, and the conditions under which it was found, on Punuk and St. Lawrence Islands, will be briefly outlined. THE PUNUK ISLAND AND CAPE KIALEGAK VILLAGE SITES The three small Punuk Islands lie four miles off the southeast end of St Lawrence Island. The largest island, on which the old village site is located, is slightly less than a half mile long and only a few hundred yards across at its widest point. The greater part of the island is covered with the usual tundra vegetation and is relatively flat, except for two rocky hillocks that rise suddenly from the southern side. To the west the island narrows and the tundra is replaced by a low sandy area covered with coarse grass. At the beginning of this sandy stretch, which is also the narrowest part of the island, is located the ex- tensive kitchen midden which marks the site of the old village. The last houses to be occupied were sunk into the top of the midden. They consisted of square excavations in which had been erected frameworks of whale bones and driftwood logs; there are 14 house pits besides numerous underground caches on the midden. The nar- row tunnel-like entrances to the houses faced the sea, sometimes to the north, sometimes to the south. Of the logs which formed part of the framework scarcely any remain on the surface, but whale ribs and jaw bones are more plentiful. NO. 14 PREHISTORIC ART OF ALASKAN ESKIMO—COLLINS 15 The area covered by the Punuk midden is approximately 400 feet by 130 feet. The average depth is around 12 feet and the greatest depth, 16 feet. It is of particular interest to note that the lower parts of the midden are below the present beach level—at one place as much as six feet—and that at the very bottom were found houses and house entrances, six in number, all of which were below the reach of storm waves. The sinking of the land to this extent, together with the enor- mous accumulation of refuse, must undoubtedly indicate the passage of a considerable period of time. Asa definite criterion of age, how- ever, stich a geological phenomenon is of questionable value at the present time, since there are not available any comparative data on recent subsidence and elevation in the Bering Sea region which might be interpreted in terms of years. The problem of the antiquity, at least the comparative antiquity, of the Punuk village site must rest mainly on cultural evidence. To the west of the midden, near the end of the island, are four recent houses with roofs still partly intact, which were occupied up to about 40 years ago. It is not clear whether these were built by fami- lies who settled on Punuk after the abandonment of the main village or whether they were occupied by the last remnants of the original population. Excavations were carried on at Punuk from June 23 to August 17, 1928. Assisting me were Mr. Harry E. Manca of Seattle and two, later three, Eskimos from St. Lawrence Island. Three of the recent houses toward the end of the island were excavated and two of the older houses on the midden, in addition to extensive cuts made through the refuse at various places. The midden sections and the two old house pits on the midden were taken down systematically, the material from the successive levels being kept separate.’ Except for a surface layer which thaws out in summer, the ground is permanently frozen, making the work of excavation difficult and slow. The cuts were taken down in lavers of a few inches, the process being repeated daily as the newly exposed frozen surface thawed by contact with the atmosphere. The many specimens excavated from the midden village stood in striking contrast to the material from the three recent house ruins at the western end of the island. The latter yielded iron, glass, and only 1In the following description of certain decorated objects from Punuk Island and Cape Kialegak, reference will not be made to the location and depth at which individual specimens were found, as this would only add unnecessary detail and call for a fuller description of the sites and method of excavation than is necessary at the present time, when the purpose is to describe only the art of the two sites. 2 16 SMITHSONIAN MISCELLANEOUS COLLECTIONS voL. 81 modern types of artifacts of ivory, bone and wood, identical to those still being used on St. Lawrence or having been used there in recent years. At the old village, however, among the several thousand specimens excavated, there were only four small fragments of iron and two glass beads, although every inch of the soil was gone over with trowels. There was likewise not a single modern type closed socket harpoon head found, and the blades of the harpoons, lances, knives and adzes were all of slate. However, the presence of even these few pieces of iron and, more significant even, an occasional file mark on some of the specimens, indicates that at least during the later years of their stay on the island the people of the Punuk village were in possession of small quantities of European metal. Late in July, I made a brief trip in an Eskimo whale boat to Cape Kialegak on the southeastern end of St. Lawrence Island where there was a deserted village with a kitchen midden even higher than the one at Punuk. The Kialegak people had occupied two villages; the older and smaller village was entirely prehistoric, judging from the objects dug from the midden, while the later and more extensive settlement only a few hundred yards distant had been established ap- parently at about the time of the abandonment of the earlier village and occupied until about 40 years ago. Proof of this was found in the midden, the lower levels of which yielded only ancient types of arti- facts, including harpoon heads with open sockets and some with side blades, while in the upper levels, beginning at about 10 feet, were implements of the modern type accompanied by glass beads and nu- merous pieces of iron. The objects from the lower levels of the later village and all of those from the older site were of the types we had been finding at the old Punuk village. Cape Kialegak thus afforded valuable supplementary evidence, for the occupancy of the later vil- lage began at a period contemporaneous with Punuk but continued without break until recent years, whereas the old village on Punuk was abandoned at some unknown time within the proto-historic period and was succeeded, perhaps after a considerable interval, by the few recent houses at the end of the island. This will serve as an outline of the conditions under which the specimens to be described were found and removed, and no further details of the procedure will be entered into at this time. Instead, discussion will be limited, with a few necessary exceptions, to the decorative designs on the objects themselves, to their relation with those previously described and those of the modern Eskimo, and to the bearing that they are believed to have on the larger problem of the NO. I4 PREHISTORIC ART OF ALASKAN ESKIMO—COLLINS V7; spread and relationships of early Eskimo culture in the Bering Sea and elsewhere. On plate 9, a, b, c and d, are shown four fragmentary harpoon heads from Punuk and Cape Kialegak, the only specimens from these localities that belong, from their decoration, with the type of material illustrated on plates 1 to 8. Plate 9, c and d, are two closed socket harpoon heads, both water-worn to such an extent that the designs are almost obliterated. The harpoon head, c, has the same ornamen- tation as those shown on plate I, but unlike them the blade slit runs parallel with the line hole. The circles are raised. Plate 9, d, in ad- dition to being badly worn, shows evidence of having been trimmed down, the only part retaining the original decoration béing that shown in the illustration. The remaining circle is elevated. This specimen Fic. 2—An ivory object of unknown use from Punuk Island. was found at the base of the larger Kialegak midden. The two frag- ments, a, and J, also bear curvilinear decoration. These, as well as c, were found at depths of from three to ten feet in the Punuk midden. With the exception of these four fragmentary specimens another and quite distinct type of decoration was found to prevail on Punuk and at Cape Kialegak. This occurs in such abundance and so nearly in isolation at these sites that it seems proper to refer to it as the Punuk type or phase of the old Bering Sea art. Plates 10 to 15 illustrate its various aspects. The exact distribution of the decorated objects from Punuk and Cape Kialegak is as follows: old curvilinear, 4 (those shown in pl. 9, a-d) ; Punuk type, 117 ; modern, 7 ; indeterminable, 13. The last group comprizes objects decorated so carelessly, or of which such a small fragment remains that it could not be determined whether they were of the Punuk or the modern type of ornamentation. In figure 2 is shown an ivory object of unknown use found at a depth of 13 feet in the Punuk midden. It is thin and convex, and in 1g SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 81 general shape is somewhat like the St. Lawrence Island wrist guard, although it has no slots or other openings through which a thong might pass except the handle-like loop at the top. This specimen is of particular interest in that the design combines features typical of the several stages of art in the Bering Sea region. The three petaloid figures are most suggestive of the older phase, while the deeply in- cised parallel lines in bands of four and the Y-shaped figure are char- acteristic of the Punuk and recent stages. The spurs attached to the lines and curves are common to all three stages, having been retained in the art of the Alaskan Eskimo from the earliest known times to the present. Plate 10, a-b, represents an object of unknown use gracefully decorated in the style typical of Punuk. One wing is broken off. A complete specimen similar to this is shown on plate 13, f, from Cape Kialegak. In the present specimen there is a rectangular socket in the base 16 mm. deep which was made by drilling. From the socket a round hole 7 mm. in diameter extends through the base of each wing. The depression at the top of the central upright section and the socket at the base are features that were seen also in the object on plate 6. Lines terminating in dots, short converging lines enclosing dots, and small squares form the design, which is often found recurring in the Punuk style of the old Bering Sea art. The designs on the two sides are made continuous by the single connecting line that crosses over at the center of the wing on the inner side. The object from Point Hope’ illustrated by Mathiassen is very similar in outline and practically identical as to the general style of ornamentation. In form it is intermediate between the present specimen and that shown on plate 6. The two objects shown in c-d and e, plate 10, while not from Punuk Island or Cape Kialegak, are introduced for the reason that they represent a common St. Lawrence type and a variant of the class of objects illustrated by a-b of this plate, a-b, plate 6, and a-b, plate 7, and corresponding specimens described by Gordon, Jenness, and Mathiassen. In c-d are shown two views of one of these objects from St. Lawrence Island, purchased by Mr. H. W. Krieger. Like the specimen just described it has a basal socket and a small de- pression in the top of the central projection which, however, is joined to the wings. This socket is round and only 12 mm. in diameter, and in it is the broken end of the wooden shaft which probably formed its handle. The decoration on both sides consists of deeply and evenly *TIndian Notes, Vol. 6, No. 1, fig. 19. eee NO. 14 PREHISTORIC ART OF ALASKAN ESKIMO—COLLINS 19 incised straight and slightly curving lines, single or in bands of from two to four, with small spurs attached. Red ochre had been rubbed into the incisions and some of it still remains in place. The object shown in e differs only in minor details from the other. It came from Kukuliak, on the north side of St. Lawrence Island, and is owned by Mr. C. L. Andrews of Seattle. These three specimens and the one on plate 6 seem to represent three rather widely variant forms of the same object. All have a basal socket, a central projection in the top of which is a shallow depression, and two wings, the latter especially being extremely variable as to size, shape, and position. My Eskimo workmen were unable to say defi- nitely what had been the use of these objects, but one of them thought they might have been ornaments for the war helmets formerly used on St. Lawrence Island. However, I am more inclined to accept the explanation given by Gordon for the specimens he described—that they are charms used in ceremonies in connection with whale hunting. As to which of these forms is the oldest, the evidence seems to favor the type shown on plate 6. These bear the old Bering Sea curvilinear patterns, while the others are ornamented after the Punuk style, which, as will be shown later, appears without doubt to be more recent, at least on St. Lawrence, than the curvilinear. A further indi- cation that the type shown on plate 10, a-b, is fairly recent is the fact that an unfinished specimen was found in the refuse from one of the last houses to be occupied on Punuk Island. On plate 11 are illustrated a number of artifacts from Punuk and St. Lawrence Island bearing decorations typical of the Punuk style of art, with nucleated circles, straight or slightly curved lines, dots and spurs. Plate 11, a and b, are two broken harpoon heads of the open socket type with slots for the lashing and with line hole parallel with the blade slit. This is the form of harpoon head most common on Punuk although a number of closed socket heads of a distinctive type, always ornamented, were also found. The decoration on the two specimens consists of lines extending from the barb toward the point, terminat- ing in evenly inscribed nucleated circles, and similar lines, together with smaller cross lines and spurs, around the central line hole. The ornamentation of the upper portion, above the line hole, is the same on both sides; but on account of the open socket on the inner or under side, the decoration beneath the line hole is restricted almost entirely to the outer side. The harpoon heads represented by d and e are of the same type as the two preceding. In d the circles are smaller and there are more 20 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 81 lines and spurs, while in e there are more circles on the barb than are necessary to balance the design. These harpoon heads are intermediate in type between those bearing the curvilinear patterns such as illustrated on plates 1 and 2 and the modern, shown on plate 20. The open socket, flat shape, and rec- tangular slots for lashing are old; the decoration, although somewhat profuse, is more rigid and lacks the graceful flowing lines of the earlier period. A pattern of lines, circles, dots and spurs is never present on modern harpoon heads, although circles alone are found on a restricted type from the Nunivak-Kuskokwim region, and crude lines and triangles on others from the north coast of Alaska (see pl. 20). However, in shape as well as design these modern decorated harpoon heads differ essentially from the Punuk type. In plate 11, c, is shown the end of a box handle simply decorated with the circle and dot, which might occur in either the Punuk or the recent period. Plate 11, f, shows the upper end of a dart foreshaft from Kukuliak, St. Lawrence Island. In the top is a cavity 19 mm. deep and 12 mm. in diameter. The object has been used secondarily as a drill or reamer. The surface etching consists of lines, dots, circles and spurs. In g is shown an object of unknown use. It is broken at one end, but the remaining decoration at this point, as well as the shape, sug- gests that there was originally a second wing similarly ornamented. The incisions are deeply and evenly cut and had been filled with red ochre. The design is made up of circles and spurs and straight or slightly curved lines, some of them forming bold Y-shaped figures. The opposite side is not ornamented. The object illustrated by / is a wrist guard purchased at Gambell, St. Lawrence Island. The leather thong is modern. The design is very simple, consisting only of nucleated circles enclosed in panels formed by straight lines. An ivory drill rest with a rather closely applied decoration of lines, dots, circles, and spurs is shown in 7. Comparison of the circles on the objects just described with those shown on previous plates reveals the significant fact that they are mechanically perfect as well as flat, while the circles and ellipses previ- ously shown were usually raised above the surface and were always somewhat irregular, having been made free hand. The Punuk circles were engraved mechanically with an implement of some kind, probably a two-pointed compass of metal. The dots are usually from 2 to 3 mm. deep and the circles slightly less than 1 mm. The extreme pre- NO. I4 PREHISTORIC ART OF ALASKAN ESKIMO—COLLINS 21 cision of the cutting and the uniform depth and width of the lines, circles, and dots, give every appearance of having been produced with steel tools. On the other hand, no metal was found in the Punuk midden except four small fragments in the upper section. Further- more, the great number of slate blades for harpoons, lances, knives, and adzes proves beyond a doubt that the Punuk people still depended on stone for their cutting, thrusting, and hewing implements, even though they may have possessed a few treasured tools of iron with which they decorated their ivory and bone implements. There is thus reason for believing that the Punuk settlement dates from the time when small quantities of European metal were first obtained through the Chukchi and Siberian Eskimo following the arrival of the Rus- sians in Northeastern Siberia in the seventeenth century. On plate 12 are illustrated eight specimens which differ from those on the preceding plate in that the decoration is made up of lines and dots instead of lines and circles. In a, b, and c are represented three different types of harpoon heads bearing a similar ornamentation. The first two, a and b, have closed sockets, but b is flat and has the line hole parallel with the blade slit, while in a the line hole is at right angles and there is a sharp high longitudinal ridge on both sides above the line hole, as was seen also on plate 1. It is an interesting fact that every closed socket harpoon head found at the old sections of Punuk and Cape Kialegak was decorated, whereas the more common open socket heads were usually undecorated. Plate 12, c, is of bone; it has an open socket, rectangular slots for lashing the foreshaft in place, and two deep slots for side blades. These were of shell and the lower parts are still in place. The slight projection brought about by cutting away a section of the edge at the end of the outer line below the lateral blade slots is a feature that was observed on several of the harpoon heads on plates 1 and 2. The designs on a and 0 are in general very similar, despite the dif- ference in shape of the two harpoon heads. Both have incised lines on the barb, around the line hole, and up toward the point. On 0 the dots are placed at the end of short lines while on a they are free; a also has spurs attached to the lines. The dots on ¢ are connected with the lines, though not always at the ends as in 0, and the spurs are shorter and more numerous than in a. The object represented by d is broken at both ends and hollowed out like a spoon on the opposite side. The lines are more curved than those on the harpoon heads just described. The dots are almost 4 mm. deep and three of them pass completely through the specimen. { i } { { 22 SMITHSONIAN MISCELLANEOUS COLLECTIONS voL. 81 The terminal barb of a large closed socket whaling harpoon head is shown in e. The arrangement of dots and short lines within a long two-pronged figure is common on the smaller seal and walrus harpoon heads of Punuk and Cape Kialegak. In f is shown an ivory object of unknown use decorated only on one side. The dots are somewhat more numerous than on most of the specimens. Two pairs of pronged figures enclosing four dots are attached to the central band where the sides gradually widen. Between these are six small dotted squares, such as are also found in modern Alaskan Eskimo art (see pl. 18, a). In g is shown an object which may possibly have been one section of a double knife handle. The under side is flat and at the large end is a rectangular groove 33 mm. long, 7 mm. wide and 12 mm. deep, too large to serve as a slot for any but a very thick stone blade. A wide sunken groove extends around the surface for the purpose of lashing. There is a narrow rectangular perforation at the lower end. The decoration is a simple pattern consisting of a large triangle with two Y-shaped figures terminating in dots, three vertical lines with similar dots, and two plain oblique lines. A very slender harpoon socket piece or foreshaft, probably for a toy harpoon, is shown in h. At the upper end is a small round hole 4.5 mm. deep for receiving the foreshaft or dart head. Somewhat above the center is a small rectangular perforation. The ornamenta- tion consists of lines and spurs, the latter being attached only to the curved lines which enclose the pattern at both sides. Within these are three pairs of straight lines meeting at acute angles, with the lower ends coinciding with slight bulges along the sides. Plate 13 illustrates seven artifacts from Punuk Island and Cape Kialegak on which the decoration is restricted to lines and spurs. A closed socket harpoon head with blade slit parallel with the line hole is shown in a. The tip of the barb has been roughly cut off, leaving serrations having somewhat the appearance of inverted barbs. The lines and spurs are incised very deeply and still contain some of the red ochre with which they were formerly filled. In addition to the absence of circles and dots the pattern differs from those on the har- poon heads previously described in having a greater number of lines on the lower portion and in having short cross incisions at several places. The second head, b, while an open socket type, has the same ar- rangement of blade slit and line hole and bears essentially the same decoration. It also has red ochre rubbed into the deep incisions. NO. I4 PREHISTORIC ART OF ALASKAN ESKIMO—COLLINS 23 The broken object shown in c represents a type of artifact of which five specimens are known thus far, three in the present collection and two in the collection of Mr. C. L. Andrews; one of the latter is shown on plate 14. In the present specimen there were originally five trans- verse perforations through which lines may have passed, a circular one at the center and each end, with two rectangular ones between. The rectangular groove at the end suggests that the object was attached to something by lashing. This groove was cut before the decoration was applied, for the lines stop just before reaching the edge. The surface rises gradually toward the center where there is a small cylindrical hole bordered by a circle and four long spurs. Around this inner circle are three larger rings, all with spurs attached, then two others which are not completed but which extend from the two central notches toward the ends; and finally, beyond the groove at the end— the other end is broken—a similar curve and three long spurs. In d is shown another object for which I can suggest no use. It is slightly convex and has been broken and partly smoothed off at the larger end. At the smaller end is a circular hole which passes through the object. The two holes at the center are 6 and 8 mm. deep. Toward the large end is another hole in which is the broken end of an ivory plug and just beyond this still another which barely misses meeting one drilled from the under side. The decoration is confined to lines, with spurs, arranged in reference to the projecting and incurving out- line. On the under side the decoration is simpler, consisting of a nar- row band down the center which widens and, at the larger end, divides into two ladder-shaped figures. It is unfortunate that so many of the objects most elaborately decorated have no analogies in modern Eskimo culture from which it would be possible to determine the purpose for which they were made. This applies also to the next specimen, e, a propeller-shaped object with two wings and on the under side a central projection through which is drilled a transverse circular perforation. The designs on the two wings are unconnected ; they differ only in that there is an additional enclosing line on the left side. The under side is plain. The small ornament, f, is of the type shown on plate 10, a-b. The slot in the base is round and g mm. deep and the central projection lacks the customary indentation on the top. The two wings are perfo- rated near the base. The decoration will be seen to be practically identical with that of the other object as to outline, lacking only the dots and such additional lines as were made possible by the greater surface to be covered on the larger specimen. As in that case, the 24 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 81 design here is in two parts, one a connected line reaching to the tip of each wing and descending to the base, and another occupying the up- right central projection, then passing up along the inner sides of the wings to continue on the opposite side in a similar but simplified design. The object illustrated by g is flat on the bottom and is broken at one end, but from the design it may be judged to have come to a point as at the opposite end. The short cross lines in pairs, such as were observed on the harpoon heads, a and b, are here still more prominent. They divide into segments the space within the four long curving bands and serve to tie these together. There are also some- what longer cross lines in the central panel, connected by a single straight line and enclosed at one end by a pair of lines that come to a point near the large circular perforation. This was drilled after the decoration was completed and passes through a double lined triangular figure similar to the one below. On plate 14 is illustrated an object similar to the fragmentary speci- men shown on plate 13, c. It is from Kukuliak, St. Lawrence Island, and is owned by Mr. C. L. Andrews. The five circular and rectangular perforations noted on the former specimen are visible here. At the center, instead of an elevation, is a circular depression. Within this are placed two roughly crescent-shaped figures which touch at the centers. Pendant from these are two short lines ending in dots, and in each crescent a central dot and a short cross line at the ends. En- circling the central concavity is a single line and beyond, in both direc- tions, three additional curving lines. Between these are slightly curved triangular figures enclosing a central dot, their apices pointing out- ward, with the exception of the first one to the right which points back toward the center. On both sides of the triangles are single de- tached dots. The triangles on this object are very similar to those shown on plate 12, d. The harpoon head shown on plate 15, a, has an open socket and as usual the line hole parallel with the blade slit. The incised lines forming the decoration are lightly applied. It has on each side two pairs of lines extending up toward the point and between these other lines that enclose the circular line hole in a narrow triangular pattern. Plate 15, b, is a closed socket harpoon head with the upper end broken away. The blade slit appears to have run parallel with the line hole. The broken end of the foreshaft remains wedged in the socket. The decoration of lines and dots is quite similar to those on the har- poon heads shown on plate 12. The lines are still filled with red ochre. In c and d are illustrated two interesting harpoon heads with en- Sees Se ee - eo eae NO. 14 PREHISTORIC ART OF ALASKAN ESKIMO—COLLINS 25 closed slots for lateral blades ; c is of bone. It has an open socket and a bifurcated terminal barb; it is broken on the right side and may originally have had an additional smaller barb. The slot for the side blades passes completely through. An interesting feature—not shown in the photograph—is that on both sides along the sharp edge opposite and below the blade slot are additional grooves. The upper groove is 2mm. deep and 17 mm. long and below it are two still smaller tri- angular depressions. All of these seem too shallow to have held stone or shell blades, and since there was originally a serviceable blade in the larger central slot these side grooves were probably only ornamental. The decoration consists of a few rather carelessly incised lines. The harpoon head, d, has an open socket and a slot for side blades like c, and one of these, a semi-circular piece of slate with a sharp edge, remains in place. As in c, there are also two additional grooves on the sides opposite the blade slot, and these are so shallow that there can be no doubt but that they are ornamental. The decoration, while almost as meagre as on the preceding specimen, is more neatly applied. It consists of a pair of converging straight lines within which are two Y-shaped figures. The three points to bird darts illustrated in e, f, and g are typical of Punuk Island and Cape Kialegak. The characters that mark the type are the three barbs on one side and two on the other; two lines down the center or just at the base of the barbs; a curving, sharpened base; a rectangular line slot near the barb; and two or three small notched projections along one edge near the base. This last feature is of especial interest since it is present as a decorative motive in modern Alaskan Eskimo art (see pl. 18, c). In h is shown a piece of a box handle bearing a simple but pleasing pattern of large Y-shaped figures with dots at their bases, enclosed within two parallel lines. In 7 is shown the end of an object of the type shown on plate 13, c, and plate 14. It bears a typical Punuk design of lines terminating in dots and with short cross lines forming small rectangular spaces. The object shown in j7 is a fragment of an elaborate wrist guard from Gambell. If we may judge from a similar complete specimen from the same locality, this had a second wing curving in the opposite direction. While the lines follow the contour, the design is very similar to that of plate 13, g, except for the addition of long sharp spurs attached to the pairs of short cross lines. Examples of carving in the round were extremely rare among the finds from Punuk Island. That these Eskimo were capable of excel- lent workmanship along this line, however, is shown by the remarkable 26 SMITHSONIAN MISCELLANEOUS COLLECTIONS voL. 81 ivory figurine illustrated on plate 16. This represents a woman with long torso, prominent abdomen, pendant breasts and very short legs and forearms. An incision has been made in the left breast as if for suspension, and striations are seen across the upper arms and shoul- ders. The well defined curves, especially of the lower part of the body and the breasts, and the realistic treatment of the figure as a whole, produce an effect quite unlike that seen in the simple and stiffly conventional dolls of the modern Alaskan Eskimo. It is, however, somewhat similar to the armless dolls of the Ammassalik Eskimo. More significant even than the scarcity of carving in the round was the total absence on Punuk of the pictographic art that is so char- acteristic of the modern Alaskan Eskimo. Etched realistic designs have not been found at any ancient Alaskan site so that this type of art must for the present be considered as recent. RELATION OF THE PUNUK ART TO, THE CURVILINEART ARLE OF THE OLD BERING.SEA CULTURE We have seen on plates 10 to 15 a number of objects from Punuk and St. Lawrence Islands decorated in the manner characteristic of what it seems proper to call the Punuk phase of the old Bering Sea art, as represented at these localities. It shows significant differences from as well as resemblances to the older phase of this art as repre- sented by the objects on plates 1 to 8 and those figured by Jenness, Hrdli¢cka, and Mathiassen. It will be observed first that both the Punuk and the older Bering Sea cultures abound in highly decorated objects of unknown use. Such are the winged objects shown on plates 6; 7, a-b; 10, a to e; 13, f; all of which are apparently related forms of the same class of highly variable objects, used most likely in whaling ceremonies or as individual charms. One of the most striking differences between the prehistoric and modern Eskimo cultures in Alaska is found in the harpoon head. The only type used in Alaska at the present time has a closed socket. It often has also a small hole in the tip for a rivet which holds the blade in place ; and often a slightly sunken area leading from the line hole to the base. The prevailing type of harpoon head from the old Alaskan sites lacks the last two features and has an open socket, with rectangu- lar slots for lashing on the foreshaft. The harpoon heads from the old villages on Punuk Island and Cape Kialegak are without exception either of this open socket type or belong to a decorated closed socket type related in form to the old closed socket heads shown on plate 1. NO. 14 PREHISTORIC ART OF ALASKAN ESKIMO—COLLINS 27 In form, therefore, the Punuk harpoon heads belong with the old Bering Sea culture. Very few other types of artifacts from old Bering Sea sites have been described, so there is as yet little with which to compare the bulk of the material collected at Punuk and Cape Kialegak. However, the resemblances to the old Thule culture of Canada and Greenland, to which it bears undoubtedly a close relation, and the corresponding absence of many modern Alaskan types, indicate beyond a doubt that the old Punuk and Cape Kialegak culture belongs almost entirely to the prehistoric phase, that is, to a period preceding the actual dis- covery of Alaska by the Russians. A somewhat different condition is observed in regard to the orna- mentation. While the forms of the objects themselves are distinctly ancient the decorations they bear depart radically from the ancient patterns and at times approach very closely the designs employed by the modern Alaskan Eskimo. First in importance, perhaps, is the circle and dot. In the old form this was seen to be always slightly irregular and often elliptical, showing unmistakably that it had been made free hand. In addition, it was usually raised above the surface. The circles of the Punuk period are not raised and are without exception perfectly round, having been made with a compass or bit, probably of metal. In some cases pairs of circles on the barbs and tips of harpoon heads still carry out the suggestion of eyes but the effect is greatly léssened by the addition of straight lines and the absence of the cuts and enclosing lines along the edges that in the older art com- bined to produce the appearance of an animal’s head. Curved lines become much less frequent. Small checked or hachured areas and lightly incised broken or dotted lines are absent. In addition to the compass-made circle and dot the Punuk art brings deeply incised straight lines, often in bands; pairs of short straight cross lines or single cross lines forming small squares or rectangles ; long deeply-cut spurs in contrast to the more delicate and pointed spurs of the old art ; the use of dots applied free or at the end of lines ; bold Y-shaped figures, though not detached; and pairs of serrations on bird dart points occupying a raised border near the base. Besides these specific features distinguishing the two art styles there is a marked difference in appearance due to technique. The old curvilinear designs were deftly applied ; some of the lines were lightly etched while others, for contrast, were deeper. The impression re- ceived is that the artist exercised selective judgment as well as manual skill in the harmonious arrangement of the lines, curves and panels 28 SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL. 81 that make up the composition. The Punuk art, on the other hand, shows no such discrimination. The lines are all uniform and usually quite deep. There is still a slight tendency to utilize the outline of the object for the enhancement of the design but not to nearly such an ex- tent as in the older art. Designs are much more formal and rigid; on harpoon heads, for example, the relatively simple ornamentation is repeated almost exactly over and over again. The same adherence to convention is seen in many of the other objects, resulting in a fixed mechanical style, which though symmetrical and graceful in its sim- plicity, distinctly lacks the elasticity and exuberance that mark the finer products of the older Bering Sea culture as works of real art. While the carving and surface decoration of the older objects bear evidence of high skill there is no reason why they should not be re- garded as the result of cutting with stone tools. There is direct evi- dence, on the other hand, that metal was employed during the Punuk stage. The possibility that the evenly inscribed circles were made with a stone bit is extremely remote. The invariable uniformity in depth and width of the circles shows plainly enough that they could only have been produced by a very narrow, sharp and smooth instrument. The extreme precision of the other lines is evidence in the same direction. Reference has been made to the four fragments of iron found in the upper levels of the Punuk midden and to the file marks on a few of the artifacts. File marks are of more definite value as an aid to chronology than small fragments of iron, for the latter, even after eliminating the possibility of its being of meteoric origin, might still have reached its destination as wreckage, to be salvaged and util- ized by the Eskimo. A file, however, could hardly have come into their possession in such a manner but must almost certainly have been ob- tained, even though indirectly, from a European or, possibly, Oriental source. With the Punuk midden yielding objects from top to bottom (along the outer edges only, however; the bottom of the midden at the center was not reached) showing decorations apparently made with metal tools and with an occasional specimen also showing file marks, we are forced to the conclusion, if the metal be regarded as European, that the Punuk settlement cannot be older than three hundred years ; for it was toward the middle of the 17th century that the Cossacks began to penetrate Northeast Siberia, bringing metal which the Chukchi received and passed on in trade to the Siberian and Alaskan Eskimos. It 1s quite possible that the greater number of objects from the old Punuk site were carved with stone tools; certainly the hun- dreds of stone knife, harpoon, and adz blades found show that stone played an important part in their industries. The first few tools of NO. 14 PREHISTORIC ART OF ALASKAN ESKIMO—COLLINS 29 iron received may have been such prized possessions that they were used only for fine cutting and for decorating implements and ornaments. It is somewhat difficult to reconcile the many outward evidences of antiquity at the old Punuk site with an age of only three hundred years. Even the houses on the top of the midden are now represented by nothing more than shallow levelled pits and fallen whale bones. Wood is absent to a conspicuous degree. The tremendous pile of refuse, reaching a total height of sixteen feet, and most significant of all, old houses at the very bottom of the midden and six feet below the reach of storm tides, would appear without doubt to be prima facie evidence of a considerable antiquity. If the metal could have reached Punuk Island from some Oriental source before the arrival of Europeans in eastern Siberia or in the North Pacific, it would be possible to allow an antiquity to the site more in keeping with its appearance. At present, however, this can be mentioned only as a pos- sibility ; it seems safer to consider the age of the Punuk site, at least provisionally, as not greater than three hundred years. Whatever the age in years of the Punuk site it is without doubt later than the sites from which come the curvilinear art of the old Bering Sea culture. This is indicated by the difference in technique re- ferred to and the fact that the Punuk ornamentation at times ap- proaches very closely that of the modern Alaskan Eskimo. Distribution affords further evidence. The old curvilinear art has been found at Point Barrow, Point Hope, Cape Prince of Wales, Northeast Siberia, the Diomede Islands, Imaruk Basin, St Lawrence Island, and one specimen is reported from Nelson Island. The Punuk type occurs on St. Lawrence and Punuk Islands and one example comes from Point Hope. While the few decorated specimens from mainland sites that have found their way into collections are of the old Bering Sea style, this need not mean that it is the only type present, for with the exception of Wales and the Diomedes we have no de- tailed first hand knowledge of any northern Alaskan sites. For the present, therefore, we must turn to St. Lawrence Island for anything like a comprehensive or comparative view. The old curvilinear art has been found at Gambell on the northwest- ern end of the Island and at Kukuliak on the north side, with traces of it—four out of 141 decorated specimens—on Punuk Island and Cape Kialegak. The Punuk type prevails at Punuk Island and Cape Kialegak and appears to be much more common than the curvilinear at Gambell and Kukuliak. The explanation that appears to best fit these facts is that the northern and western St. Lawrence sites, such 30 SMITHSONIAN MISCELLANEOUS COLLECTIONS voL. 81 as Gambell and Kukuliak, are older, having passed successively through the stage when the curvilinear art flourished, into that of the Punuk type, and finally into the modern. Punuk Island and Cape KKialegak, however, appear to have been settled either after or near the close of the curvilinear period, the cccupancy continuing until recent years. This eastward movement would be in keeping with the reason- able assumption that the first settlements of Siberian Eskimo on St. Lawrence were made at the western end of the Island. By any chance, could the Punuk art have coexisted with the curvilinear? In view of the distribution and considerations of tech- nique this seems very unlikely. The presence of two contemporaneous art styles as distinct as these are, one consisting of deftly incised free hand circles, ellipses, curves and lines, and the other of cleanly cut, straighter, and more rigid designs evidently produced with metal tools, both, furthermore, purely decorative in character and both present on the same types of artifacts, would be a most unusual situation and one without parallel in Eskimo history. We must await stratigraphic studies at the western sites where the two types are known to be present before definitely settling the question of their chronological positions. But even assuming the two styles to have coexisted on St. Lawrence Island it would still be extremely difficult to account for the presence of only the single type at Punuk and Cape Kialegak, since according to all available evidence there has always been close inter- communication between the different villages on the Island. This exists today and that it existed formerly is indicated by the identity of much of the archeological material from one end of the Island to the other. It must certainly be regarded as significant that at Punuk Island, the only St. Lawrence site where intensive and systematic excavations have been made, there were found among the several thousand specimens, including 141 decorated objects, only four ex- amples of the old curvilinear art and those fragmentary and water- worn; while at Gambell and Kukuliak the random digging of the Iéskimos has brought to light a considerable number of objects deco- rated in both styles. The evidence, therefore, though not yet as direct and conclusive as might be desired, points plainly to the two art styles as representing different periods, with the curvilinear style as the older. : The question as to whether the Punuk style represents a purely local development on St. Lawrence Island must for the present remain unanswered. As to its being a firmly established type on St. Lawrence there can be no doubt; a greater number of specimens of the Punuk type have been found there than of the cuvilinear. It is also significant e R [ _ ‘ a ei NO. 14 PREHISTORIC ART OF ALASKAN ESKIMO—COLLINS ei that only one example of it has been described from a site outside of St. Lawrence—the winged object from Point Hope described by Mathiassen. Thus, the present evidence seems to indicate that the Punuk style of art, if not entirely restricted to St. Lawrence Island, at least played a more dominant role there than elsewhere. We must have further knowledge of other old north Alaskan sites before the actual range of the Punuk style of ornamentation can be determined. OBJECTS FROM PUNUK AND ST. LAWRENCE ISLANDS SHOWING MODERN DESIGNS Reference has been made on page 17 to the finding of seven speci- mens in the old sections of Punuk and Cape Kialegak showing modern decoration. On plate 17 are shown six of these, together with two similarly decorated specimens from Gambell. Plate 17, a, is probably one piece of a double knife handle. The decoration is thoroughly modern, consisting of two narrow bands con- taining alternating spurs, two small Y-shaped figures, and pairs of parallel lines. It should be mentioned perhaps that this specimen was found on the surface at the old village, where it might possibly have been lost by the later people who occupied the houses at the end of the Island within recent years. Plate 17, e, represents a cord handle of modern type from Punuk Island. It is carved in the shape of a seal with a longitudinal hole through the base for receiving the line. Short straight lines are the only decoration. In f is shown a wrist guard collected by Dr. Riley D. Moore at Gambell. It is of deeply stained ivory and bears the decoration of narrow bands and numerous alternating spurs generally applied to St. Lawrence wrist guards. In g is shown another object from Gambell, the use of which is doubtful. It is decorated like the preceding specimen but with larger and more widely spaced spurs. The object shown in / is a ferrule used on the end of the dog whip handle for disentangling the harness lines. The ornamentation of lines and spurs is carelessly applied. The three bone tubes, b, c, and d, plate 17, from Punuk Island and Cape Kialegak, are probably needle cases. The simple decoration consists of encircling lines, spurs, and detached dots. A similar specimen was purchased which had been excavated from the old village at Gambell. Assuming these tubes to have been used as needle cases, it will be observed that they are practically identical with those 3 OO EE eeeoorormrmrmrmreee———V—————— 32 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 8&1 in use by the present Eskimo of the Yukon-Kuskokwim district. Two of these modern needle cases are shown for comparison on plate 18. We seem to have evidence in the St. Lawrence and Punuk speci- mens of a definite association between a certain ornamental type and a particular class of objects. The same simple ornamentation was found on a few additional specimens, some of which are shown on plate 17, but the bone tubes represent the only class of objects in which every example bears this simplified modern decoration. Its presence at the old sites shows that at a comparatively early period the designs that we think of as characteristic of the modern Eskimo - were already established, although submerged or overshadowed by the more elaborate designs typical of Punuk. It is merely a simpli- fication of these designs, however, and not something apart, for spurred lines and dots are among the constant features of the Punuk style. On plate 9, figure 3, is shown the upper part of an old needle case of different form excavated at Gambell. It is of especial inter- est in connection with Boas’ interesting study of needle cases* and Mathiassen’s recent references thereto.” Boas concluded that the flanged tubular needle case of the Norton Sound region in Alaska and the winged needle case of the eastern Eskimo were derived from the same origin: “‘It seems to me very plausible that the Alaskan type and the Eastern type represent specialized developments of the same older type of needlecase, and that the flanges and diminutive knobs of the Alaskan specimens are homologous to the flanges and large wings of the Eastern specimens.” Furthermore, he showed conclusively that the animal heads and human figures found on some of the Alaskan specimens were secondary adaptations of the original flanges and knobs: “ The conclusion which I draw from a comparison of the types of needlecases here represented is that the flanged needlecase represents an old conventional style, which is ever present in the mind of the Eskimo artist who sets about to carve a needlecase. The various parts of the flanged needlecase excite the imagination of the artist; and a geometrical element here or there is developed by him, in accordance with the general tendencies of Eskimo art, into the representation of whole animals or of parts of animals. In this manner small knobs or the flanges are developed into heads or animals.” Mathiassen questioned the validity of this view ‘Boas, Franz. Decorative Designs of Alaskan Needlecases: A Study in the History of Conventional Designs, Based on Materials in the U. S. National Museum. No. 1616, Proc. U. S. Nat. Mus., Vol. XXXIV, pp. 321-344, 1908. * Archeology of the Central Eskimo, Vol. II, pp. 92-97. ee NO. I4 PREHISTORIC ART OF ALASKAN ESKIMO—COLLINS 33 and concluded that ** The Alaskan type seems to be a very locally re- stricted, special form, which has hardly anything to do with the origin of the winged needle case.” On the other hand he considered that the Alaskan type of needle case in human form was the prototype of the winged needle case of the East. As evidence of this he points to two ancient needle cases of a modified winged type from Point Barrow, one of which has on each wing two oblique lines which “ are appar- ently intended to indicate the hands bent in front of the body.” The interpretation of these lines as hands is extremely doubtful. No single feature of the two needle cases suggests the human form and as for the pairs of oblique lines, they may be compared much more readily with the similar oblique lines on the lower parts of the flanges of many of the Alaskan specimens. The significant point, however, is that these two Point Barrow needle cases are in reality intermediate between the winged needle case of the East and the Alaskan flanged needle case, and not, as Mathiassen has supposed, between the winged case and the Alaskan case in human form. Comparison of the three types will, I believe, bear this out. The upper portions of the Point Barrow specimens are widened out like the Alaskan forms although there are no distinct flanges. The bands and spurred lines are known to both regions but their arrangement is more suggestive of the Alaskan cases. The “ wings” are of the Eastern type but are much longer and narrower than in the typical Eastern needle case. In the fragmentary needle case from Gambell, plate 9, figure 3, we have a modification of the Point Barrow type still further in the di- rection of the Alaskan flanged type. The general shape is that of the Alaskan case except that there are no flanges at the enlarged end ; the bands with alternate spurs are common to both types; the two long incurving lines down the sides are the same as those on the Point Bar- row specimens but they do not contain “ wings.” I consider that the Point Barrow and Gambell needle cases furnish further and con- clusive evidence of the relation between the Alaskan and Eastern needle cases as demonstrated by Boas. It appears to me also that we may have here a possible explanation of the origin of the Eskimo needle case. The Gambell form may have been the prototype from which developed on the one hand the restricted Alaskan form with its flanges and knobs and on the other the Eastern form with its promi- nent projecting wings, the Point Barrow specimens representing an early stage in the development of the winged type. I do not care to stress this hypothesis, however. The Gambell type might as reason- ably have been derived from the Point Barrow type, having retained the long incurving lines while losing the wings. Whatever the expla- 34 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 8&1 nation may be, it now seems very probable that the origin of the needle case when fully traced, will be found to be in Alaska; and that whatever form it may have had originally it was not a human or animal form, the occurrence of which among the Eskimo, as Boas has shown, can so often be attributed to the strong tendency of the artist to enliven and vary his handiwork by the occasional replacement of existing simple or geometric elements by life forms. RELATION BETWEEN THE ANCIENT AND MODERN ART OF THE BERING SEA REGION The objects illustrated on plate 18 are from Nelson’s collection of modern Alaskan Eskimo material and are included to show certain designs of the Punuk period that have continued in use to the present time. Plate 18, f and g, are the two needle cases previously referred to from the Kuskokwim and the lower Yukon, respectively. They are shown for comparison with the old needle cases from St. Lawrence and Punuk Islands illustrated on plate 17. An animal carving from Bristol Bay is shown in a. Opposite rows of six oblique lines suggest the ribs while the seven small squares with enclosed dots are no doubt supposed to represent the vertebral column. Squares of this kind were also seen on plate 12, f, from Punuk, but they were not employed in a manner to suggest a realistic meaning. In b is shown the under side of a woman’s hair ornament from Agiukchugumut, to the south of Nelson Island. Like the preceding specimen it has a row of small squares but no dots. This is a fairly common design among the Alaskan Eskimo; it is to be compared with similar designs from Punuk shown on plates 10, a-b, and 15, i. Plate 18, c, is a bodkin from Sledge Island. Two of the edges are carved with a series of notches or serrations in the tops of seven small elevations. This is the principal decorative motive on the lower end of bird dart points from Punuk, plate 15, e, f, g. Other mod- ern examples are given by Hoffman,’ plates 37, 5; 38, 4; 30, 3-4; and by Nelson, plate x11, 23; and figure 20. In d is shown a woman’s workbag fastener and bodkin from the lower Yukon. It is introduced here to show the continuation among the modern Eskimo of the familiar decoration of lines with spurs attached. It should be compared with many objects of both the Punuk and the curvilinear stages shown on previous plates. * Hoffman, W. J., The Graphic Art of the Eskimo. Rep. U. S. Nat. Mus., 1897. * Nelson, E. W., The Eskimo about Bering Strait. 18th Ann. Rep. Bur. Amer. Ethnol., 1890. NO. I4 PREHISTORIC ART OF ALASKAN ESKIMO—COLLINS 35 Another workbag fastener, from Norton Sound, is shown in e, on which the decoration is restricted to bands of parallel lines. Similar designs were observed on specimens from the Punuk period shown on plate 10, c-d, and figure 2. On plate Ig are illustrated additional objects from the modern Alaskan Eskimo showing typical designs that occur also in the Punuk stage. It is of considerable interest to note that in the modern material from St. Lawrence Island there are very few decorated objects of any kind, so that in order to find modern examples with which to compare the old art of St. Lawrence we must turn to the Alaskan mainland. This poverty of decoration on St. Lawrence is paralleled on the Asiatic side of Bering Strait, where the Yuit, the Siberian kinsmen of the St. Lawrence Islanders, also exhibit a striking deficiency in art. The reduction of the modern St. Lawrence Island and Siberian Eskinio to such a low artistic level can perhaps be best explained as the result of a relatively late Chukchee influence. The Eskimo of the Alaskan mainland, practically free from such influence, have merely retained a more abundant residuum of the highly developed ancestral art common to the entire region. Plate 19, a, b, and c, are three objects from the modern St. Law- rence Eskimo, collected by Dr. Moore. The first, a, is a broken wrist guard with decoration similar to the one shown on plate 17, f, but with the addition of pronged figures to the lines and spurs. The two small bird figures, b and c, are simply ornamented with dots. In d and ¢ are illustrated a workbag fastener and belt buckle from the Kuskokwim region collected by Nelson, showing the well known nucleated concentric circles which are directly comparable to the compass-made circles of the Punuk period. It will also be observed that the centers of the circles, as is so often the case, have wooden insets. This is a feature that was also observed in the old curvilinear art, but which was not present on any of the objects from Punuk or Cape Kialegak. Plate 19, f and g, are two modern specimens from Norton and Kotzebue Sounds, on which the spurs within the bands are so evenly applied as to make the uncut space between them appear as a continu- ous zig-zag. On plate 20 are illustrated six modern harpoon heads showing the nature of ornamentation applied to these objects by the modern Alas- kan Eskimo. By far the greater number of modern harpoon heads are undecorated, in contrast to the old specimens which often bear elabo- rate designs. The large whaling harpoon head of bone, a, is from Point Hope. It bears a simple ornamentation of lines and spurs arranged about ne ae eos 36 SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL. 81 the line hole in the manner characteristic of the Point Hope region, and likewise of Punuk, though the decoration on the harpoon heads from the latter locality is much more elaborate. The two smaller harpoon heads, b and c, from Point Barrow, are simply decorated with lines, and on b, small hachured triangles. Red pigment has been rubbed into the incisions. In d is shown a small harpoon head from Nunivak Island. It has a three pronged barb and a decoration of two concentric circles with spurs attached. Nunivak Island and the neighboring mainland are the only localities where circles and dots are applied to harpoon heads at the present time. The small bone harpoon head shown in e is from the Semidi Islands, south of the Alaska Peninsula. Across one barb and extend- ing obliquely up toward the center is a decoration consisting of a nar- row band enclosing six small lines. In f is shown a bone harpoon head excavated at Metlatavik, 22 miles above Cape Prince of Wales. On both the upper and lower sides is an elongated depression bordered by two lines, the outer one con- tinuing to the base where it follows the bifurcated barb. Immediately above the line hole are two small grooves 5 mm. long, one on each side, which from the position and shape may be regarded as representing ornamental remnants of grooves for side blades. Two modern seal dart foreshafts from Southwest Alaska, carved to represent the sea otter, are illustrated on plate 21. These are intro- duced for the purpose of comparison with the ancient ivory object on plate 5 and the somewhat more recent foreshaft shown on plate 9, figure 2. These two modern foreshafts illustrate the well-known tendency of the Eskimo of the Bristol Bay-Nunivak region to utilize life forms for the embellishment of their implements and weapons. An animal with open mouth and exposed teeth is a favorite decoration applied to the foreshaft of the seal dart. In a the eyes and nostrils are small cylindrical plugs of baleen, but in b they are merely shallow depressions filled with a bluish clay. Plate 9, figure 2, represents the foreshaft for a light dart. This was bought at Teller, Seward Peninsula, and was reported to have been excavated on one of the Diomedes. It is 84 cm. long, but the lower end is missing. There is a circular perforation on the lower side for holding a thong, and in the forward end a circular hole 8 mm. deep for the dart head. The projecting ears are suggestive of a land mammal but the curves about the head suggest the gills of a fish. Apparently there was no intention to clearly represent any particular animal. The discs forming the eyes are of baleen. The object is a rich chocolate ae NO. I4 PREHISTORIC ART OF ALASKAN ESKIMO—COLLINS 37 brown but the ornamentation shows no direct relation with the old Ber- ing Sea art with the exception of the spurs bordering the curving lines. The decoration is also of neither the Punuk nor the modern type. It may be that it represents an intermediate stage of art on the Diomedes between the old curvilinear and the modern. On the other hand it may be largely the result of individual fancy and have little or no signifi- cance as an art type. Its shape, however, is of more importance, and along with the older object shown on plate 5, might be taken to indi- cate a closer connection in early times between the art of Bering Strait and Southwest Alaska, or even, as was suggested before, to point to a possible ancient connection between the Bering Strait region and the Northwest Pacific Coast. I would again stress, however, that these few specimens are wholly inadequate from either the standpoint of numbers or of closeness of form and design, to afford more than a suggestion that future archeological investigations may reveal more dependable evidences of such a possible contact. If no such evidence should be forthcoming the realistic and symbolic art of Southwest Alaska could no doubt be safely regarded as the result of compara- tively late Indian influence that furnished life motives around and within which these Eskimos continued to employ the geometric ele- ments they possessed in common with the Alaskan Eskimo to the northward. Comparison of the decorative art of the Punuk period with that of the modern Alaskan Eskimo reveals numerous striking similarities as well as certain important differences. On plates 18 and 19 are shown examples in which individual designs have been carried over without change. However, these designs are differently applied on the modern objects. The decorative elements are usually detached, or, if con- nected, are repetitive. The decorated objects of the Punuk period, on the other hand, are generally marked by a certain continuity of design. This may be observed on practically all of the typical Punuk examples, whether as on plate 12, where the lines and dots are some- what sparingly applied, or on plate 13, where the lines and spurs cover all of the available surface. The three principal elements in Eskimo art: the spurred line, the Y figure, and circle and dot are seen to have been present, though usually in different form, in either or both the curvilinear stage of the old Bering Sea culture and the succeeding Punuk stage. The spurred line is a common feature to both stages; the Y figure does not appear in the earlier curvilinear art but in the more angular art of the Punuk period it is a common design although it differs from the modern pronged or Y figure in being larger and in being connected with a unit 38 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 81 design; whereas in the modern art it usually rises from a base line and stands detached. The nucleated circle occupies a most important place in the decorative scheme of early Eskimo art. It was shown to occur in the old Bering Sea culture as a slightly irregular, often elliptical figure, engraved free hand and apparently with stone tools. It then follows in the Punuk stage as a perfectly symmetrical, cleanly cut circle, made with a compass or bit which almost certainly was of metal. In the modern art it is made in exactly the same way, although it is usually represented as a more or less detached element instead of an integral part of a connected design as in the old curvilinear and Punuk stages. The distribution of the circle and dot design in Northwestern America has recently been studied by Dr. Leslie Spier and Miss A. Dorothy Smith.” The following statement is made: “ This has often been looked oh as a typical Eskimo decoration. But we are able to show by its distribution that it is more clearly characteristic of the Indians of the northwest, with only a limited distribution among the Eskimo.” In conclusion, the following statement is made: “ In western Alaska the great elaboration of the dot and circle into a series of concentric circles numbering frequently five and six may be de- pendent upon iron tools. The extreme regularity of the circles speaks for the likelihood of the use of bits of various sizes. This, however, does not solve the problem of the simple nucleated circle which is probably older and, together with the alternate spur design, the basic unit from which the elaborate decorations are made. Two rea- sons can be given for this view. First, it is simple and possible to accomplish with stone implements. Second, in its simple form as a single dot and circle it has a wide and fairly continuous spread down the Pacific coast, and a wide if sporadic distribution in Eskimo ter- ritory. If the Alaskan decoration had been imitated, we would expect to find some similar examples elsewhere.” The principal value of such studies of spatial distribution lies in the light they may be able to throw on the problem of the origin and spread of culture traits when more dependable data revealing a direct time sequence are lacking. In the present case there was some justifi- cation for regarding the circle and dot in Alaska as derivitive, in view of its greater spread among the Indian tribes to the southward and in the absence of conclusive archeological evidence to the contrary. Con- sidered in the light of recent archeological developments in Alaska, however, the validity of this conclusion can no longer be upheld. *The Dot and Circle Design in Northwestern America. Journ. Soc. Ameri- canistes de Paris, XIX, pp. 47-53. NO. I4 PREHISTORIC ART OF ALASKAN ESKIMO—COLLINS 39 There is now clear evidence of the antiquity of the circle and dot de- sign in northwestern Alaska where it is seen to have formed the basic element in a very old art style. There is, on the other hand, no evidence of the antiquity of the design over the wide area outside of Alaska where it is now found except possibly at certain sites excavated by Harlan I. Smith in British Columbia and Washington.’ The antiquity of these finds was regarded as questionable, however, Smith being of the opinion that the design in this region was relatively recent. Where- ever the circle and dot may have originated, among the Alaskan Eskimo at least it was indigenous, having developed from an earlier Alaskan culture. That this earliest known form of the Eskimo circle and dot may have had its origin to the southward is, of course, pos- sible, but there is at present no evidence pointing in that direction. The various elements that enter into the composition of the designs of the old curvilinear Bering Sea art, the later Punuk stage, and the modern have been examined. In order that these three stages may be directly compared the observed resemblances and differences are given below in tabular form. A + sign indicates the presence and a — sign the absence of a feature. Curvilinear Punuk Stage Stage Modern incom lac mel min Cle Sie yevwomr racine ons olsuoreaovapenavase neve =vs an — a Pep ASSUINAGENEINCLCS tk jo. c sci oe eed nese es oe os te ae MNIEC ATI SILES oe. fects fd be pce ee secs owt als + _ — ieeised ‘circles and ‘ellipses. 3. .....5..6.0--%5 _ ~ — Circles suggesting a pair of eyes.............. + ate — Circles between converging lines.............. + — — Small plugs at centers of circles.............- + ? a SHrved liplike= projections... ........¢.-ce00-s + — — Grriay troup ites wate ee sie css c s 6 ale: «o.c's) abies slese me | ae Rare BGEMICLOSS ee IMCS te sales. ccc ose s fs see oes satus _ ae oe EGORem VINES feu ass oe lsosia sie Re a atade mete + = _ IEE Phyas CUPP MIAMES Meese et 9 ciesaig « s.a'< een 4)d inl ol = 4. se NEVE ly Ctber lime Seamer sc cistseis-o. a;s-s Selo ereieretidons + Rare — Hines in Strateht bands. ....0....cse0et seces -- + at SESE WITLI (CICCIES Sisc2. a. .c «s/c sia vs «2 cw elak.cle ees + + ID GESHAGMENGSHOL VINES... eia.c-x.'s sale cies eee deorte _ ss — MBs SCH ACH EMME ReS © cfs oS 5.6 40% bare s'ais $5.2 slsie eae _ dle a Srediehity avid “ODMGUE SPULS:,. 0. .'s.. S gQ PLATE I0 a-h, Ivory object, Punuk Island. 9 cm. high. Cat .No. 343141, U. S. Nat. Mus. c-d, Ivory object, Alaska. 6.5 cm. long by 8 cm. wide. Cat. No. 344677, U. S. Nat. Mus. e, Ivory object, Kukuliak, St. Lawrence Island. 5.1 cm. long. Owned by Mr. C. L. Andrews. PLATE II a, Harpoon head, Punuk Island. 7.1 cm. long. Cat. No. 344034, U. S. Nat. Mus. b, Harpoon head, Punuk Island. 7 cm. long. Cat. No. 343945, U. S. Nat. Mus. c, End of box handle, Punuk Island. 8.8 cm. long. Cat. No. 343199, U. S. Nat. Mus. d, Harpoon head, Punuk Island. 9.1 cm. long. Cat. No. 344021, U. S. Nat. Mus. e, Harpoon head, Cape Kialegak, St. Lawrence Island. 10 cm. long. Cat. No. 342062, U. S. Nat. Mus. f, Upper end of dart foreshaft, Kukuliak, St. Lawrence Island. 7.7 cm. long. Cat. No. 344601, U. S. Nat. Mus. g, Ivory object, Punuk Island. 10.8 cm. long. Cat. No. 343370, U. S. Nat. Mus. h, Wrist guard, Sevuokok, St. Lawrence Island. 10 cm. long. Cat. No. 342744, U. S. Nat. Mus. » Drill rest, Punuk Island. 13.7 cm. long. Cat. No. 343427, U. S. Nat. Mus. >. PLATE I2 a, Harpoon head, Punuk Island. 8.5 cm. long. Cat. No. 343215, U. S. Nat. Mus. b, Harpoon head, Punuk Island. 9.1 cm. long. Cat. No. 343162, U. S. Nat. Mus. c, Harpoon head, Cape Kialegak, St. Lawrence Island. 9.7 cm. long. Cat. No. 342061, U. S. Nat. Mus. d, Ivory object, Punuk Island. 8.1 cm. long. Cat. No. 343716, U. S. Nat. Mus. e, Base of harpoon head, Cape Kialegak, St. Lawrence Island. 8.8 cm. long. Cat. No. 342877, U. S. Nat. Mus. 50 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 81 f, Ivory object, Punuk Island. 17.2 cm. long. Cat. No. 343220, U. S. Nat. Mus. g, Part of knife handle, Punuk Island. 15.8 cm. long. Cat. No. 343230, U. S. Nat. Mus. h, Harpoon foreshaft, Punuk Island. 14.5 cm. long. Cat. No. 343231, U. S. Nat. Mus. PLATE 13 a, Harpoon head, Punuk Island. 9.1 cm. long. Cat. No. 344110, U. S. Nat. Mus. b, Harpoon head, Cape Kialegak, St. Lawrence Island. 7.1 cm. long. Cat. No. 342875, U. S. Nat. Mus. c, Ivory object, Punuk Island. 14 cm. long. Cat. No. 343372, U. S. Nat. Mus. d, Ivory object, Punuk Island. 16.2 cm. long. Cat. No. 343228, U. S. Nat. Mus. e, Ivory object, Punuk Island. 15.6 cm. long. Cat. No. 343371, U. S. Nat. Mus. f, Ivory object, Cape Kialegak, St. Lawrence Island. 4.4 cm. long. Cat. No. 342876, U. S. Nat. Mus. g, Ivory object, Punuk Island. 12 cm. long. Cat. No. 343613, U. S. Nat. Mus. PLATE 14 Ivory object, Kukuliak, St. Lawrence Island. 14.2 cm. long. Owned by Mr. C. L. Andrews. PLATE 15 a, Harpoon head, Cape Kialegak, St. Lawrence Island. 11.1 cm. long. Cat. No. 3429093, U. S. Nat. Mus. b, Harpoon head, Punuk Island. 8 cm. long. Cat. No. 344062, U. S. Nat. Mus. Bone harpoon head, Punuk Island. 8 cm. long. Cat. No. 343160, U. S. Nat. Mus. d, Harpoon head, Punuk Island. 7.4 cm. long. Cat. No. 343213, U. S. Nat. Mus. Bird dart point, Punuk Island. 9 cm. long. Cat. No. 343173, U. S. Nat. Mus. , Bird dart point, Punuk Island. 8.6 cm. long. Cat. No. 343172, U. S. Nat. Mus. g, Bird dart point, Cape Kialegak, St. Lawrence Island. 12.5 cm. long. Cat. No. 342001, U. S. Nat. Mus. h, Piece of box handle, Punuk Island. 10.2 cm. long. Cat. No. 343681, U. S. Nat. Mus. ; 1, Broken ivory object, Punuk Island. 5.5 cm. long. Cat. No. 343076, U. S. Nat. Mus. j, Piece of wrist guard, Sevuokok, St. Lawrence Island. 7.3 cm. long. Cat. No. 344530, U. S. Nat. Mus. Q 3 n 3 Hh PLATE 16 Ivory figurine, Punuk Island, 11.7 cm. long. Cat. No. 344107, U. S. Nat. Mus. PLATE 17 a, Part of knife handle, Punuk Island. 10.2 cm. long. Cat. No. 344683, U. S. Nat. Mus. b, Bone needle case, Punuk Island. 5.1 cm. long. Cat. No. 343472, U. S. Nat. Mus. NO. I4 PREHISTORIC ART OF ALASKAN ESKIMO—COLLINS 51 c, Bone needle case, Punuk Island. 7.2 cm. long. Cat. No. 343955, U. S. Nat. Mus. ; d, Bone needle case, Cape Kialegak, St. Lawrence Island. 5.5 cm. long. Cat. No. 343017, U. S. Nat. Mus. e, Cord handle, Punuk Island. 5 cm. long. Cat. No. 343956, U. S. Nat. Mus. f, Wrist guard, Sevuokok, St. Lawrence Island. 8.7 cm. long. Cat. No. 280385, U. S. Nat. Mus. g, Ivory object, Sevuokok, St. Lawrence Island. 9.7 cm. long. Cat. No. 344525, U. S. Nat. Mus. h, Ferrule for dog whip, Punuk Island. 4.6 cm. long. Cat. No. 343430, U. S. Nat. Mus. PLATE 18 a, Ivory object, Bristol Bay. 9.8 cm. long. Cat. No. 168626, U. S. Nat. Mus. b, Hair ornament, Agiukchugumut, south of Nelson Island. 3.7 cm. long. Cat. No. 37008, U. S. Nat. Mus. c, Bodkin, Sledge Island. 12.4 cm. long. Cat. No. 45339, U. S. Nat. Mus. d, Workbag fastener, Lower Yukon. 12.6 cm. long. Cat. No. 48870, U. S. Nat. Mus. e, Workbag fastener, Norton Sound. 12.5 cm. long. Cat. No. 33285, U. S. Nat. Mus. f, Bone needle case, Lower Kuskokwim. 8.6 cm. long. Cat. No. 36787, U. S. Nat. Mus. g, Bone needle case, Lower Yukon. 12.2 cm. long. Cat. No. 48604, U. S. Nat. Mus. PLATE 19 a, Wrist guard, Gambell, St. Lawrence Island. 8.2 cm. long. b, Bird figure, Gambell, St. Lawrence Island, 4.6 cm. long. c, Bird figure, Gambell, St. Lawrence Island, 2.9 cm. long. d, Workbag fastener, Lower Kuskokwim. 16.3 cm. long. Cat. No. 176225, U. S. Nat. Mus. e, Belt buckle, Lower Kuskokwim. 6.4 cm. long. Cat. No. 37332, U. S. Nat. Mus. f, Box handle, Kotzebue Sound. 8.6 cm. long. Cat. No. 48562, U. S. Nat. Mus. g, Bodkin, Norton Sound. 12.4 cm. long. Cat. No. 43837, U. S. Nat. Mus. PLATE 20 a, Harpoon head, Point Hope. 22.4 cm. long. Cat. No. 201058, U. S. Nat. Mus. b, Harpoon head, Point Barrow. 10.8 cm. long. Cat. No. 56616, U. S. Nat. Mus. c, Harpoon head, Point Barrow. 8.2 cm. long. Cat. No. 56611, U. S. Nat. Mus. d, Harpoon head, Nunivak Island. 7.6 cm. long. Cat. No. 339598, U. S. Nat. Mus. e, Harpoon head, Semidi Islands. 6.5 cm. long. Cat. No. 72547, U. S. Nat. Mus. f, Harpoon head, Metlatavik, Seward Peninsula. 6 cm. long. Cat. No. 342617, U. S. Nat. Mus. 52 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 81 PLATE 21 a, Dart foreshaft, Alaska Peninsula. 21.5 cm. long. Cat. No. 127766, U. S. Nat. Mus. b, Dart foreshaft, Lower Kuskokwim. 21.4 cm. long. Cat. No. 38442, U. S. Nat. Mus. PLATE 22 Fic. 1. View oi the Punuk Island midden. Excavation along the outer edges showed that it extended as deep as six feet below the present beach line. Total height 16 feet. Fic. 2. Excavation in House No. 3, Punuk Island, recent, showing skeleton and partially exposed wooden floor. PLATE 23 Section of the Punuk Island midden at Cut B, showing timbers and whalebones of an old house at bottom, now six feet below reach of storm tides. PLATE 24 Recent house ruin at Cape Kialegak, St. Lawrence Island. Framework of drift- wood logs and whale ribs and jaws. Abandoned 50 to 60 years ago. (‘gb pure -£ sosrd 99s uolvurydxe 10,]) ‘AIOAL SHLAPVM JO spol uoodae fy SNOILO3711090 SNOS NV113 0SIW NVINOSHLIWS SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 81, NO. 14, PL. 2 i | | ; | Harpoon heads of walrus ivory. (For explanation see pages 5-6 and 48. ee SSS —— (‘gr pue 4-9 so3ed 90s uorjeur[dxe 10.7) “CIOAL pue auoq JO sa_puey XO € “Id ‘tl “ON ‘18 “10A SNOILO31100 SNOSNV11S0SIN NVINOSHLIWS (gr pue Z sased 99s uoneueldxa 104) ‘ash UMOUyUN JO sydo[qGo AIOAT b “Id ‘pl “ON ‘LB “I0A SNOILO3171100 SNOANVIISOSIW NVINOSHLIWNS (gr pue g-Z saded 3aas uoljeurldxa 10,1) ‘peo S[eunrue ue JUdSaIda1 0} paased joalqo Gd ‘bl "ON ‘18 “0A AOAT SNOILO31109 SNOANV1ISOSIW NVINOSHLIWS Cy pUe UY 2P90CU BEd UUIFOUTUAD Aus) ‘ash UMOUNUN FO JOIfGO AITOAT 9 “Id ‘PL “ON ‘18 “10A SNOILO31100 SNOANVIISZOSIN NVINOSHLIWS (gr pure OI-6 sased 3as uoljeur[dxa JOT) ‘2091d-JayIOS U0K daey AIOAT Jo yred pue ‘ 9sh UMOUYUN JO yalqo AIOAT uayorg : ; (eS ) £"ld ‘tl “ON ‘18 “10A SNOILO31109 SNOANV17390SIN NVINOSH.LIWS (‘6h pue ri-o1 saded 9as uoljeurldxa 107) ‘Q001d-JayIos uoodiey AIOAT Jo yaed pue [eas b Juasaidat 0} padres yoofqo AIOAT 8 "Id ‘bl “ON ‘18 “10A SNOILO31100 SNOANV11IS0SIW NVINOSHLIWS eee (6b pue gf ‘z& ‘41 saded vas uoleurldxe 1047) ‘AIVJUIWISCIF [Je ‘AsBdo]P9ou pue }peYysetO} JAep ‘speoy uoodiey AIOAT I c aa Tt EN VIE RON NU ae OS at ee aS a a wv ~$ SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 81, NO. 14, PL. 10 i —— ——— Ivory objects of unknown use. (For explanation see pages 18-19 and 49.) (6p pur oz-61 saded vas uoneurldxe 10,7) ‘spuv]s] aouaIMey JS puw yNuNg wWosz syoalqo A1OAT LL ‘Id ‘bb “ON ‘18 “10A SNOILO31100 SNOANVTISOSIN NVINOSHLIWS oL “Td ‘pL “ON 'L8 “110A (‘6h pue zz-1z sased 99s uoneurldxe 10,7) ‘SpuR|ST 9OUZIMET “JG puke yYNUNG wWosF spofqo auoq puke K10AT SNOILOS1100 SNOANVITSOSIN NVINOSHLIWS (‘oS pue 2-2 sased 99s uorjeurldxa 10...) “spuels] JOUIIMET YS pue hun wot $}0fqo AIOAT EL “Id ‘tl “ON ‘18 “10A (‘oS pue tz sased 9as uoljeurldxa IO.T ) “puels] JIUIIME T YS “eIpN yy wood} osn uMOUAUN JO yoafqo AJOAT vl .Id ‘yt “ON 'L8 “IOA SNOILO31100 SNOANVITSOSIW NVINOSHLINS (‘oS pue Sz-bz sa8ed aas uorjeurldxa 10,7) ‘spur]sT s0UeIMeT “YS pue ynung wolf syalqo AIOAT SHLINS . SNOILO31109 SNO3 NV1130SIW NVINO ciocqa ‘+t “ON ‘LB IOA SMITHSONIAN MISCELLANEOUS COLLECTIONS = » a ae Gs eh, te ha “ee PaaS gt gl OE Ps a ee a Ivory figurine from Punuk Island. (For explanation see pages 25-26 and 50.) VOL. 81, NO. 14, PL. 16 ay LL ‘1d ‘pL “ON ‘E8 “10A (‘oS pue ze-1e sosevd sas uoneuridxa 10,1.) *spue]s] DUITIMET 4S pue hun wot} $}99 [qo ouo0q pue AIOAT SNOILO31100 SNOANVT1S0SIN NVINOSHLIWS BL “Id ‘bh “ON ‘L8 “TOA CIS pue Sf-r€ sased aas uoneurldxa 10,7) ‘9U0g puke AJOAT JO sjoafqo OLULYSy] URYsE]Yy UsOpoy P 2 S57 U Yeeror SNOILO31109 SNOANVIISOSIN NVINOSHLIWS (‘18 pue S€ sased vas uoneurldxa 10.7) “KIOAT JO Spoafqo oumysy UPYseTy UtOpPoW a 6L “Id ‘bl “ON ‘18 “1OA SMITHSONIAN MISCELLANEOUS COLLECTIONS VOES 81), INO 14, (Pie320 i ; ; Modern Alaskan Eskimo harpoon heads of bone and ivory. (For explanation see pages 35-36 and 51.) ue gf saded 998 uoneurldxa IO) 0} poAred LIOAL JO S}FRYSI1O} JAP [BOS UIIpO fy "ZS ( P “Ja}}0 BIS dy} JUese1doI Rhine re L LZ “Id ‘pL “ON ‘18 “10A SNOILO311090 SNOANV11S0SIN NVINOSHLIWS SMITHSONIAN MISCELLANEOUS COLLECTIONS Excavations on Punuk Island. (For explanation see pages 14-16 and 52.) VOL. 81, NO. 14. PL. 22 a SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 81, NO. 14, PL. 23 Section of Punuk Island midden. (For explanation see page 52.) ¥S “Id ‘PL “ON ‘L8 “10A CZs a8ed 39s uoneurldxa 10.7) PULST SUSIMET “IS ‘Yesoyery ode je ums asnoy yusooy A * SNOILD31100 SNOANV11S0SIN NVINOSHLIWS ! IT sO NIAN MISCELLANEOUS COLLECTIONS ua VOLUME 81, NUMBER 15 Ng e Det (End of Volume) : RTHROPODS AS INTERMEDIATE HOSTS OF HELMINTHS BY - MAURIGE G. HALL , Zoological Division, Bureau of Animal Industry, U. S. Department of Agriculture : x oe ¢ VIN es a : aft cs ui ) : | \ % = z ; / (PUBLICATION 3024) k GITY OF WASHINGTON | a -pUBLISHED BY THE SMITHSONIAN INSTITUTION ee SEPTEMBER 25, 1929 Eee SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLUME 81, NUMBER 15 (End of Volume) ARTHROPODS AS INTERMEDIATE HOSTS OF HELMINTHS BY MAURICE CGC. HALL Chief, Zoological Division, Bureau of Animal Industry, U. S. Department of Agriculture (PUBLICATION 3024) CITY OF WASHINGTON PUBLISHED BY THE SMITHSONIAN INSTITUTION SEPTEMBER 25, 1929 | A a a %® 2 “ » a se ee ° = & 2 2 2 ° al s a » BALTIMORE, MD., U. S. A. ARTHROPODS AS INTERMEDIATE HOSTS OF HELMINTHS By MAURICE C. HALL, . CHIEF, ZOOLOGICAL DIVISION, BUREAU OF ANIMAL INDUSTRY, U. S. DEPARTMENT OF AGRICULTURE INTRODUCTION The phylum Arthropoda contains numerous forms which serve as intermediate hosts of many parasitic worms, including nematodes, acanthocephalids, flukes, and tapeworms. This fact follows naturally from the fact that the arthropods are an exceedingly large group of animals, including the ubiquitous insects and the numerous and widely distributed crustaceans. It also follows from the fact that these arthro- pods constitute the food supply, wholly or in part, for so many higher animals, especially for such forms as fish, many amphibians, some reptiles, numerous birds, and some mammals. To a lesser extent it follows from the fact that in feeding on various plants the higher animals are certain to swallow the arthropods habitually present on or in these plants. It follows from the fact that many insects feed on or breed in manure and consequently are exposed to infection from the eggs or larvae of worms parasitic in the hosts responsible for the manure. Last, but not least, the importance of arthropods as inter- mediate hosts of parasitic worms follows from the fact that large numbers of anthropods, especially the innumerable biting insects, whether transient or permanent ectoparasites, feed on blood and so serve as intermediate hosts of worms which have larval stages living in the blood of vertebrates. The worm parasites may be classified from one point of view as monoxenous or heteroxenous. The monoxenous worms have life histories in which the worms pass from one host animal to a similar host animal without the intervention of an intermediate host. The heteroxenous worms have life histories in which in most cases the worms pass from mature stages in one host animal to larval stages in a host animal of a different sort, the intermediate host, and then return to a host animal of the first sort or a more or less closely related species and develop in this animal to maturity. In some instances two intermediate hosts are utilized in sequence for larval stages. SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 81, NO. 15 (END OF VOLUME) I ho SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL. 81 Of the four worm groups named, the cestodes are almost exclu- sively heteroxenous. We have the rare exception of Hymenolepis nana of the rat, which develops as an adult in the small intestine of the rat, produces eggs which pass out in the feces and by contamina- tion of the rat’s food infects the rat with the larval stage of the tapeworm, a small cysticercoid which develops in an intestinal villus of the rat, and which then returns to the lumen of the intestine to become an adult worm, the rat serving as both the primary and the intermediate host for the worm. Even in this case it has been claimed that rat fleas may act as intermediate hosts, but this has not yet been confirmed. This may be one of those cases in which a parasite can use an intermediate host or do without it. We seem to have similar cases in such parasites as the common gape-worm of poultry which can utilize the earthworm as an intermediate host or can infect chickens directly, and the blackhead organism which can use the cecum worm as an intermediate host or can infect turkeys directly. In the great majority of cases, the tapeworm is adult in an animal which eats the intermediate host animal and thereby becomes infested with the adult worm as the larval worm from the intermediate host comes to maturity in the primary host. In some of the bothriocepha- lids, in cases in which the life histories are well known, the eggs of the adult tapeworms present in the primary host, a higher vertebrate, hatch on entering water, infect such small animals as the copepods, and develop in the body cavity of these first intermediate hosts to an early larval stage, the procercoid. When such infested entomostracans are eaten by such intermediate hosts as fish, the procercoid undergoes further development and becomes a plerocercoid in the flesh of the fish. When infested fish are eaten by a suitable higher vertebrate, such as a human being or dog, the plerocercoid develops to the adult tape- worm in the small intestine of this host. Among the flukes we have one large group, the Monogenea, which are usually ectoparasitic, mostly on fish, but sometimes endoparasitic, as in the respiratory tract of turtles or the urinary bladder of amphib- lans, and these flukes are monoxenous, developing without an inter- mediate host; another large group, the Digenea, are regularly endoparasitic and are heteroxenous. The digenetic flukes occurring in vertebrates produce eggs which pass out in the feces or urine and hatch after entering water. Usually the newly hatched worm (miracid- ium) attacks a mollusk host and develops in this host to the stage known as a cercaria. It may now be eaten by its primary host, or may escape and encyst in water or on vegetation and be swallowed by its primary host, developing in either case to an adult worm, or it may a = = —— SS | NO. 15 ARTHROPOD HOSTS OF HELMINTHS—HALL 3 enter a second intermediate host, an aquatic arthropod or a small fish, and encyst in this host. When such a second intermediate host is eaten by a primary host, the fluke develops to maturity in the new host. Among the nematodes we have several groups which are usually monoxenous, although some of the ascarids, belonging to a super- family, the Ascaroidea, which is ordinarily monoxenous, may be heteroxenous, as in the case of a seal ascarid having a larval stage encysted in fish. One large and important group, the Filariata, composed of two superfamilies, the Filarioidea and the Spiruroidea, is a heteroxenous group with larvae developing in blood-sucking arthropods or in arthropods which feed in some stage of development on the feces of the primary host or on food contaminated with these feces, Among the acanthocephalids we-know of the occurrence of inter- mediate hosts, but for the most part we must assume that this is the rule, as very few life histories are known in this group. In the known cases the worm eggs passing from the primary host infect secondary hosts, develop to a larva and infect primary hosts when these eat infected secondary hosts, or else re-encyst in another intermediate host and infect the primary host when it eats the second intermediate host. The lists of heteroxenous worms and their arthropod hosts, given in this paper, are the most complete of those published and the omis- sions are probably few. The lists for certain groups have been com- piled from time to time, some of the more important and more recent being those of Joyeux (1920), Ransom (1921), Van Zwaluwenburg (1928), Seurat (1916, 1919), MacGregor (1917), and Henninger (1928), and, of course, the indispensable catalogues of Stiles and Hassall, but no previous paper has attempted to cover all the arthro- pod hosts of the parasitic worms of vertebrates. On the basis of the lists given here this paper includes a consideration of the general facts and of the broad principles which may be derived from a correlation of these facts. While it will serve as a reference for the trained scientist in the groups involved, its principal value will be as a ref- erence and guide to the younger worker and student and to the man who works in places remote from adequate library facilities and the specialized literature on arthropods or parasitic worms. The subject of the paper excludes from consideration the worms which have arthropods as primary hosts, and the arthropods which are inter- mediate hosts for Protozoa or animal parasites other than the worm SSS 4 SMITHSONIAN MISCELLANEOUS COLLECTIONS voL. 81 groups occurring as parasites in vertebrates. The intermediate arthro- pod hosts are listed here as completely as possible; the primary host list is frequently abbreviated to only representatives of groups. In the lists arranged on a basis of parasite groups the names of hosts are given as they are found in the literature, regardless of spell- ing, Synonymy, recognizable status, or validity. This is to enable the reader to trace the records if desired. In the final lists, arranged on a basis of intermediate host groups, the parasites are listed under the valid names of their arthropod hosts as far as possible. Synonyms of host names are indicated as synonyms, but names which cannot be recognized as valid or synonyms are retained. The insect host names have been checked by Dr. E. A. Chapin and the late Dr. H. G. Dyar of the Federal Bureau of Entomology through the courtesy of Mr. Harold Morrison, Chief of the Division of Taxonomy, and the crus- tacean host names have been checked by Dr. Waldo Schmitt of the U. S. National Museum, and I wish to acknowledge my indebtedness to these workers for their assistance. ARTHROPODS AS INTERMEDIATE HOSTS OF CESTODES The known number of arthropods acting as intermediate hosts for tapeworms is so small that this subject can be covered rather compre- hensively. At the same time, one must generalize here as elsewhere rather carefully, since we know the life histories of only about 1 per cent of the known tapeworms. In addition to arthropods, the inter- mediate hosts of tapeworms include mammals, birds, reptiles, am- phibia, fish, mollusks, annelids, and other animals. In all probability many worms now known only as having one intermediate host will be found to require two successive intermediate hosts. The following list will show the tapeworms, their primary hosts, and their inter- mediate hosts, for such tapeworms as have arthropods as intermediate hosts. ANOPLOCEPH ALIDAE It is still true that the life histories of the anoplocephaline tape- worms are unknown. The larval cestode which has been reported from Aphodius obscurus and tentatively referred to Cittotaenia marmotae has not been definitely coupled with that worm by the test of success- ful feeding experiments, and the record is of value primarily as a possible clue to solving the unknown life histories in this group. 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I5 ARTHROPOD HOSTS OF HELMINTHS—HALL 27 An analysis of the records given shows the following: PLAGIORCHIIDAE In the Plagiorchiidae we have a group of flukes which have a wide range of intermediate hosts, including the insect groups Odonata, Diptera, Trichoptera, Plecoptera, Ephemerida, and Coleoptera, and the crustacean groups, Decapoda and Amphipoda. This range of intermediate hosts is associated with the range of primary hosts, which include fish, amphibians, and birds. Considered on the basis of primary hosts, the plagiorchids in birds utilize Odonata as inter- mediate hosts, those in fish use the Diptera, Odonata, Trichoptera, Plecoptera and Decapoda; while those in frogs use the Odonata, Tri- choptera, Coleoptera, Ephemerida, Plecoptera, and Amphipoda. The Trichoptera and the Odonata appear to be the most important inter- mediate hosts. LECITHODENDRIIDAE In the Lecithodendriidae the insects serve as intermediate hosts and they include the Plecoptera, Ephemerida, Coleoptera, Diptera, Tri- choptera, Odonata and “ amphibious insects.” The frog flukes of this family use Odonata and Coleoptera as intermediate hosts; the bat flukes use Plecoptera, Ephemerida, Diptera, and Trichoptera. Here also the insects have the double réle of intermediate host for the fluke and of food for the primary host. OPISTHORCHIIDAE In the Opisthorchiidae, insects, specified by Stafford (1927), as amphibious insects, are the only reported hosts. Since this is a large family with a wide range of hosts, little of a general nature could be concluded from the foregoing. ALLOCREADIIDAE In the Allocreadiidae, parasitic for the most part in fish, the inter- mediate hosts include Ephemerida, Trichoptera, Diptera, and Deca- poda, the more important being the Ephemerida and the Decapoda. The intermediate hosts probably serve as such by virtue of their role as food for fish. The record for Astacotrema cirrigerum of a bird as primary host is found in a footnote reference based apparently on correspondence and lacks evidence or detail. } } | \ ¢ t Soe ge te I ts ee 28 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 81 GORGODERIDAE In the Gorgoderidae, commonly parasitic in frogs, the intermediate . hosts known at present are mostly Odonata, the role of intermediate host here being combined with the role of food for frogs. One of the Decapoda, a crayfish, is the host for a gorgoderid parasitic in fish. HALIPEGIDAE In the Halipegidae, the only reported life history involves the Odonata as intermediate hosts, the primary hosts here being frogs. TROGLOTREM ATIDAE In the Troglotrematidae the only known life history, that of the human lung fluke, involves several species of decapods, crabs being known hosts and crayfish probable hosts. HEMIURIDAE In the Hemiuridae, which are fish parasites, all known intermediate hosts are crustaceans, those for two flukes being copepods and those for one fluke being decapods. DICROCOELIIDAE The one dicrocoelid with a known life history utilizes an amphipod as an intermediate host, the primary hosts being fish. BRACHYCOELIIDAE The one brachycoelid with a known life history has a trichopteran as an intermediate host, the primary hosts being amphibians. FAMILY UNCERTAIN The three flukes of uncertain relationship for which we know primary as well as secondary hosts, and not merely secondary hosts for larval stages, all have carnivores as primary hosts and crabs as sec- ondary hosts. ARTHROPODS AS INTERMEDIATE HOSTS OR NEMATODES In listing thé nematodes having intermediate stages in arthropods, no attention has been paid to nematodes listed only as nematodes without reference to whether the nematodes were mature or immature. Nematodes occurring consistently as larvae in insects may be the RENE ENE RL ES A PME NT BET NO. 15 ARTHROPOD HOSTS OF HELMINTHS—HALL 29 larvae of worms which will develop to maturity on reaching a suitable host, although larval nematodes specified as such with no further discussion may be the larvae of such worms as the mermithids which will develop to maturity as free-living forms. In this paper the mermithids and gordians are not considered, as they are not regarded as true parasites of vertebrates in the scope of treatment of that sub- ject as limited here. The gordians may parasitize immature frogs in the course of development of the worms, but this topic is disregarded here owing to a lack of space for its consideration. All records which are merely surmises to the effect that a certain arthropod is the intermediate host of some nematode are likewise disregarded. Such surmises have their value in directing exploratory research, but for the purpose of analyzing existing records to obtain valid data they are worthless. There is sufficient uncertainty in con- nection with a number of existing records to introduce certain ele- ments of possible error as it is. The following list covers the important cases of arthropod hosts for nematodes. The worms involved fall in the Filariata or Filarida and most of them fall in the superfamilies Spiruroidea and Filarioidea, two closely related superfamilies which are markedly heteroxenous and hence in sharp contrast with most of the other nematode groups which are usually monoxenous. In the exceptional cases in which members of other superfamilies utilize intermediate hosts, the hosts are never arthropods so far as the writer is aware, but are such forms as fish or earthworms. SPIRURIDAE As intermediate hosts of nematodes of the Spiruridae, which is made up predominantly of mammalian parasites and to a lesser extent of bird parasites, the Coleoptera are of outstanding importance. In this family the common mode of transmission of the larval worm to the primary host is by means of the ingestion of the secondary host, either as a deliberate act of eating or because of the more or less accidental presence of the secondary host in the food of the pri- mary host. In general, dogs, sheep, cattle and horses cannot be called insectivorous animals, but the presence of beetles in their customary food seems to be sufficiently common to enable various spirurid parasites of these animals to maintain themselves with the aid of these beefle hosts. It is evident that some of the spirurids utilizing beetle hosts may have alternative life histories which are more compli- cated than the mere infection of the beetle host by means of infective worm eggs and the infection of the primary host as a result of swal- lowing infected beetles. 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OlEUNIS |p te ost eye coe eres hos Dom e1oydiq reresess spynUMS (¢) eee ewes ‘ds vajng (2) seer eee ee OVC fence les orca hea ae aacael uejyy eee SUIUYNIIDI DIAIIOINC “MUN{D] -nIvUuiiAy UiNndAquaIod Giz] pDAIHIpAOI DIOGO;DUAD FY es1azdiq { e1WIG sae wnsouupp wnynuns eee ee eeoe ayeullig Pe ee ue eeeee snjnajo2 DIAIIOING eee eeeree aVaWav 1 dno1y ysoy ATepuosasg dnoin ysoy AIvUIIg apozeuls Ny Apimae yy See ee ee ee ee SS ES eS ee panulju0j—sapojnmany fo sjsoyy podosyjap 4 co =| o > SMITHSONIAN MISCELLANEOUS COLLECTIONS “snaumnb epluyoery -ups snyoydasidiyy|******* JtOATUTeD | ttt ttt tts Bog|*** ussps6 pmauojpjagiq ‘ds spbap epluyoely “DyDg -NOU SNAOPOY}UAC elaydeuoydig} "+ +++ * Suppad xajng *s1u0 -tuanjuad Sdostsy) (2) tree eeee dg sapydoung “smmsof “lun Saprommosun py “** aupypéb Saproaryny “8% quagsnp Sapiozyny eloyuqd ‘snyouuad -oasnf sdouozoshkay) "su -uadynspu sajayqdoup “'* sypjsod sajaygoup eee eeee suargrd xvamng sesesss Suains sapap 0) 6 e0sepe10 udxban Sapap sere cones oye g a afieXe,00 0), 4)'0'7s).0\\07,0 eee 6 ueyq **supjsiad puauojpjagiq cece eer ee AVGIINV IL] dnoin ysoy AIBpUODaS dnoin ysoy ALVWIIG apozeula Ny Ayre T ponunuoj—sapojpmany fo sjsopy podosyjp © ~ “**pupuays Didnauobhr “MUNAD ~ epliowialydy { iat 40 P : “*'* pipbyna vaamaydy see eee e eee eens AT kcattes eG PN (es OSE are é -luamay ga p1ajzgouig S$ e1oj}diq “""* SNIDPUNIAI SNUDQD J, see ewe stew ewes é is) A) Be Le), 6 (ehefeleleles) 6) (06) 97elere . é **pjooupqn4 nippy oul p eia}diq eee apuibId-IDAAI4 DISHETAT Nn oes eee ee dl | fetenno ers olsir nice Bei Ne a Pp SLE DANALGSOULDB P| * +2080" NIVIYION cece eces 01906 Snjj0J i he mpj4aga SNAIUMS (-) S88 SNIYNA SNISIINAT eter eters omens tee epodatio>) foccttttss cds snmozqoiq tees pquua snupsgp eee a inlets “Us asgorsig) "* 8 SLUDBINA SnissDADI]|** Snamnbups DAJamopiYy J “SND epodado)|**** suzopidsnzig sgopahD| +++ +++" apyday|:*** puyuadsas vapkjay)| -yda20qoj6 snjnounapaq epodadoy eee eee en esos ‘ds sgojaky se wee wen ee atyday sees seeee eiqoo ueIpuy OI ‘ds SNINIUNIDAC se snnuays $gojak) “*snyopidsna1q sqoj9ky vse snuspig sqojaky sess 1499uUuaG DIJIZD4) epodaido7 see eee SUpt aia sdojaty y ee yeory e- smsoo.aponb sdopat9 see eeee soyepnsuy) sie raiekavelwidveivne elena daays "t9* mgapyona, $qoj9ky ar ae onda apieg “*** snypuosod sdoj9ky | Msensile Ke Nohen eden nee hens ds1OF] cece cccsecee piedooy POOR OR SOS ODP 751g aS eet OE LUD TMT hei peat ere tee ia 2 ae eG “sisuauipaut SNJNIUNIDA se ee* FTVGITNIONNIDVA rd f 4 ARTHROPOD HOSTS OF HELMINTHS—HALL ponies SSOIOATUAES) { dno1in ysoy Arepuossg dno1n ysoy AreUIIg apoyeUis Ny Ayre NO. I5 psnunu0j—sapojpmayy fo sjsozy podoayjap ae a mh ll a oO J Oo > SMITHSONIAN MISCELLANEOUS COLLECTIONS e19qdiq $06) 01000 a efecetecve ‘ds DIUUD elaydoajog|****** snuypoapa snjourg e19}d0a[0D|**** Sniupjauyf Smipoyd Pr erojdeuoydis fo cg sudoays vyksqouay L sngwvospf snygtydowasg elaydogjog}******* vbpsyaom sdvjg we « eee eroydiq|**** sunjdas mnynuusng|****** dnoin ysoy Aepuosas . . . . . eee ee ewes @ |]/0: ¢, exe ee eee ence elecese eer ees eoe es leeee teres sng fo cct snuayjns sasuadvup mlL cette: osny sasuagiap dnoiy ysoy ATeULIg ‘OzOI ‘jJoroueg ¢|uojsuyof JO vluoU [eAIeT a. a. é ‘veOr ‘Wweig jo euou [eAIeT ‘ZzO1 ‘qqod jo eWou [eAIey “E161 ‘uojsuyof ‘ds vinauowpl p ‘aDHVSyAoUut -Sgv]q wnpoymmauouDnb py “** stuasuagian sisgojsk) opoyeUuta Ny cgieleieieinis © NIT VLUGION () Ayruregy ponunuoj—sapojnman fo sjsozy podowypip NO. 15 ARTHROPOD HOSTS OF HELMINTHS—HALL 45 sexalatus, a spirurid parasite not uncommon in the stomachs of swine and peccaries, will develop to an infective third-stage larva in its beetle host, and when these beetles are fed to some unusual host, such as rodents, birds, or even cold-blooded animals, the larval worm will re-encyst as a third-stage larva in the unusual host ; but if the infected unusual host is fed to a suitable primary host, the larva will continue its development to maturity. How extensive this device is we do not know, but it may prove to be a common means of transmitting the spirurid worms of rapacious birds, as Cram has suggested, the spiru- rids of these birds producing eggs which infect some arthropod host, such as a beetle, the beetle being eaten by a small mammal, bird, amphibian or reptile, which is infected in turn with the third-stage larva, and the bird of prey eating these animals and becoming infested with the adult worms. The investigation of these life histories is a thing on which the mammalogist, the ornithologist, the herpetologist, the entomologist and the parasitologist might collaborate to great’ advantage, and the results might show some very interesting and surprising biological interrelationships. As intermediate hosts of spirurids, the Orthoptera are also of some importance. The arthropods in question are all cockroaches, and they are probably of special importance as intermediate hosts for parasites of such rodents as rats and mice. These rodents seem to eat cockroaches with dependable certainty, and the association of rats, mice and roaches in the household provides a suitable and, so to speak, natural combination of factors for the benefit of these spirurids. On the other hand, the development of spirurid parasites of sheep, cattle and horses in cockroaches must be regarded as a case in which the roach merely serves as a host for a worm which cannot depend on such a host for its transmission, but which is capable of developing in that host as a case of accidental parasitism. In this connection it may be noted that roaches will serve as intermediate hosts for so many worms in this way that these insects make excellent experiment ani- mals for carrying out life-history experiments in the laboratory. The plentiful supply of these insects in winter, a thing so unfortunate from some points of view, is a fortunate thing for the parasitologist who obtains interesting worms in winter at a time when other insects are searce, and wishes to carry out feeding experiments on some insect. The Diptera appear as intermediate hosts of spirurids with 14 species serving as hosts for 3 known species of spirurids, all species of the genus Habronema and all parasitic in horses and other mem- bers of the Equidae. This association obviously depends in part on the importance of the manure of horses and other Equidae as a breeding a eS ———— | f \ : 46 SMITHSONIAN MISCELLANEOUS COLLECTIONS voL. 81 place for Diptera. The transmission of the worm from the fly to the horse appears to present several complications. It was surmised by Ransom that horses might swallow flies which had fallen in drinking troughs or were benumbed in feed troughs on cold mornings. Sub- sequent work has indicated that the worms may escape from the proboscis of flies as they feed on the moist lips of horses, and pre- sumably these worms may get to the stomach and develop to adult worms. However, if the fly feeds on the conjunctiva of the eye the larval worms may escape to the eye, remaining there as larvae and causing a habronemic conjunctivitis. If the fly feeds on a wound, the worms may escape and remain in the wound as larvae, causing “ sum- mer sores” or cutaneous habronemiasis. Finally, the worms may be found as larvae in the lungs, causing pulmonary habronemiasis, but the precise method of infection here remains to be ascertained. These cases illustrate the fact that there are numerous deviations from the cut-and-dried rule that intermediate hosts either transmit worms by being eaten by the primary host, or else transmit the worms by biting the primary host. One member of the Siphonaptera occurs as a somewhat doubtful host of a rat spirurid, Protospirura muris, but the case for this should be developed by feeding experiments. The one bird nematode of the family Spiruridae having a known life history is Hartertia gallinarum, and this worm utilizes a termite as its intermediate host, the host here serving as food for chickens which devour them with great eagerness. THELAZIIDAE In the Thelaziidae, we have a member of the Orthoptera, the roach Pycnoscelis surinamensis, serving as the intermediate host of the chicken eyeworm, Oxyspirura mansoni, and also for the somewhat dubious species, O. parvovum, distinguished from O. mansoni only by the smaller size of the egg. This life history was worked out by Fielding in Australia and somewhat later, but independently, by Sanders in Florida in the United States. At present the eyeworm, O. mansoni, appears to be confined in the United States to Florida, so far as our records show, but the intermediate host now has a much wider range in this country and unless measures are taken to stamp out the worm in Florida we can confidently expect it to spread beyond the confines of that state. The movements of the infected primary and secondary hosts by the swift methods of modern transportation over wide areas can hardly fail to ensure this result. [Since the above was written, the eyeworm has been found outside of Florida in this country. | NO. I5 ARTHROPOD HOSTS OF HELMINTHS—HALL 47 ACUARIIDAE In the Avuariidae, we are dealing with bird parasites. Of the two worm species involved, one is a parasite of water birds, Anseriformes, and it is not surprising to find that this worm, Echinuria uncinata, uses Cladocera as its intermediate hosts, the one known intermediate host being Daphnia pulex. The other worm is a parasite of land birds, Galliformes and Columbiformes, and utilizes an isopod, Por- cellio laevis. [Cram has since found grasshoppers to be intermediate hosts for Acuariidae of terrestrial birds. | TETRAMERIDAE In the Tetrameridae we are again dealing with bird parasites, and here again the intermediate hosts are Entomostraca, a cladoceran, Daphnia pulex, and an amphipod, Gammarus pulex. The one worm for which we know the life history, Tetrameres fissispina, is usually and normally a parasite of water birds, Anseriformes, and its occur- rence in land birds must be regarded as following from the accidental swallowing of the infected entomostracans while drinking, whereas in water birds we are dealing with a dependable arrangement, from the standpoint of the parasite, based on Entomostraca in the double role of food for the primary host and of secondary host for the worm. [Cram has recently found grasshoppers serving as inter- mediate host of tetramerids of terrestrial birds. ] CUCULLANIDAE For the one cucullanid with a known life history, a fish nematode, copepods and aquatic isopods serve as intermediate hosts, the hosts also serving as food for fish. CAMALLANIDAE For the two camallanids with known life histories, one a fish nematode and one a turtle nematode, copepods are hosts for both and dragonflies also serve as hosts for one. These hosts are also food for the primary hosts. HEDRURIDAE Of two species of hedrurids, parasitic in reptiles, amphibians and fish, one uses aquatic isopods and one amphipods as intermediate hosts. The foregoing families are regarded by many parasitologists as part of the superfamily Spiruroidea, and in this superfamily the life history is usually one in which the transfer of the larval nematode to the primary host is accomplished when this host swallows the sec- 48 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 81 ondary host, either as food or accidentally, such apparent exceptions as in the case of Habronema being the unusual thing. We now take up a group of worms which all fall in the family Filariidae, regarded by those who recognize the superfamily Spiruroidea as described above, as being part of the superfamily Filarioidea, the two super- families being grouped on their affinities as the Filariata. FILARIIDAE In the Filariidae the customary mode of transmission of the worm is by the bite of the secondary host, this host becoming infected when it bites an infested primary host and in turn infecting a primary host by biting it after an interval in which the worm develops to the in- fective stage in the secondary host. It is to be expected, then, that the biting Diptera will show up prominently in this list of intermediate hosts, and we find a long list of such hosts recorded as transmitting numerous species of filarid worms. Here we have a number of im- portant worm parasites of man and dogs, including such filarids as Wuchereria bancroftt and Loa loa of man, and the heart worm, Dirofilaria imnutis, a serious pest of dogs in the hunting field. Mosquitoes take first place in this group of Diptera, many species transmitting W. bancrofti and D. immutis, while the tabanids, espe- cially Chrysops spp., function for Loa loa. The Siphonaptera, Mallophaga, Anopleura, and Arachnida are all charged with the transmission of filarid worms. DRACUNCULIDAE In the peculiar genus Dracunculus, including the guinea worm of man, D. medinensis, the worms usually infest superficial body parts of the primary host, and when these come in contact with water, the worms release large numbers of embryos, some of which are swal- lowed by copepods. The larval worms develop in these hosts to the infective stage and when these hosts are swallowed by suitable pri- mary hosts the worms develop to maturity. ARTHROPODS AS INTERMEDIATE HOSTS OF ACANTHOCEPHALIDS The acanthocephalids quite generally utilize at least one inter- mediate host, and sometimes two such hosts, the second one a fish, amphibian, or bird in some cases, in their life history. These first intermediate hosts are sometimes snails or leeches, but in most of the reported cases the first intermediate host is an arthropod. The follow- ing list shows the reported hosts for the species having known life histories : 49 ARTHROPOD HOSTS OF HELMINTHS—HALL NO. 15 vreees ueiqrydury |‘ * °°" DyuanIsa DUDY sete teens apday| ttt * Stapjnaiqso sku Freese sess pou DIME seen e wees orf omjps : sees SUDAN DIAG teeeeeeeeess p07 D407 sets s SNYNd SNISIINAT snuxoyg snasianay “snuyDyy -Ygodypisa SnIsinaTy srsstss Sup snIzsianaT "** snosiana] snasignaT er eee eee ne 01gob 01g0+) smyibundg Snajsosajsv+) ps Re SNJDIINID SNIJSOAAISD+) seseeeeees gmigany xosy srrees ordapa snuisghy seeeeess mruany s1441q09 ** SUSDU DUOJSOAPUOY 4) *** SMISSDADI SNISSDADD sseees sngang snqwg . ero\do.mayy fcr ct ss: pianyny simi sorts pinbun pynbup ere ci ory SONG sss QnUdad DULADIL “** snuangip Snuéng 7 sees pUDAg SWWDAGP “dvd "59s puydsolg sumpaégP |\'yins SnysutysouryII0d N'| -IHONAHYONIHOIOAN, dnoin ysoy AIepuUddIG dnoin ysoy AIeULIg preqdoooyjueoy Ayrure iT spypydaso0yjunop fo sjsopy podosywsp = co S| S > SMITHSONIAN MISCELLANEOUS COLLECTIONS 50 focrtt syymioany snovisp Lott snovjsp sniqompjog **** pISNI0] SHADULULDS) diyd epodiydury { sres xvamndg SnADULULDS) epodessq [ "++ snakyos saqaksojnx “e+ snupynl snbaypa4g ‘+ psobna vbpygoXy J ** suaasaf vbopy doy gq eta}dozJOD}< suamayaa vbvygozjcyg * snaapqn snaapoqgop” "+ srapbjna DYy4Uoj0]2 YY DYJUOJO]AIM DYJUOIOIa PY Loi tttt* pypann piu0ja7y eB19}d09]09) i ‘ds sqv1g dnoir ysoy AIepuosasg so 5+ -IMOy JOJVM PIL AA Snrysamop 4010 snub\) "t 98 SQUIIOJIIOsSUYy ‘sng -SIMOP SNI4IUMII ABSUpy l DIUSIMOP SPYISOg SDUP eens sajomag ls) ae uryy Losses: smpyaniwl snqaa °°* saJOATUIeZ) i Sa sovensug |f ; ose 52°" SIOATOOSUT |“ * ees esa PUTT seeeee SOIOATUIC*) S forreresssssss eudy sya seeees SnadID SUD) sees SHIDISIAI SNS aisle serene SUT * snavbyp SsnaIDUIag “** snauunag AniuaT] “** snippnnIa SNNUT "** snjjanjof sngay ses" 4040] UOhIOAd ete e eens rut] SY ay “*sippyssog snydsomfjog|***** AvdIINOSONAYOD “snagDUIpn.Ary ‘aVaIHO SNYIUKYLOYIUDIDAID | + -NKHYOHLNVOVDITIO "888 snaanp suipy|*ojnaids snysudys0j;uvb14) | *IVdIHONAHUAOLNVODIL) ysoy ATVUTIG preydsooyjueay Ape penurju0j—'spypydar0yjupop fo sjsopyy podosyjap ARTHROPOD HOSTS OF HELMINTHS—HALL 5! NO. I5 epodiydury|****"** vajynd snapunun+y avavenctale SYDJUIIAO D}IDIG e19}d0 f , OMZON -punaisaun vyaunjgisag|*** f ses DIMUOsAINUL sqoig e.193d03]07) Lotter spb sdoig dnoin ysoy AIepuods9S SOWIOFLIYDOY | °° ayeullig|*: OIOATUIPD | °° ee oeee SJUspoyy dno1n Feee ss usiy tayo ARTY ‘dds owns eevee Nenacs “+ smonj xosy snapgpr snuraday eee eee s++ “dds snqjoD uuuvutzapar Snuobasoy DjNjog4nqg $141G0) “++ snap auojag se stupbhjna vypnbup osny dasuaqi py MADBINAZ DUIAIIP mrs pQUa SUMDAGY DULDAG SIUMDAQY "" podyg SuuDAgRr SNIMAIMII OI]DT uey Smisojng D1aISNn TY "2558" SUDQAD DIOP snuzsanb snxok py 788" SUIDZAD SNIOLIN TY SmapjuauMnAf SNJIIIAD mre srusomosnf snpy reeeeeseeseees genow Serene etree eeeeee gpye ee ee www eee wee ewe cee. ysoy AleWIIg “*s1aan] snysukysoyquod|* AVGIHONAHAYONIHOY “smmsof -ymom Simsofymopy|**** aAVvAaIWuOATTINOW preyqdoooyjyuroy Ayre panunuoj—'spypydas0yjuvop fo sysop] podosyjip VoL. 8I SMITHSONIAN MISCELLANEOUS COLLECTIONS 52 "* WIAA UOIPAJSOSYS "+ Snsojnqau snamaup "'* pypaqsod DINbUp™ shhh) SNIDINIYIA LOST lee Veer StI OS ET sees * SUIISIQBDY DIAI Peeeeeeeeees using ¥ d : se SapoAdpa DULIAI TI Sijsadna sayygojqup SIPIOULIDS SNAIZFOAILY NIWMOJOP SNAIIGOAI WUOSAIWUMOD SNULOJSOJDI epodiydury| imayoquayouy 1]2]08 "*) SnuDI4aULD SNIIOY |SNIDIIYL SNYyIUAYsLOUIY IZ tere ee eens odds UOpld [ "5 DAD DAPUDUDIDS ee eee eee ee SUPA ofng sreceees srupbna ofng “8 snauby Ao;pUMquio g seesess piuanImsa DUDY "t+ mlapsoguiay Duby |*** avUDA SnYyIUKYsOUIY IF se eee sueiqiyduy epodosy|******* sn2yonbo snpyjaspy epodiydury|***** ++ 2oy piasogojuod sc #9 °USig. TouIO: AUBIN, eee e eee see 01qg06 01qg0+) SNIDIINID SNIJSOAIISD!) reeeeeeess gmgnp xosq seeees organs snuigey Gu eilen@ \0.:6 10:,6, 9119), elle ysty soe ei > 91g0b SNI09 sseeees “dds snuo0bai07 vresess snaing Ssnq4ng ss pyinbun vppnbup sts DNUNAII DULAIIP epodosy|****** snoonbo snjjasy ( "rots puipAg SuuDAgE?|***uINn, SNDYgaI0YJUDIP |* AVGIHONAHYONIHOY | dnoir ysoy AIBpuodaS dno1ry ysoy ATEUITIg preqdacoyjueoy ATIWMe es nS penunuoj—'spypygas0yjuva py fo siso fy podoayiap NO. 15 ARTHROPOD HOSTS OF HELMINTHS—HALL 53 An inspection of the tables indicates, first of all, that we have but limited knowledge of the life histories of acanthocephalids in any one family, and that, it will not be possible to generalize to any great extent on such limited data. What may be said is as follows: NEOECHINORHYNCHIDAE In the Neoechinorhynchidae we know the life history of one acanthocephalid, a parasite occurring in a large number of fish and in some reptiles and amphibians, and the known intermediate hosts are species of Sialis, one an unrecognized species, in the Neuroptera, other hosts being leeches and snails. GIGANTORHY NCHIDAE In the Gigantorhynchidae we again have only one known life history. In this case the echinorhynch occurs as an adult in mammals of various groups, including primates, carnivores, and insectivores, and has a species of Blaps, a coleopteron, as an intermediate host. OLIGACANTHORH Y NCH IDAE In the Oligacanthorhynchidae we again have one acanthocephalid with a known life history, the well-known thorn-headed worm of swine, occurring in such animals as swine, carnivores, and man and other primates. This worm has a number of species of scarabaeid beetles as its intermediate hosts. CORY NOSOMIDAE In the Corynosomidae we have one known life history, that of an acanthocephalid of water fowl, Anseriformes, using crustaceans, amphipods and crayfish, as intermediate hosts. MONILIFORMIDAE In the Moniliformidae we have a parasite with a wide range of primary hosts, from man, carnivores and rodents to rapacious birds, and having as its intermediate hosts two species of Coleoptera and two of Orthoptera. ECHINORHYNCHIDAE In the Echinorhynchidae we have three acanthocephalids parasitic in fish of numerous species, two of them with an amphipod as an intermediate host and one with an aquatic isopod as an intermediate host; and one acanthocephalid parasitic in various amphibians and with an amphipod as its intermediate host. 54 SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL. 81 By way of summary it may be noted that of three acanthocephalids parasitic in mammals, all develop in insects, all with Coleoptera and one with Orthoptera also as intermediate hosts ; the one acanthocepha- lid habitually parasitic in water birds uses crustaceans as intermediate hosts ; and that of five acanthocephalids parasitic in fish and amphib- ians, four use crustaceans as intermediate hosts, these being amphi- pods in two cases, isopods in one case, and both amphipods and isopods in one case. In the case of one acanthocephalid in fish, the Neurop- tera serve as hosts. Insects are apparently of major importance for acanthocephalids of mammals and crustaceans for acanthocephalids of fish. In the foregoing lists of parasites arranged by orders and families, the names given for the arthropod hosts are those under which they are reported in the literature and no attempt is made in these lists to eliminate synonyms for the reason already given that it is easier to trace these references in the literature under the names quoted. In the following lists arranged on the basis of intermediate hosts, syn- onyms are cross-referenced to the names accepted by the authorities already mentioned in the first part of the paper. ARTHROPOD HOSTS OF HELMINTHS, ARRANGED BY HOST GROUPS INSECTA ANOPLEURA Ephemerid Haematopinus piliferus. See Linog- Lecithodendrium lagena nathus piliferus. Cercaria secunda Linognathus piliferus Dipetalonema reconditum DERMAPTERA Anisolabis annulipes Hymenolepis diminuta Hymenolepis microstoma EPHEMERIDA Blasturus cupidus. See Leptophlebia cupida. Cloeon dipterum ? Opisthioglyphe endoloba Ephemera danica Stephanophiala farionis Ephemera vulgata Allocreadium isoporum Opisthioglyphe endoloba Spiroptera ephemeridarum Hlexagenia sp. Crepidostomum cornutum Stephanophiala farionis Leptophlebia cupida Allocreadium commune Oligoneuria rhenana Spiroptera ephemeridarum COLEOPTERA Ablattaria laevigata Weinlandia uncinata Akis goryi Spirura gastrophila Spirocerca sanguinolenta Akis spinosa Hymenolepis diminuta Aphodius castaneus. See Aphodius rufus castaneus. Aphodius coloradensis Gongylonema scutatum NO. 15 Aphodius distinctus Gongylonema scutatum Aphodius femoralis Gongylonema scutatum Aphodius fimetarius Protospirura gracilis Gongylonema scutatum Larval nema of Cobb, 1922 Aphodius granarius Hymenolepis carioca Gongylonema scutatum Aphodius haemorrhoidalis ? Gongylonema pulchrum Aphodius obscurus ? Cittotaenia marmotae Aphodius rubeolus Gongylonema scutatwin Aphodius rufus Arduenna strongylina Aphodius rufus castaneus Arduenna strongylina Aphodius sp. Gongylonema scutatum Aphodius vittatus Gongylonema scutatum Ateuchus sacer. See Scarabaeus sacer. Ateuchus sp. Physaloptera abbreviata Blaps appendiculata Gongylonema scutatum Blaps emondi Gongylonema scutatum Blaps gigas Moniliformis moniliformis Blaps mortisaga Agamonematodum blapis-mortisagae Blaps mucronata Moniliformis moniliformis Blaps spp. Spirura gastrophila Gongylonema scutatum Gongylonema brevispiculum Gigantorhynchus spirula Blaps strauchi Spirura gastrophila Gongylonema scutatum Gongylonema brevispiculum Caccobius schreberi ? Gongylonema pulchrum ARTHROPOD HOSTS OF HELMINTHS—HALL 55 Canthon sp. Spirocerca sanguinolenta Cetonia aurata Spirura talpae Macracanthorhynchus hirudinaceus Chironitis irroratus Gongylonema mucronatum Copris hispanus Spirocerca sanguinolenta Diloboderus abderus Macracanthorhynchus hirudinaceus Geotrupes douei. See Geotrupes (Ste- reopyge) douei. Geotrupes (Stereopyge) douet Spirocerca sanguinolenta ? Physocephalus sexvalatus Gongylonema mucronatum Geotrupes (Anoplotrupes) stercorosus Choanotaenia infundibulum Hymenolepis serpentulus ? Physocephalus sexalatus Geotrupes stercorarius Physalocephalus sexalatus Geotrupes stercorosus. See Geotrupes (Anoplotrupes) stercorosus. Geotrupes sylvaticus Choanotaenia infundibulum Hymenolepis serpentulus Gymnopleurus mopsus Gongylonema mucronatum Gymnopleurus sturmt Spirocerca sanguinolenta Gongylonema mucronatum Ilybius fuliginosus Haplometra cylindracea Ilybius sp. Cercaria prima Melolontha melolontha Macracanthorhynchus hirudinaceus Melolontha vulgaris. See Melolontha melolontha. Onticellus fulvus Gongylonema scutatum Onitis irroratus. See Chironitis irro- ratus. Onthophagus bedeli Physocephalus sexalatus Gongylonema mucronatum 56 SMITHSONIAN MISCELLANEOUS COLLECTIONS Onthophagus hecate Arduenna strongylina Physocephalus sexalatus Gongylonema scutatum Onthophagus nebulosus Physocephalus sexalatus Onthophagus pennsylvanicus Gongylonema scutatum Onthophagus sp. Spirura gastrophila Onthophagus taurus ? Gongylonema scutatum Phyllophaga arcuata Macracanthorhynchus hirudinaceus Phyllophaga fervens. See Phyllophaga fusca. Phyllophaga fusca Macracanthorhynchus hirudinaceus Phyllophaga rugosa Macracanthorhynchus hirudinaceus Phyllophaga vehemens Macracanthorhynchus hirudinaceus Pinotus carolinus Larval nema of Cram, 1924 Scarabaeus sacer Spirura gastrophila Spirocerca sanguinolenta Physocephalus sexalatus Gongylonema mucronatum Scarabaeus variolosus Spirocerca sanguinolenta Physocephalus sexalatus Scaurus striatus Hymenolepis diminuta Silpha laevigata. See Ablattaria lae- vigata. Strategus julianus Macracanthorhynchus hirudinaceus Tenebrio molitor Hymenole pis arvicolae ? Hymenolepis nana Hymenole pis diminuta Hymenolepis microstoma Onchoscolex decipiens Protospirura muris Gongylonema neoplasticum Tenebrio obscurus Gongylonema sp. Tribolium ferrugineum Hymenolepis diminuta VoL. 81 “Water beetles ” Pleurogenes medians Pleurogenes claviger Pleurogenes confusus Xyloryctes satyrus Macracanthorhynchus hirudinaceus DIPTERA Aedes aegypti Filaria ozzardi Wuchereria bancrofti Dirofilaria immuitis Dirofilaria repens Dipetalonema perstans Aedes albolineata Wuchereria bancrofti Aedes albopictus Wuchereria bancrofti Aedes caspius Dirofilaria immitis Aedes fasciatus. See Aedes aegypfti. Aedes (Finlaya) togoi Wuchereria bancroftt Aedes gracilis. See Bironella gracilis and Anopheles gracilis. Aedes perplexus Wuchereria bancrofti Aedes pseudoscutellarts. variegatus. Aedes punctatus. See Aedes caspius. Aedes scutellaris. See Aedes albopic- tus. Aedes sugens. See Aedes vittatus. Aedes vagans Dirofilaria immitis Aedes variegatus Wuchereria bancrofti Aedes vexans Dirofilaria immitis Aedes vigilax Wuchereria bancrofti Aedes vittatus Dipetalonema perstans Anastellorhina augur Habronema sp. Anopheles albimanus Filaria oggzardi Wuchereria bancrofti Anopheles albitarsis Filaria ozzardi See Aedes NO. 15 Anopheles algeriensis Dirofilaria immitis Anopheles annulipes Wuchereria bancrofti Anopheles argyritarsis Wuchereria bancrofti Anopheles barbirostris Wuchereria bancrofti Anopheles bifurcatus Dirofilaria immitis Agamodistomum martiranot Anopheles claviger. See Anopheles bifurcatus. Anopheles costalis. gambiae. Anopheles culifaciens Agamodistomum sintoni Anopheles fuliginosus Cercaria of Stephens & Christophers, 1902 Anopheles funestus listoni. See Ano- pheles listonit. Anopheles gambiae Wuchereria bancrofti Dipetalonema perstans Anopheles gracilis ? Wuchereria bancrofti Anopheles hyrcanus pseudopictus Dirofilaria immitis Anopheles hyrcanus sinensis Wuchereria bancrofti Dirofilaria immitis Anopheles listonii Agamodistomum sintoni Anopheles maculipennis Lecithodendrium lagena Agamodistomum anophelis Filaria ozzardi Filaria sp. Fuelleborn, 1909 Dirofilaria immitis Dirofilaria repens Dipetalonema perstans Anopheles palestinus. See Anopheles super pictus. Anopheles rossi. See Anopheles sub- pictus. Anopheles sinensis. hyrcanus sinensis. Anopheles sinensis peditaeniatus Wuchereria bancrofti See Anopheles See Anopheles ARTHROPOD ‘HOSTS OF HELMINTHS—HALL 57 Anopheles sinensis pseudopictus Dirofilaria immitis Anopheles sinensis vanus. bheles barbirostris. Anopheles subpictus Cercaria of Soparkar, 1918 Cercaria of Stephens & Christophers, 1902 Wuchereria bancrofti Anopheles superpictus Wuchereria bancrofti Dirofilaria immitis Anopheles tarsimaculatus Filaria ogzardi Bironella gracilis ? Wuchereria bancrofti Chironomus libiferus Lissorchis fairporti Chironomus plumosus Lecithodendrium lagena Chrysoconops fuscopennatus. See Mansonia fuscopennatus. Chrysops centurionis Loa loa ? Dipetalonema perstans Chrysops dimidiatus Loa loa Chrysops longicornis Loa loa Chrysops silaceus Filaria sp. of Med. Rept., Lagos, Nigeria, 1918 Loa loa Corethra sp. Cercaria prima Cercaria secunda Culex ciliaris. (May be Aedes cin- ereus, fide Dyar.) Wuchereria bancrofti Culex fatigans. See Culex quinque- fasciatus. Culex fuscocephalus Wuchereria bancrofti Culex gelidus Wuchereria bancrofti Culex hortensis Cercaria of Joyeux, 1918 Culex malariae. See Aedes vexans. Culex microannulatus. See Culex siti- ens. See Ano- 58 SMITHSONIAN MISCELLANEOUS ‘COLLECTIONS Culex penicillaris. See Aedes caspius. Culex pipiens Wuchereria bancrofti Dirofilaria immitis Dipetalonema perstans Culex procax. See Aedes vigilax. Culex quinquefasciatus Filaria ozzardi Wuchereria bancrofti Dirofilaria immitis Dipetalonema reconditum Culex sitiens Wuchereria bancrofti Culex sp. ? Onchocerca caecutiens Culex teniatus. See Aedes aegypti. Culex vigilax. See Aedes vigilax. Culicoides austeni Dipetalonema perstans Culicoides grahami Dipetalonema perstans Eusimulium reptans Cystopsis acipenseris Fannia sp. Larval nema of Johnston & Ban- croft, 1920 Haematopota cordigera Loa loa Hippocentrum trimaculatuin Loa loa Howardina albolineata. See Aedes al- bolineata. Lyperosia exigua Habronema microstoma Mansonia africana Wuchereria bancrofti Mansonia annulipes Wuchereria bancrofti Mansonia fuscopennatus Dipetalonema perstans Mansonia pseudotitillans Wuchereria bancrofti Mansonia sp. Dipetalonema perstans Mansonia uniformis Filaria sp. Castellani & Chalmers, 1913 Wuchereria bancrofti Dipetalonema perstans VOL. 8I Mansonioides annulipes. See Mansonia annulipes. Mansonioides pseudotitillans. See Man- sonia pseudotitillans. Mansonioides wuniformis. sonia uniformis. Musca bezzi Habronema spp. Musca domestica Choanotaenia infundibulum Davainea tetragona Davainea cesticillus : Habronema microstoma Habronema megastoma Habronema muscae See Man- Musca fergusoni Habronema megastoma Habronema muscae Musca humilis Habronema megastoma Habronema muscae Musca lusoria Habronema megastoma Habronema muscae Musca terrae-reginae Habronema megastoma Habronema muscae Agamospirura muscarum Musca ventrosa Habronema me'gastoma Habronema muscae Musca vetustissima Habronema megastoma Habronema muscae Myzomyia superpicta. See Anopheles superpictus. Myzorhynchus pseudopictus. See Ano- pheles hyrcanus pseudopictus. Panoplites africanus. See Mansonia africanus. Panoplites sp. Dipetalonema perstans Pseudopyrellia sp. Habronema megastoma Habronema muscae Sarcophaga melanura Habronema microstoma Sarcophaga misera Habronema muscae P NO. 15 Scutomyia albolineata. See Aedes albo- lineata. Simulids Oncocerca caecutiens Simulium damnosum Oncocerca volvulus Stegomyia fasciata. See Acdes acgypti. Stomoxys calcitrans Hymenolepis carioca Habronema microstoma ? Habronema muscae ? Setaria labiato-papillosa Tabanus circumdatus Agamofilaria tabanicola Taeniorhynchus annulipes. See Man- sonia annulipes. Taeniorhynchus domesticus. Probably Culex pipiens, q. v. Wuchererta bancrofti Tanypus decoloratus Lissorchis fairporti ISOPTERA Hodotermes pretoriensis. See Macro- hodotermes mossambicus trans- vaalensis. Macrohodotermes mossambicus trans- vaalensis Hartertia gallinarum LEPIDOPTERA Aglossa dimidiata Hymenolepis diminuta Aphornia gularis Hymenolepis diminuta Asopia farinalis Hymenolepis diminuta Nymphula nymphaecata Fluke Paralipsa gularis. See Aphornia gularis. Pyralis farinalis. See Asopia farinalts. Tinea granella Hymenolepis diminuta ARTHROPOD HOSTS OF WELMINTHS—HALL 59 MALLOPHAGA “Bird louse ” ? Filaria cypseli Trichodectes latus Dipylidium caninum NEUROPTERA Mystacides nigra Distomum mystacidis Sialis lutaria. See Sialis favilatera. Sialis flavilatera Distomum notidobiae Distomum sialidis Neoechinorhynchus rutili Sialis niger Neoechinorhynchus rutili ODONATA Aeschna sp. Prosotocus confusus Agrion puella. See Coenagrion puella. Agrion spp. Gorgodera pagenstecheri Gorgodera varsoviensis Pleurogenes medians Distomum sp. of Villot Procercoid of Galli-Valerio, 1923 Camallanus lacustris Agrion virgo Pneumonoeces variegatus Pneumonoeces similigenus Halipegus ovocaudatus Calopteryx virgo. See Agrion virgo. Coenagrion puella Tatria acanthorhyncha. Cordulia sp. Prosotocus confusus “ Dragonfly ” Plagiorchis ameiurensis Cercaria prima Epitheca sp. Gorgodera pagenstecheri Gorgodera varsoviensis Gorgodera cygnoides Libellula quadrimaculata Prosthogonimus intercalandus Prosthogonimus pellucidus Tetragoneuria sp. Prosthogonimus sp. of Kotlan and Chandler 60 SMITHSONIAN MISCELLANEOUS COLLECTIONS PLECOPTERA Perla bicaudata Opisthioglyphe endolobum “ Perlid larva” Plagiorchis maculosus Lecithodendrium lagena ORTHOPTERA Blatta orientalis Spirura gastrophila ? Spirocerca sanguinolenta Gonaylonema neoplasticum Gongylonema sp. Moniliformis moniliformis Blattella germanica Protospirura columbiana Gongylonema neoplasticum Gongylonema scutatum Gongylonema pulchrum Periplaneta americana Gongylonema neoplasticum Gongylonema orientale Gongylonema sp. Moniliformis moniliformis Periplaneta australasiae Gongylonema orientale Pycnoscelus surinamensis Oxyspirura mansoni Oxyspirura parvovum SIPHONAPTERA Ceratophyllus fasciatus Hymenole pis diminuta ? Hymenolepis nana Hymenolepis microstoma Agamonema sp. Johnston, 1913 Ctenocephalus canis Dipylidium caninum Hymenolepis diminuta Dirofilaria imimitis Dipetalonema reconditum Ctenocephalus felis Dipylidium caninum Dirofilaria immitis Dipetalonema reconditum Leptopsylla muscult Hymenolepis diminuta VOL. 81 Mesopsylla eucta Cysticercoid of Dampf, 1910 Pulex irritans Dipylidium caninum Hymenolepis diminuta Dipetalonema reconditum Dipetalonema perstans Nenopsylla cheopis Hymenolepis diminuta ? Hymenolepis nana ? Protospirura muris Agamonema sp. Johnston, 1913 TRICHOPTERA Anabolia nervosa Allocreadium isoporum Opisthioglyphe endolobum Chaetopteryx villosa Allocreadium isoporum Drusus trifidus Plagiorchis maculosus Limnophilus flavicornis Opisthioglyphe endolobum Limnophilus griseus Opisthioglyphe endolobum Limnophilus lunatus Opisthioglyphe endolobum Limnophilus rhombicus Opisthioglyphe endolobum Distomum limnophili Notidobia ciliaris Distomum notidobiae Phryganea grandis Opisthioglyphe endolobum Lecithodendrium cheilostomum Brachycoelium retusum Distomum phryganeae Phryganea sp. Lecithodendrium cheilostomum Rhyacophila nubila Fluke UNPLACED “Amphibious insects ” Plagioporus sp. Eumegacetes sp. “ Raubinsekten ” Gorgodera vitelliloba NO. I5 ARTHROPOD HOSTS OF HELMINTHS—HALL 61 ARACH NIDA ACARINA Argas sp. Dipetalonema perstans Ixodes ricinus ? Filaria martis Ornithodorus moubata Dipetalonema perstans Rhipicephalus sanguineus Dipetalonema reconditum Dipetalonema grassii Dirofilaria immitis Rhipicephalus siculus ? Dipetalonema reconditum oil Clcew ? Filaria mitchelli MYRIAPODA Fontaria virginiensis Hymenolepis diminuta Glomeris limbata Cestode larva Julus guttulatus Nematode larva Julus sp. Hymenolepis diminuta CRUSTACEA AMPHIPODA Allorchestes sp. ? Hedruris orestiae Gammarus locusta Distomum gammari Rentsch Polymorphus boschadis Gammarus ornatus. See Gammarus locusta. Gammarus pulex Opisthioglyphe endolobum Distomum agamos Distomum gammari Linstow Distomum pulicis Hymenole pis collaris Hymenolepis tenuirostris Aploparaksis dujardini Echinocotyle mrazeki Cysticercoides sp. Mrazek, 1806 Cysticercus bifurcus Cysticercus hamanni Cysticercus taeniae-pachyacanthae Cysticercus sp. Luehe, 1910 Cysticercus sp. Mrazek, 1890 Taenia sp. Daday, 1900; 168 Tetrameres fissispina Polymorphus boschadis Pomphorhynchus laevis Ayallela azteca Echinorhynchus thecatus Proteocephalus ambloplitis Hyallela knickerbockeri. See Hyallela azteca, Pontoporeia hoyi Echinorhynchus ranae Themisto libellula Sinistroporus simplex BRA NCHIOPODA Apus sp. Agamodistomum apodis CLADOCERA Bythotrephes longimanus Proteocephalus agonis Daphnia pulex Echinuria uncinata Tetrameres fissispina Leptodora kindtu Proteocephalus agonis COPEPODA Acartia clausa Hemiurus appendiculatus Acartia sp. Derogenes varius Boeckella braziliensis. See Pseudo- boeckella braziliensis. Cyclops agilis. See Cyclops serrulatus. Cyclops albidus Proteocephalus ambloplhitts ms 7 = we Ca ee 62 SMITHSONIAN MISCELLANEOUS COLLECTIONS Cyclops bicuspidatus Hymenolepis tenuirostris Drepanidotaenia lanceolata Schistocephalus solidus Bothriocephalus cuspidatus Corallobothrium fimbriatum Dracunculus globocephalus Cyclops brevicaudata. See Cyclops stvenuus. Cyclops brevispinosus Bothriocephalus cuspidatus Diphyllobothrium latum Cyclops coronatus. See Cyclops fuscus. Cyclops crassicornis Hymenolepis brachycephala Cyclops fimbriatus. See Platycyclops fimbriatus. Cyclops fuscus Dracunculus medinensis Cyclops leuckartt Proteocephalus ambloplitis Bothriocephalus cuspidatus Diphyllobothrium mansoni Dracunculus medinensis Cyclops lucidulus Hymenolepis collaris Cyclops oithonoides. See Mesocyclops oithonoides. Cyclops prasinus Proteocephalus ambloplitis Corallobothrium giganteum Bothriocephalus cuspidatus Dracunculus medinensis Cyclops pulchellus. See Cyclops bicus- pidatus. Cyclops quadricornis Cucullanus elegans Dracunculus medinensis Cyclops robustus Diphyllobothrium latum Cyclops serratus. See Cyclops bicus- pidatus. Cyclops serrulatus Hymenolepis collaris Hymenole pis tenuirostris Hymenolepis fasciculata Hymenolepis microsoma Proteocephalus torulosus Proteocephalus longicollis Proteocephalus percae VoL. &I Corallobothrium giganteum Corallobothrium fimbriatum Bothriocephalus cuspidatus Schistocephalus solidus Abothrium infundibuliformis Abothrium crassum Cysticercoid of Rossiter, 1803 Cysticercus quadricurvatus Cysticercus grubert Cysticercus sp. Luehe, 1910 Cyclops sp. Fimbriaria fasciolaris Camallanus lacustris Camallanus microcephalus Dracunculus sp. Philometra sanguineum Cyclops strenuus Proteocephalus torulosus Proteocephalus longicollis Proteocephalus percae Ichthyotaenia sp. Fuhrmann, 1926 Hymenolepis setigera Diphyllobothrium latum Abothrium crassum Abothrium infundibuliformis Triaenophorus nodulosus Cysticercus gruberi Dracunculus medinensis Cyclops tenuicornis. Probably Cyclops albidus q. v. Distomum sp. Herrick Cyclops varius. See Cyclops serrula- tus. Cyclops vernalis Hymenolepis anatina Hymenolepis collaris Cyclops viridis Hymenolepis collaris Hymenolepis gracilis Hymenolepis fasciculatus Dracunculus medinensis Diaptomus africanus Plerocercus africanus Diaptomus alluaudi Hymenolepis anatina Dicranotaenia dubia Diaptemus asiaticus Echinocotyle linstowi Echinocotyle polyacantha Taenta zichyi NO. 15 Diaptomus castor Proteocephalus torulosus Diaptomus coeruleus Hymenolepis collaris Hymenolepis gracilis Hymenolepis tenuirostris Hymenolepis fasciculatus Hymenolepis setigera Diaptomus gracilis Diphyllobothrium latum Diaptomus graciloides Diphyllobothrium latum Diaptomus oregonensis Diphyllobothrium latum Diaptomus sp. Cysticercoides sp. Mrazek, 1808 Cercocystis dendrocercus Philometra sanguineum Diaptomus spinosus Hymenolepis anatina Hymenolepis gracilis Drepanidotaenia lanceolata Echinocotyle linstowi Diaptomus vulgaris Fimbriaria fasciolaris Leptocyclops agilis. See Cyclops ser- rulatus. Mesocyclops oithonoides Proteocephalus percae Platycyclops fimbriatus Hymenolepis brachycephala Triaenophorus nodulosus Pseudoboeckella braziliensis Echinocotyle mrageki OSTRACODA Candona candida Hymenolepis coronula Candona neglecta tuberculata Hymenolepis gracilis Candona rostrata Hymenolepis gracilis Cyclocypris globosa Hymenolepis gracilis Hymenolepis coronula Hymenolepis liophallos Hymenolepis venusta Echinocotyle rosseteri Cysticercoides sp. Rossiter, 1890 5 ARTHROPOD HOSTS OF HELMINTHS—HALL 63 Cyclocypris laevis Hymenolepis coronula Cyclocypris ovum Hymenolepis coronula Cypria ophthalmica Hymenolepis anatina., Hymenolepis gracilis Hymenolepis coronula Echinocotyle rosseteri “ Cypris agilis” Hymenolepis venusta Cypris cinerea. See Cyclocypris glo- bosa. Cypris compressa. See Cypria ophthal- mica. Cypris elongata Taenia sp. Daday, 1900 Cypris incongruens. See Heterocypris mcongruens. Cypris ophthalmica. See Cypria oph- thalmica Cypris ovata. See Cypris pubera. Cypris ovum. See Cyclocypris ovum. Cypris pubera Hymenolepis anatina. Cypris virens. See Eucypris virens. Cypris viriens. See Eucypris virens. Dolerocypris fasciata Hymenolepis gracilis Eucandona hungarica Hymenolepis anatina Eucypris crassa _Hymenolepis anatina Eucypris virens Hymenolepis collaris Hymenolepis coronula Hymenolepis gracilis Heterocypris imcongruens Hymenolepts anatina “Ostracod ” Cysticercus sp. Luehe, 1910 DECAPODA Astacus astacus Astacotrema cirrigerum Hymenolepis collaris Hymenolepis tenuirostris Polymorphus boschadis 64 SMITHSONIAN Astacus fluviatilis. cus. Astacus japonicus. japonicus. See Astacus asta- See Cambaroides Astacus leptodactylus Disiomum reinhardii Caimbaroides japonicus Paragonimus westermani Cambaroides similis ? Paragonimus westermani Cambarus propinquus Microphallus opacus Cambarus spp. Crepidostomum cornutum Cerataspis monstrosa Dinurus tornatus “Crabs.” Distomum kalapai “ Crayfish ” Astacotrema cirrigerum Acrolichanus petalosa Plagiorchis ameiurensts Distoma of Cooper, 1883 Distomum of Linton, 1892 Eriocheir japonicus Paragonimus westermani Stephanolecithus parvus Geothelphusa dehaani. See Potamon (Geothelphusa) dehaani. Geothelphusa obtusipes. See Potamon (Geothelphusa) obtusipes. “ Marine decapods ” Rhynchobothrius ruficollis MISCELLANEOUS COLLECTIONS VoL. 81 Potamobius astacus. See Astacus asta- cus. Parathelphusa (Parathelphusa) sinen- SIS Stephanolecithus parvus Potamon dehaant. See Potamon (Geothelphusa) dehaani. Potamon obtusipes. See Potamon (Geothelphusa) obtusipes. Potamon sinensis. See Parathelphusa (Parathelphusa) sinensis. Potamon (Geothelphusa) dehaani Paragonimus westermani Macroorchis spinulosus Stephanolecithus parvus Potamon (Geothelphusa) obtusipes Paragonimus westermant Stephanolecithus parvus Pseudothelphusa iturbei Paragonimus westermant Sesarma dehaanii Paragonimus westermant Stephanolecithus parvus ISOPODA Asellus aquaticus Camallanus elegans Hedruris androphora Acanthocephalus luciu Echinorhynchus ranae Porcellio laevis Dispharynx spiralis GENERAL DISCUSSION On the basis of the foregoing lists, the arthropod hosts are arranged below in their approximate order of relative importance for each order of parasites, with a résumé of the numbers of host and para- site species involved. As intermediate hosts for tapeworms with primary hosts living in water or feeding on arthropods which live in water, the Copepoda are of outstanding importance, the next most important group being the Ostracoda. The Amphipoda, Decapoda, and Cladocera are much less extensively involved as intermediate hosts for tapeworms so far as is known at the present time. NO. 15 ARTHROPOD HOSTS OF HELMINTHS—HALL 65 a Cestoda No. of parasite : No. of host No. of parasite spp. Tee ae eee Intermediate spp. with known hosts known hosts host group involved for adults for adults CRUSTACEA : COME Hoda neccsawe se cisetes 27 20 8 CISC 60 EI a a 15 6 3 mp nt podal ines isc << « I 5 7 Wecapodae cece « > s1<1- 2 3 0 (lad Ocerairce hoc cassSiontes 2 I 0 INSECTA: (ColeOpteray caccsetrciow noms cc 8 9 I Siphonaptera 05.5. .56 50. 7 4 I DiMheraMeea eyes sieve loos Senet 2 4 O IEepicdoptenal yas. oes oe 4 I O Odorataee era ke cs cscsccieis 2 I I Wermeaptena aces ce cies c< ene I 2 oO Mia ophamayenesee oem. «+ src I I oO IMGMRMAPODA) Qs csc ccs ceecsseee- 2 I oO As intermediate hosts for tapeworms of land animals, the Coleop- tera are distinctly the most important group, other insect groups being of much less importance so far as we know. Trematoda No. of parasite spp. reported as No. of host No. of parasite spp. larvae without Intermediate spp. with known hosts known hosts host group involved for adults for adults CRUSTACEA : Wecapodays samciiocaxs oa eis 15 8 4 AtTID A pPOMae re cteceller -e-e10 010 3 I 4 (Camnenock) “sosaceasoomesecs 2 2 I BranehtopoOdal qeyraceee =<) «1s I oO I INSECTA: Dice tcp cers sere: sos teisiae eves II 2 8 Mrichoptetay ... as «82/6 II 2 4 @clomataee sees cote ava s titers 8 6 2 Biphementday sajc6 2. o-sle- 5 5 I Coleoptera, oieiecmieccie's eiayse'6 3 3 I (Wnplaced insects <......... 2 4 0 INetmnopterde «acces os caer 60s 2 oO 8 ITecOntehamaeanes chhs a sete 2 I 0 Wepidopteray Ge sce siie cess + I O I Among the Crustacea, the Decapoda are of outstanding importance as hosts for flukes, some of these flukes occurring in land mammals which eat raw crabs or crayfish. 66 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 81 Among the Insecta, the apparent importance of the Diptera and Trichoptera is not well established. These groups rate high in number of species serving as hosts, but as the flukes reported from them are mostly larval forms of which the adults are not known and which may later prove to represent no more known species than are already known and recorded here from these hosts, or only a few more, these groups cannot be rated at the present time as any more important than the Odonata or Ephemerida as hosts for flukes. Nematoda No. of parasite spp. reported as No. of host No. of parasite spp. larvae without Intermediate spp. with known hosts known hosts host group involved for adults for adults CRUSTACEA: Copepoda ua.nacmccsts smell 8 / 0 Ann plitpodasrstersiem eerie 2 2 oO Iisopoday erie oe nome 2 2 0 Cladoceray iiss. sete «eats 1 I 0 INSECTA: ¢ Diptenay wes sccm eee 67 17 4 Coleoptera ce eeeecee 39 10 3 Onthopteragetst acca eee 5 5 0 Siphonapteray sani. cnicem ces 5 4 I Ephemeridas 222). 2 sere ets 2 0 I Anoplenunay yee wicncen nee I I 0 Tisoptetay Waly. taceticltenudene I I 0 Mallophagam jase eee: I I O Odonata so ee I I oO HARA GIETENIDIAUS oer toga Pens se rei oie 6 6 0 Among the Crustacea, the Copepoda are the important group as carriers of parasitic nematodes. Among the Insecta the Diptera are of striking importance, no less than 67 of the Diptera being reported as carriers for a total of 17 nematode species, this fact being the result largely of the role of the mosquitoes as carriers of filarids. The Coleoptera take first rank as carriers of spirurids. Of lesser impor- tance are the Orthoptera and Siphonaptera, and the other groups of insects show but few host species and these accused of carrying but one nematode parasite. The Arachnida as a whole have been accused of carrying 6 nematodes, and but 6 arachnids are incriminated. The arachnids have not been reported as carriers of any parasitic worms other than nematodes. NO. 15 ARTHROPOD HOSTS OF HELMINTHS—HALL 67 Acanthocephala No. of parasite ; No. of host No. of parasite spp. eee rae Intermediate spp. with known hosts known hosts host group involved for adults for adults CRUSTACEA: ENT POCA eee sels eee A 4 0 SOIC ersten oles Se snus es I O MeeapOdate sis iets ones ss I I O INSECTA: GGlEOpieRal Piss ace coe seas II 3 INetnOptera i. .isec ssn. ssn s os 2 I O Oxthoptenaw sess 6s soe ck 2 I 0 Among the Crustacea, the Amphipoda are of special significance as hosts for acanthocephalids of aquatic animals, so far as the life his- tories of such worms are known. The only other crustaceans involved are Isopoda and Decapoda. Among the Insecta, the Coleoptera are of major importance as carriers of acanthocephalids with known life histories. The only other insects involved are Neuroptera and Orthoptera. If we take the outstanding groups of intermediate hosts for each order of parasites, we have the following: For Cestoda: Copepoda and Ostracoda; Coleoptera. For Trematoda: Decapoda; Diptera, Trichoptera, Odonata, and Ephemerida. For Nematoda: Copepoda; Diptera and Coleoptera. For Acanthocephala: Amphipoda ; Coleoptera. Among the insects, the importance of the Coleoptera is indicated by the fact that this group is of decided significance for Cestoda, Nematoda, and Acanthocephala. The Diptera are important as car- riers of Trematoda and Nematoda. The Trichoptera, Odonata, and Ephemerida only figure as outstandingly important for Trematoda. Among the crustaceans, the Copepoda are the major group as hosts for both Cestoda and Nematoda. The Ostracoda are only known to be important as hosts for Cestoda, the Decapoda as hosts for Trema- toda, and the Amphipoda as hosts for Acanthocephala. The following table is inserted to give a rapid check on the known occurrence in the different arthropod hosts of parasites of the groups involved in this paper. If an arthropod group is known to contain intermediate hosts for the worm groups involved, an X is placed under the worm group and opposite the host group. If there are no such hosts known, an O is placed in the corresponding position. Wh ha 68 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. ot LIST SHOWING RECORDS (X) OR EACK OF RECORDS (@)9OE PARASITE GROUPS IN HOST GROUERS Arthropod group Cestoda Trematoda Nematoda Acanthocephala Atm phipoday semn cise cece xX xX xX xX Branchtopoday. ae aaniosseceee: O xX O O Cladocera secs viene xX O xX O Copepodawee ane eee eee ox xX x O Mecapodaweenncadec ooo ecrine X xX O xX Tsopodavee: mreachiac caneceieneiers O O xX xX Ostracodaiwac. scr seen eee oe x O O O ANOpleura .. cuacteristnes cert eee O O xX O Coleoptera nceererer cree xX xX xX xX Wermapterameremecacee ete oe xX O O O Dipteray ces seen Ptcevetuerts xe xX xX O Eiphemernicdairacnce dectncsom aria: O xX X O SOPtera eehiie es wae ee eee O O aK O icenidopteta, vee cemen a eee x De O O Mallophagapreer emia cence sete xX O X O INetiroptera, <. 9. ss tenis ton Bene cave O xX O X Odonatareeserst esses eee eee xX xX xX O @xthopteras eee eee ieee eee O O xX xX Plecopterar aacnoacncer ene oe oe O xX O O Siphonapteraanse ccs onece cei xX O xX O Mrichoptena mime sacereseleios O X O O Insecta munplacedamaestecentsnince O xX O O Miyriapodais 2 tcc casei seer eee xX O O O NACH NiGaamacecnt teh ects O O xX O It is of interest to note that of the 24 arthropod groups listed above, the number of groups used as hosts by cestodes, trematodes, and nema- todes is the same or almost the same, namely, 13 by cestodes and trematodes and 14 by nematodes ; only 6 are used by acanthocephalids. From the foregoing something may be indicated as to the range of parasites on the part of the various intermediate host groups, as fol- lows: Hosts for 4 worm groups: Amphipoda and Coleoptera. Hosts for Cestoda, Trematoda and Nematoda: Copepoda; Diptera and Odonata. Hosts for Cestoda, Trematoda and Acanthocephala: Decapoda. Hosts for Cestoda and Trematoda: Lepidoptera. Hosts for Cestoda and Nematoda: Cladocera; Mallophaga and Siphonaptera. Hosts for Trematoda and Nematoda: Ephemerida. Hosts for Trematoda and Acanthocephala: Neuroptera. Hosts for Nematoda and Acanthocephala: Isopoda; Orthoptera. Hosts for Cestoda only: Ostracoda; Dermaptera; Myriapoda. Hosts for Trematoda only: Branchiopoda; Plecoptera, Trichoptera, and un- placed insects. Hosts for Nematoda only: Anopleura and Isoptera; Arachnida. No group is yet reported as a host group for Acanthocephala only. NO. 15 ARTHROPOD HOSTS OF HELMINTHS—HALL 69 Taking the major host groups, the Crustacea, Insecta, Myriapoda, and Arachnida, as a whole and the four worm groups as a whole, we may make the following summary: There are 48 species in the Cestoda which have arthropods as intermediate hosts and for which we know the primary hosts; there are 22 larval forms in addition for which the primary hosts are not known. There are 37 species in the Trematoda which have arthropods as intermediate hosts and for which we know the primary hosts ; there are 27 larval forms in addition for which the primary hosts are not known. There are 49 species in the Nematoda which have arthropods as intermediate hosts and for which we know the primary hosts; there are 12 larval forms in addition for which the primary hosts are un- known. There are 9 species in the Acanthocephala which have arthropods as intermediate hosts and for which we know the primary hosts. There are altogether 143 species of worms parasitic in vertebrates which have arthropods as intermediate hosts and for which the pri- mary hosts are known; there are 61 larval forms in addition for which the primary host is unknown. In the Crustacea there are 49 species which serve as intermediate hosts for Cestoda, 22 for Trematoda, 12 for Nematoda, and none for Acanthocephala. In the Insecta there are 25° species which serve as intermediate hosts for Cestoda, 46 for Trematoda, 122 for Nematoda, and 15 for Acanthocephala. In the Myriapoda there are 2 species which serve as intermediate hosts for Cestoda, and none for Trematoda, Nematoda, or Acantho- cephala so far as we know at present. In the Arachnida there are 6 species which serve as intermediate hosts for Nematoda, and none for Cestoda, Trematoda, or Acantho- cephala so far as we know at present. , The Insecta are far in the lead as regards number of species known to serve as intermediate hosts for parasitic worms, as there are 186 species of insects, 77 species of crustaceans, 6 species of arachnids, and only 2 species of myriapods included in these lists of intermediate hosts. The total number of arthropod species listed here as inter- mediate hosts for the worm groups involved is 271. CONCLUSION It should be reiterated that one must not draw too sweeping con- clusions in regard to the importance of host groups or in regard to 79 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 81 several other things at this time. For one thing, the lists given here are such as could be compiled in the time at the writer’s disposal and while reasonably comprehensive must necessarily be incomplete. For another thing, our total knowledge in regard to the life histories of heteroxenous helminths is very slight. As already stated, we know the life histories of approximately I per cent of the known tapeworms, and this status is sufficiently representative of conditions for all heter- oxenous worm groups to need no detailed statement in regard to the other groups. There may be important intermediate host groups of which no member has yet been incriminated. We know about 143 life histories involving arthropods ; there are certainly hundreds, per- haps thousands, of such life histories still to be ascertained. Admitting all of these defects in our data, we are nevertheless justified in saying that the lists presented here will be of value in affording the student a clue as to the probabilities in beginning a search for the intermediate host of a heteroxenous worm parasite, or in considering the probable identity of a larval worm found in an arthropod. This will fulfill one of the purposes of this paper—to aid the student. The young students of to-day will include among their ranks the competent scientists of to-morrow. Another purpose of this paper is to point out the opportunities for cooperation among scientists in adding to our knowledge of the life histories of parasitic worms. Zwaluwenberg, an entomologist, has said recently: “The interrelationships of insects and nematodes is a subject of which most entomologists seem to have little adequate conception.” Some months ago, in discussing the scope of this paper with Dr. L. O. Howard, the writer told him that he expected to call attention to the fact that our knowledge of these life histories had come almost entirely from the parasitologists, and that the workers on insects and crustaceans had aided very little in the process. Dr. Howard, characteristically, suggested that this be done very diplo- matically. It is the writer’s intention to do this diplomatically. It is primarily the business of the parasitologist to ascertain the life his- tories of the parasites with which he deals. It would not be in order to ask the specialist on insects or crustaceans to ascertain the life histories of the larval worms which he finds in these insects and crustaceans, nor would it add greatly to our knowledge to have per- sons unfamiliar with parasitic worms publish findings in an unfamiliar field. Nevertheless, there is an opportunity for cooperation between the workers on parasitic helminths and the workers on their arthropod hosts, and little advantage has been taken of this fact in the past. NOP 15 ARTHROPOD HOSTS OF HELMINTHS—HALL 71 My friend, Dr. Wm. A. Riley, has called my attention to Stein’s pioneer contributions ; conceding the point, Stein’s good example has not been followed by most entomologists. The competent worker in either field should be primarily a zoologist, and as such able to see the possibilities for cooperation with other zoologists in connection _with incidental findings which come to his attention. The larval helminth in an arthropod is an animal which concerns the helmin- thologist in one direction and the “ arthropodologist ” in another. A sound consideration of the worm calls for a sound consideration of its host, and vice versa. Prophylactic measures directed against het- eroxenous worms call for control measures for intermediate hosts, and if this host is an arthropod the helminthologist must draw on the knowledge of the man who knows about arthropods. One of the promising developments in this connection is the fact that whereas the entomologist in the past has devoted his attention to the outside of the insects with only casual attention to the internal anatomy, there is now a tendency to devote more attention to the internal structures. In examining the interior of the insects, the entomologist is certain to find larval worms in some of them. In such cases he would be rendering a service if he would do one of the fol- lowing things: If the entomologist is well trained in zoology, and has the time, facilities, and inclination to carry out an adequate investigation of these worms, he can proceed with feeding experiments and ascertain the life history. Lacking the training, time, facilities, or inclination to to do such work, he can turn the material over to a parasitologist for investigation, or call attention in his publications to his findings in order that they may serve as a guide to the parasitologist who is working along this line. Some of the hosts given in this paper are not well established, but are included for completeness. In establishing a life history for a parasitic worm, one may be guided with profit by the remarks of Stiles in 1896 in connection with the life histories of bird tapeworms: The known or supposed life history has been based upon four different methods of work, i. e.: 1. Experimental infection of the fowls by feeding to them known larval stages found in invertebrates, and thus raising the adult stage. 2. Experimental infection of invertebrates by feeding to them the eggs of tapeworms found in birds, and thus raising the larval stage. 3. Comparison of the hooks upon the heads of adult tapeworms of birds with the hooks of larvae found in invertebrates, and thus associating the young and the old stages. 4. Wild speculations as to the intermediate hosts, based upon negative results and totally devoid of any scientific foundation. 72 SMITHSONIAN MISCELLANEOUS COLLECTIONS voL. 81 Of these four methods of work the first two give positive proof of the life history when the experiments are successful; the third gives a probability to the statements, but not a proof; the less said about the fourth method the better. In this later epoch it is advisable to establish a life history by both of the first methods, not ignoring the third, if adequate evidence is desired. Raising Diphyllobothrium latum in dogs by feeding plerocer- coids from fish did not show that a copepod was the first host ; failure to infect fish with the tapeworm eggs necessitated further search and so led to the discovery of the copepod host. Another thing deserves emphasis: Finding that one arthropod is an intermediate host does not settle the problem of a life history of a worm. The worm may have a score of intermediate hosts, and the most important one may not be an arthropod. ADDENDUM Since the foregoing was written the following records have come to hand and are given here without discussion: EIST BY PARASELES (Crust. = Crustacea) CESTODA CESTODARIA Amphilina foliacea—Corophium curvispinum; Crust.; Amphipoda Dikerogammarus haemobaphes; Crust.; Amphipoda Gammarus platycheir; Crust.; Amphipoda Metamysis strauchi; Crust.; Mysidacea DIPH YLLOBOTHRIIDAE Diphyllobothrium ranarum—Cyclops fuscus; Crust.; Copepoda Diphyllobothrium decipiens—Cyclops sp.; Crust.; Copepoda Diphyllobothrium erinacei—Cyclops sp.; Crust.; Copepoda Diphyllobothrium mansoni—Cyclops sp.; Crust.; Copepoda Cyclops strenuus; Crust; Copepoda PROTEOCEPH ALIDAE Proteocephalus ambloplitis—H yalella azteca; Crust.; Amphipoda Cyclops serrulatus; Crust.; Copepoda Cyclops viridis; Crust.; Copepoda Proteocephalus pinguis—Cyclops serrulatus; Crust.; Copepoda Cyclops viridis; .Crust.; Copepoda Ophiotaenia testudo—Cyclops sp.; Crust.; Copepoda NO. I5 ARTHROPOD HOSTS OF HELMINTHS—HALL es CESTODA (Continued ) HY MENOLEPIDIDAE Hymenolepis collaris—Cypris sp., Crust.; Ostracoda Hymenolepis anatina—Cypris sp.; Crust.; Ostracoda Hymenolepis coronula—Cypris sp.; Crust.; Ostracoda Hymenolepis carioca—Choeridium histeroides; Insecta; Coleoptera Hister (Carcinops) 14-striatus; Insecta; Coleoptera Anisotarsus agilis; Insecta; Coleoptera ? Choanotaenia infundibulum—Cratacanthus dubius; Insecta ; Coleoptera * DAVAINIIDAE Raillietina cesticillus—Anisotarsus agilis; Insecta; Coleoptera Anisotarsus terminatus; Insecta; Coleoptera Choeridium histeroides; Insecta; Coleoptera Aphodius granarius; Insecta; Coleoptera * Selenophorus ovalis; Insecta; Coleoptera * Selenophorus pedicularis; Insecta; Coleoptera Triplectrus rusticus; Insecta; Coleoptera * TREMATODA PLAGIORCHIIDAE Plagiorchis maculosus—Chironomus plumosus; Insecta; Diptera Chironomus sp.; Insecta; Diptera HETEROPH YIDAE Microphallus minus—Macrobrachium nipponensis; Crust.; Decapoda TROGLOTREMATIDAE Paragonimus westermani—Cambaroides dauuricus; Crust.; Decapoda Eriocheir sinensis; Crust.; Decapoda FAMILY UNCERTAIN Distome of Eckstein—Culex pipiens; Insecta; Diptera Metacercaria of Joyeux, 1928—Anopheles maculipennis; Insecta; Diptera Cercaria X.1 of Harper, 1929—Gammarus pulex; Crust.; Amphipoda Orchestia littorea; Crust.; Amphipoda - Chironomus plumosus; Insecta; Diptera Culex pipiens; Insecta; Diptera Tipula maxima; Insecta; Diptera Pedicia rivosa; Insecta; Diptera Dysticus marginalis; Insecta; Coleoptera Sialis Iutarius; Insecta; Diptera Halesus sp.; Insecta; Trichoptera Limnophilus centralis; Insecta; Trichoptera Limnophilus rhombicus; Insecta; Trichoptera Plectrocnemia conspersa; Insecta; Trichoptera Rhyacophila dorsalis; Insecta; Trichoptera * Unpublished work by M. F. Jones. 74 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 81 NEMATODA SPIRURIDAE Gongylonema ingluvicola—Copris minutus; Insecta; Coleoptera * Phanaeus carnifex—tInsecta; Coleoptera * Physocephalus sexalatus—Canthon laevis; Insecta; Coleoptera Gymnopleurus sinnatus; Insecta; Coleoptera Phanaeus carnifex; Insecta; Coleoptera Larval spirurid (?)—Campodea sp.; Insecta; Thysanura PHYSALOPTERIDAE Proleptus scillicola—Carcinus maenas; Crust.; Decapoda Eupagurus bernhardus; Crust.; Decapoda ACUARIIDAE Cheilospirura hamulosa—Melanoplus femurrubrum; Insecta; Orthoptera * Cheilospirura spinosa—Melanoplus femurrubrum; Insecta; Orthoptera Acuaria anthuris—Melanoplus femurrubrum; Insecta; Orthoptera * Crickets; Insecta; Orthoptera * TETRA MERIDAE Tetrameres americana—Melanoplus differentialis; Insecta; Orthoptera Melanoplus femurrubrum,; Insecta; Orthoptera * FILARIIDAE Wuchereria bancrofti—Aedes albolateralis; Insecta; Diptera Aedes chemulpoensis; Insecta; Diptera Aedes galloisi; Insecta; Diptera Aedes subpictus; Insecta; Diptera Armigeres obturbans; Insecta; Diptera Culex annulus; Insecta; Diptera Culex bitaeniorhynchus karatsuensis; Insecta; Diptera Culex japonicus; Insecta; Diptera Culex pipiens pallens; Insecta; Diptera Culex tipuliformis; Insecta; Diptera Culex tripunctatus; Insecta; Diptera Culex tritaeniorhynchus; Insecta ; Diptera Culex whitmorei; Insecta; Diptera ; DRACU NCULIDAE Philometra nodulosa—Cyclops brevispinosus; Crust.; Copepoda FAMILY UNCERTAIN Cystopsis accipenseris—Dikerogammarus haemobaphes,; Crust.; Amphipoda Gammarus platycheir; Crust.; Amphipoda Cyclopsinema mordens—Pachycyclops signatus; Crust.; Copepoda * Unpublished work of E. B. Cram. 7 by iS , ©. a a ee NO. 15 ARTHROPOD HOSTS OF HELMINTHS—HALL 75 EIST BY HOSTS (Cest. = Cestoda; Trem. = Trematoda; Nem. = Nematoda) CRUSTACEA AMPHIPODA Corophium curvispinum—Amphilina foliacea; Cest.; Cestodaria Dikerogammarus haemobaphes—Amphilina foliacea; Cest.; Cestodaria Cystopsis accipenseris; Nem.; Family ? Gammarus platycheir—Amphilina foliacea; Cest.; Cestodaria. Cystopsis accipenseris; Nem.; Family? Hyalella azteca—Proteocephalus ambloplitis; Cest.; Proteocephalidae COPEPODA Cyclops brevispinosus—Philometra nodulosa; Nem.; Dracunculidae Cyclops fuscus—Diphyllobothrium ranarum; Cest.; Diphyllobothriidae Cyclops serrulatus—Proteocephalus pinguis; Cest.; Proteocephalidae Proteocephalus ambloplitis; Cest.; Proteocephalidae Cyclops sp.—Diphyllobothrium decipiens; Cest.; Diphyllobothriidae Diphyllobothrium erinacei; Cest.; Diphyllobothriidae Ophiotaenia testudo; Cest.; Proteocephalidae Cyclops strenuus—Diphyllobothrium mansoni; Cest.; Diphyllobothriidae Cyclops viridis—Proteocephalus pinguis; Cest.; Proteocephalidae Proteocephalus ambloplitis; Cest.; Proteocephalidae Macrocyclops signatus—Cyclopsinema mordens; Nem.; Family? Pachyclops signatus—See Macrocyclops signatus DECAPODA Cambaroides dauuricus—Paragonimus westermani; Trem.; Troglotrematidae Carcinides (Carcinus) maenas—Proleptus scillicola; Nem.; Physalopteridae Carcinus maenas—See Carcinides (Carcinus) maenas Eriocheir sinensis—Paragonimus westermani; Trem.; Troglotrematidae Eupagurus bernhardus—See Pagurus bernhardus. Macrobrachium nipponensis—Microphallus minus; Trem.; Heterophyidae Pagurus bernhardus—Proleptus scillicola; Nem.; Physalopteridae MYSIDACEA Metamysis strauchi—Amphilina foliacea; Cest.; Cestodaria OSTRACODA Cypris sp.—Hymenolepis anatina; Cest.; Hymenolepididae Hymenolepis collaris; Cest.; Hymenolepididae Hymenolepis coronula; Cest.; Hymenolepididae 76 SMITHSONIAN MISCELLANEOUS COLLECTIONS — VOL. 8I INSECTA COLEOPTERA Anisotarsus agilis—Raillictina cesticillus; Cest.; Davainiidae Hymenolepis carioca; Cest.; Hymenolepididae Anisotarsus terminatus—Raillietina cesticillus; Cest.; Davainiidae Aphodius granarius—Raillietina cesticillus; Cest.; Davaintidae Canthon laevis—Physocephalus sexalatus; Nem.; Spiruridae Choeridium histeroides—Raillietina cesticillus; Cest.; Davainiidae Hymenolepis carioca; Cest.; Hymenolepididae Copris minutus—Gongylonema ingluvicola; Nem.; Spiruridae Cratacanthus dubius—? Choanotaenia infundibulum; Cest.; Hymenolepididae Dysticus marginalis—Cercaria X.1 of Harper, 1929; Trem.; Family? Gymnopleurus sinuatus—See Gymnopleurus sinuatus Gymnopleurus sinuatus—Spirocerca sanguinolenta; Nem.; Spiruridae Hister (Carcinops) 14-striatus—Hymenolepis carioca; Cest.; Hymenolepididae Phanaeus carnifex—See Phanaeus vindexr Phanaeus vindex—Gongylonema ingluvicola; Nem.; Spiruridae Physocephalus sexalatus; Nem.; Spiruridae Selenophorus ovalis—Raillietina cesticillus ; Cest.; Davainiidae Selenophorus pedicularis—Raillietina cesticillus; Cest.; Davainiidae Triplectrus rusticus—Raillictina cesticillus; Cest.; Davainiidae NEUROPTERA Sialis lutarius—See Sialis flavilatera Sialis flavilatera—Cercaria X.1 of Harper, 1929; Trem.; Family? TRICHOPTERA Halesus sp.—Cercaria X.1 of Harper, 1929; Trem.; Family? Limnophilus centralis—Cercaria X.1 of Harper, 1929; Trem.; Family? Limnophilus rhombicus—Cercaria X.1 of Harper, 1929; Trem.; Family? Plectrocnemia conspersa—Cercaria X.1 of Harper, 1929; Trem.; Family? Rhyacophila dorsalis—Cercaria X.1 of Harper, 1929; Trem.; Family? DIPTERA Aedes albolateralis—W uchereria bancrofti; Nem.; Filariidae Aedes chemulpoensis—W uchereria bancrofti; Nem.; Filariidae Aedes galloisi—W uchereria bancrofti—Nem. ; Filariidae Anopheles maculipennis—Metacercaria of Joyeux, 1928; Trem.; Family? Anopheles rossi—See Anopheles subpictus Anopheles subpictus—Wuchereria bancrofti; Nem.; Filariidae Armigeres obturbans—See Desvoidya obturbans Chironomus plumosus—Plagiorchis maculosus; Trem.; Plagiorchiidae Cercaria X.1 of Harper, 1929; Trem.; Family? Chironomus sp.—Plagiorchis maculosus; Trem.; Plagiorchiidae Culex annulus—See Culex tritaeniorhynchus NO. 15 ARTHROPOD HOSTS OF HELMINTHS—HALL Tf INSECTA (Continued) DIPTERA (Continued) Culex bitaeniorhynchus karatsuensis—Wuchereria bancrofti; Nem.; Filariidae Culex japonicus—Wuchereria bancrofti; Nem.; Filariidae Culex pipiens—Distome of Eckstein; Trem.; Family? Cercaria X.1 of Harper, 1929; Trem.; Family? Wuchereria bancrofti; Nem.; Filariidae Culex pipiens pallens—See Culex pipiens Culex tipuliformis—Wuchereria bancrofti; Nem.; Filariidae Culex tripunctatus—Wuchereria bancrofti; Nem.; Filariidae Culex tritaeniorhynchus—W uchereria bancrofti; Nem.; Filariidae Culex whitmorei—W uchereria bancrofti; Nem.; Filariidae Desvoidya obturbans—W uchereria bancrofti; Nem.; Filariidae Tipula maxima—Cercaria X.1 of Harper, 1929; Trem.; Family? Pedicia rivosa—Cercaria X.1 of Harper, 1929; Trem.; Family? THYSANURA Campodea sp.—Larval spirurid (?); Nem.; Spiruridae? ORTHOPTERA Cricket—Acuaria anthuris; Nem.; Acuariidae Melanoplus differentialis—Tetrameres americana; Nem.; Tetrameridae Melanoplus femurrubrum—Tetrameres americana; Nem.; Tetrameridae Cheilospirura hamulosa; Nem.: Acuartidae Cheilospirura spinosa; Nem.; Acuariidae Acuaria anthuris; Nem.; Acuariidae