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In the course of study of the fossil rodent collections of the Carnegie Museum, the writer, through the kindness of Mr. Charles Gilmore of the United States National Museum, and of Dr. Walter Granger, of the American Museum of Natural History, has been privileged to examine the extensive rodent collections preserved in the latter two institutions, and to describe the material which is the subject of the present paper. The illustrations for this article are from drawings by Mr. Sydney Prentice. Order SIMPLICIDENTATA Lilljeborg Family ISCHYROMYIDT: Alston Genus Pseudocylindrodon gen. nov. Genotype: Pseudocylindrodon neglectus sp. nov. 1903. Cylindrodon fontis Matthew, Bull. Amer. Mus. Nat. Hist., Vol. XIX, Art. VI, fig. 8A (p. 212). Holotype: Left ramus of mandible, U. S. N. M. 13758. Paratype: Left ramus of mandible, U. S. N. M. 13757. Referred specimens: Left ramus of mandible with DP4 Mi_2, A. M. N. H. 9644; right ramus of mandible with P4 and Mi, U. S. N. M. 13759; part of left ramus of mandible with P4, A. M. N. H. 9646. Horizon: Pipestone Springs Oligocene. Locality: Pipestone Springs, Jefferson County, Montana. Diagnosis: Cheek teeth with lower crowns than those of Cylindrodon, and with shallower basins, the floors of which are visible in crown view. P4 with trigonid more elevated than in Cylindrodon and trigonid cusps less widely separated, protolophid absent or merely indicated, ex- ternal intermediate cuspule present but hypolophid absent. Inferior molars with central basins broader, anterior and posterior basins narrower, antero-posteriorly, than in Cylindrodon', internal inter- mediate cuspules prominent, their flanks fused with entoconids and 1 Mar 25 mo Issued October 21, 1935. 2 Annals of the Carnegie Museum VOL. XXV metaconids, blocking outlets of central basins; posterior basins converted into fossettes, their outlets dammed due to complete fusion of entoconids and hypolophulid crests. Talonids of molars, except in M3, wider, relative to trigonids, than in Cylindrodon\ hypoconids showing some hypertrophy, consisting of strong rounded buttresses, jutting externally (except in M3) and not attenuate as in Cylindrodon. DP4 relatively smaller than in Cylindrodon. Mental foramina two, placed posterior to diastema. The present species has been confused with Cylindrodon fontis Douglass, from which, as the diagnosis indicates, it is quite distinct. An occlusal view of the dentition of A. M. N. H. 9644, was figured by Matthew^ as an example of an early stage of tooth wear in Cylin- drodon fontis Douglass. In Matthew’s figure (fig. 8A) however, Mi_2 differ from the molars of Cylindrodon fontis Douglass, illustrated in the same drawing, in showing robust, rounded hypoconid buttresses, broad central valleys and hypolophulid crests fused with entoconids — characteristic features of P send 0 cylindrodon neglectus m. My figures of the holotype mandibular ramus (figure i of the present paper) bring out more clearly the details of the pattern, which may be com- pared with that of a specimen of Cylindrodon fontis Douglass, A. M. N. H. 9638, shown in figure 2. The discrepancy in crown height Fig. I. Pseudocylindrodon neglectus Burke, holotype, U. S. N. M. 13758. External view of mandibular ramus and occlusal view of cheek teeth, x 5. ^Matthew, W. D., “The Fauna of the Titanotherium Beds of Montana.” Bull. Amer. Mus. Nat. Hist., Vol. XIX, Art. VI, fig. 8a (p. 212), 1903. 1935 Burke: Pseudocylindrodon, A New Rodent Genus 3 Fig. 2. Cylindrodon fontis Douglass, referred specimen, A. M. N. H. 9638. Ex- ternal view of mandibular ramus and occlusal view of cheek teeth, x 5. between Pseudocylindrodon neglectus m. and Cylindrodon fontis Douglass is quite evident in these figures. In some respects, Pseudocylindrodon is intermediate between Cylindrodon and Ardynomys. The cheek teeth are nearer those of Ardynomys in crown height and their tendency toward hypertrophy of the hypoconid region; the tooth pattern in general shows similari- ties. The incisor of P seudo cylindrodon is not flattened, however, as it is in Ardynomys; the central basins of the cheek teeth are deeper in P seudocylindrodon, have rounded floors, and are closed internally by the upgrown intermediate cuspule. The jaw of Ardynomys is deeper anteriorly than that of P seudo cylmdrodon, the posterior basins of the cheek teeth of Ardynomys have internal exits, and the pro- tolophids are less transverse than in Pseudocylindrodon. Cylindrodon, Pseudocylindrodon, and Ardynomys, are probably Oligocene representatives of the stock of rodents typified in the Bridger Eocene by the forms which TroxelE has included under Tillomys. In North America this line appears to culminate in the lower Oligocene, at which time it is also represented by Ardyyiomys in Asia. Whether or not Tsaganomys and Cyclomylus of the Upper Oligocene of Mongolia are also derivatives of this stock is not determinable without further ^Troxell, Edward L., “The Eocene Rodents Sciuravus and Tillomys.” Amer. Jour. Sci., Vol. 5, pp. 387-396, 1923. 4 Annals of the Carnegie Museum VOL. XXV study, although there are some interesting points of resemblance be- tween the latter genera and Ardynomys. M E ASUREMENTS Holotype Paratype U. S. N. M. 13758 U. S. N. M. 13757 Inferior I antero-posterior 2 •3 mm. 2 . 2 mm. Inferior I transverse I •7 mm. I . 7 mm. P4 antero-posterior I . ,6 mm. I . 7 mm. P4 transverse I . .8 mm. I . 8 mm. Ml antero-posterior I . ■7 mm. I . 7 mm. Ml transverse 2 . IS mm. 2 . 15 mm. M2 antero-posterior I , ■7 mm. I . 8 mm. M2 transverse 2 , •3 mm. 2 . 15 mm. M3 antero-posterior I , .8 mm. I . 9 mm. Ms transverse 2 , . 0 mm. 2 . 3 mm. P4-M3, greatest length 6 ■5 mm. 6. 8 mm. M1-3, greatest length 5 •5 mm. 5 ■4 mm. 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A V-- / A I -70^- 4S A /A -, ’ >y / ■ .-.V vv >'' A rAVApy - ' AfA VI',* v*'^ /4 V ' £ s> A ssA sy>s A ■ y A#:' A A4 V ASS:VAAv/i/yyS'?\A*V s ■'//;/': sVsxS//7s//.':v/ysyi A's'is-,-- ”/ ' ■■ y 7- :|'iy/v As Ayy yy ^ ., ,^s AV4v,S;S, .., ■,;SS-, r At.: tS/i .if /f. ^.a/I'' V£' A S- 'C 'Sr4' -.' rV /7Ss S.S''A/AS'ft,/‘'y' ,sSAj /Ai'7//yvA«toy'SyiVSSAAAi%.j:| '■S / S-SA: ,. : ..y/AY A A-q: ' JAA,;A'-S'\r "Sf w ■ sA' ' S I- yf -^’-'s'.x A; oSAA-'vr '.'A’P-yAx A A AVAA 'yA:':A-yA"'A^Ay A y'A yA' ■ Aa y AAPAAy| A'''y'''AAA • A^;. yAA./ssv 7.' '^'Av-V^y'V'’ y ■■•^ y''^' ■ 'V-a-'J’s - .y 'A 'A ART. VI. A NEW NOTOSTRACAN GENUS FROM THE ORDOVICIAN OF SIBERIA. By B. F. Howell and Teiichi Kobayashi In the Ordovician strata of Europe and North America there have long been known to occur peculiar little fossils which paleontologists have had difficulty in classifying. They have been considered by some authors to be the remains of the carapaces of crustaceans of the order Notostraca. As a number of new species and genera have recently been discovered by one of us (Kobayashi)* in Upper Cambrian and Ordovician strata in Eastern Asia, and the distribution of these peculiar fossils is of unusual interest, he is now making a further comprehensive study of the group. Meanwhile it seems desirable to record the dis- covery of one new species, apparently the representative of a new genus, in a collection, secured from northern Siberia some years ago by Dr. I. P. Tolmachoff, Curator at the Carnegie Museum of Pittsburgh, which we are now studying. The collection contains besides several species of Wanwania, Ribeiria, Ribeirella, Ischyrma, and Eopteria. Our specimens came from a boulder, believed to be of Medial Ordovician age, that was found on the right bank of the Moyero River, six miles above the mouth of the Ukdama River, in northern Siberia. We are indebted to Dr. Tolmachoff for the opportunity to examine and describe them, and we take pleasure in naming the genus in his honor. Only the characterization of the genus and the descrip- tion of the genotype are presented here. The description of the other species will be included in a memoir, describing all the species in the collection, which we are preparing. *T. Kobayashi (1933) Faunal Study of the Wanwanian (Basal Ordovician) Series with Special Notes on the Ribeiridae and the Ellesmereoceroids, Jour. Fac. Sci. Imp. Univ. Tokyo, Sect. II, Vol. Ill, Pt. 7. 2 a }a4Q 59 Issued April ii, 1936. 60 Annals of the Carnegie Museum VOL. XXV Class CRUSTACEA Subclass EUCRUSTACEA Superorder BRANCIIIOPODA Order NOTOSTRACA Sars Family RIBEIRID.E Kobayashi Genus Tolmachovia, new genus The chief diagnostic characters of this genus are the fine concentric striations on the outer surface and, internally, the pair of vertical clavicles connected by a linear thickening, several concentric folds in the anterior half, and a radial fold running diagonally across the pos- terior. The genus resembles Billings’ genus, Ischyrina, in its outline, double clavicles, and posterior plication, but differs fundamentally from that genus in its external markings, which are radial in Ischyrina, but concentric in Tolmachovia. The genotype is the new species, Tolmachovia concentrica, de- scribed below. The genus is at present known only from the Middle Ordovician of northern Siberia. Tolmachovia concentrica, new species (Text figs. 1-4.) The carapace is moderately convex, depressed, and ovate, with a subangulate, subcentral umbo. The clavicles are equally developed and are connected by a ridge. There are five or six additional ridges, running parallel to the preceding ridge, in the anterior portion and on the ventral side of the ridge. A radial ridge crosses the posterior por- tion diagonally, starting from the posterior clavicle. Sometimes one or two additional, finer, ridges diverge from the beak to the venter on the inner side of the preceding radial. Only one of our specimens has the carapace preserved on its an- terior portion. This carapace is composed of a thick layer of calcare- ous material. Its surface is gently folded. Five folds were counted. They presumably correspond to the internal concentric ridges of the valve. The whole surface is marked by innumerable fine concentric lines. As the texture of the shell of the posterior portion cannot be seen on any of our specimens, we have not been able to determine whether or not the radial internal ridge affects the surface to form a radial keel. In the dorsal view of the internal mould the umbo is 1936 Howell & Kobayashi: A New Notostracan Genus 61 somewhat pyramidal in shape, and its four edges are sub-angulate. The scars of the clavicles are found a short distance below the apex, where the faces of the pyramid are very flat. The other faces are Figs. 1-4. Tolmachovia concentrica, new species. somewhat convex. In some of our specimens the concentric markings are not confined to the ventral part, but are present all the way up to the umbo. The holotype is No. the Geological Museum of the Russian Academy of Science. Paratypes are Nos. 2mb> 2027c^ ^274. All are figured except d. The holotype is illustrated in figures i and 2. MYCORRHIZiE FROM THE UINTA BASIN By l. k. henry 7' I ' _K ■ f\ I ' '>■ A f: >■ rA ■K ,■ : ,1 V ^'' r'"7,, f: V Reprinted from the Annals of the Carnegie Museum Vol. XXV, p. 63-72, 1936 Issued April 11, 1936 v_ I J- 4 ^ 4 r i, > 4 f / t > . , y \' ^ » \ 1 I , \ "r li ' x''|. ' f V u . '? ,V.M - X ix^Vx„:x:,-4 , I - ‘ ^ ,4' :":i ;vv''. '-: . 4^: . ; - : i'. ■ ' ;■ --■' .y V 4,1 ;•■, /L , , -r - Xj V .aVx,.- :-^,Xj 4-:: ■ , ,, ■ f - " y yy: /y-r/"/: . if . , - ' ^ ,1-:,, 4-"x:4 :r-r.xx-y: x: -X />''X , ' ' r; X' X‘ ^ . ,. , ; ' ' : ■ "X ^ ' -.'' ''',.■' -' ■; -• ’-4- '-'i X 1 : . . -■,,4', ■ I ' ■"- : . f ;■, . ' -x'-" ■'/•:' ■'■ / x; ' 0- :j ./ '; ’ 1 x" _ . X W--2 ART. VII. MYCORRHIZ^ FROM THE UINTA BASIN By L. K. Henry Sometime ago, I received for investigation a collection of tree and shrub rootlets from the Uinta Basin, Utah. These rootlets were col- lected during April, May, and June, 1933, by Dr. Edward H. Graham, Assistant Curator of Botany at the Carnegie Museum, while on his second expedition to the region. Microscopical examination of these rootlets revealed ectotrophic mycorrhizae on eight out of fifteen plants collected. The mycorrhizae occurred on one Aspen, one Birch, one Fir, two Spruces, and three Pines. No mycorrhizae were found on the Juniper, Sagebrush, Grease- wood, Maple, Service-berry, or Sumach. The ectotrophic mycorrhizae on all three pines were very similar. Externally, they consisted of coralloid clusters and swollen short roots; and internally, of thin prosenchymatous fungal mantles with hyphae projecting from the outer edge, intercellular fungal nets, and cortical cell hypertrophy. These eight hosts have been checked with an unpublished Mycorrhizal Host Index check-list and four are, I believe, new mycorrhizal hosts. The fifteen specimens examined may be arranged according to presence or absence of mycorrhizae: ECTOTROPHIC Abies lasiocarpa (Hooker) Nuttall. — Dwarf tree, Mt. Emmons, 11,400 feet elevation. *Betula fontinalis Sargent. — Dinosaur National Monument, in narrow ravine near spring, 5,000 feet elevation. Picea Engelmanni Parry. — Dwarf tree, Mt. Emmons, 11,400 feet elevation. Picea Engelmanni Parry. — Peak north of Trout Creek Ranger Station, 11,500 feet elevation. *Pinus brachyptera Engelmann. — Big Park, wooded base of Uinta River Canyon, 7,500 feet elevation. *New mycorrhizal host plants. 63 HARZs 1846 Issued April ii, 1936. 64 Annals of the Carnegie Museum VOL. XXV *Pinus edulis Engelmann. — North Slope of Grouse (Summit) Creek, circa 7,000 feet elevation (north slope of north rim of Basin). Finns Murrayana Balfour. — Trout Creek Ranger Station, 9,300 feet elevation. ^Populus aurea Tidestrom.- — Big Park, wooded base of Uinta River Canyon, 7,500 feet elevation. NO MYCORRHIZ^ Acer inlerius Britton.— Dinosaur National Monument, 5,000 feet elevation. Amelanchier utahensis Koehne. — Dinosaur National Monument, 5.000 feet elevation. Artemisia tridentata Nuttall. — Dinosaur National Monument, 5,000 feet elevation. Jimiperus utahensis (Engelmann) Lemmon. — Five miles north of Vernal, 6,000 feet elevation. Juniperus sihirica Burgsd.— Low shrub. Trout Creek Ranger Station, 9.000 feet elevation. Rhus trilobata Nuttall.- — Dinosaur National Monument, 5,000 feet elevation. Sarcobatus vermiculatus (Hooker) Torrey. — South of Skull Pass Quarry, east side of Green River, 20 miles south of Vernal, 5.000 feet elevation. The specimens examined were collected in the following vegetation zones^ beginning at the desert and ascending to the mountain top: Mixed Desert Shrub Zone. — 4,500 to 5,500 feet, typical desert, soil strongly alkaline. Artemisia tridentata Betula foyitinalis Sarcobatus vermiculatus *New mycorrhizal host plants. iThe names used foi the altitudinal z es m ‘^e suggested by Dr. Graham and those which he plans to use in describing the vegetation of the south slope of the Uinta Mountains in connectior 'd his rl n the flora and phytogeo- graphy of the Uinta Basin of northeas; , r I ' rthwestern Colorado. 1936 Henry: Mycorrhiz^ from the Uinta Basin 65 Juniper-Pinyon Zone. — 5,500 to 7,000 feet, desert-woodland, soil strongly alkaline. Acer interius A melancJiier utahensis Juniperus utahensis Rhus trilohata Sub-montane Shrub Zone. — 7,000 to 8,000 feet, desert-woodland. Pinus hrachyptera — in wooded base of Uinta River Canyon. Pinus ediilis — north slope of Grouse (Summit) Creek. Populus aurea — in wooded base of Uinta River Canyon. Aspen Zone. — 8,000 to 8,700 feet, pH reading neutral. No rootlets collected here, see above zone for Aspen. Lodgepole Pine Zone. — 8,700 to 10,000 feet, pH reading slightly acid. Juniperus sihirica — low shrub. Pmus murrayana Spruce-Fir Zone. — 10,000 to 11,000 feet, pH reading slightly acid at junction of this zone with former one. Picea Engelmanni Alpine Zone. — 11,000 to 13,500 feet, pH reading neutral. This zone is above tree line. Abies lasiocarpa — dwarf shrub. Picea Engelmanni — dwarf shrub. According to Dr. Graham’s findings, made with a La Motte Soil Teskit, the soils in the Mixed Desert Shrub zone and the Juniper zone were strongly alkaline with a pH reading of 7.5 as shown at Skull Pass Quarry and the Dinosaur National Monument. Certain locali- ties in the Aspen zone showed a pH of 7.0, while at the Junction of the Lodgepole Pine and the Spruce-Fir zones readings of pH 6 and pH 6- were obtained. The majority of the plants which possessed no mycorrhizse were growing in the alkaline desert-shrub or desert-woodlands below the Aspen zone, while all but three possessing mycorrhizae were located in neutral or acid soils at elevations above the Aspen zone. Juniperus sihirica, a low spreading shrub growing in the Lodgepole Pine zone, was the only non-mycorrhizal plant to appear above the alkaline desert habitats. Populus aurea, Pinus edulis, and Pinus hrachyptera from the Sub- montaine Shrub zone harbored ectotrophic mycorrhizse. However, they were growing in the forested sections and were not under typical 66 Annals of the Carnegie Museum VOL. XXV desert conditions. A reading of pH 6-|- was obtained from the soil around Finns hrachyptera indicating a slightly acid condition, due possibly to the decaying forest litter present there. Likewise, Betula fontinalis growing in a narrow canyon near a spring in the Mixed Desert Shrub zone showed ectotrophic mycorrhizae. Although Abies lasiocarpa and Picea Engelmanni have already been reported as mycorrhizal hosts, I feel that a detailed description of the ectotrophic mycorrhizae from these two dwarfed species growing at an elevation of 11,400 feet is not out of place here. Both species were growing among broken rock, Uinta quartzite, which forms the crest of the mountains throughout this region, and were above tree line which is at 11,000 feet elevation. They were deformed, dwarfed trees about four to five feet high located on a broad morainal mound above the Chain lakes on the southeast slope of Mt. Emmons. Al- though no pH readings were made near these dwarfed trees, a pH of 7 was obtained in small meadow-like patches among the same type of rocks in this Alpine zone. Sufiicient litter was collected under these dwarfed trees to form soil. Externally, the rootlets of Abies lasiocarpa possessed typical coralloid clusters of mycorrhizae, some of which were gray in color, and gray swollen mycorrhizal short roots (plate X, fig. 2a) ; while internally, the mycorrhizae showed thick prosenchyma- tous fungal mantles with hyphae projecting from the outer edge, some of which have cross-walls and clamp connections, intercellular fungal nets, and cortical cell hypertrophy (plate X, fig. 2b). Mycorrhizae of Picea Engelmanni possessed similar external char- acteristics (plate X, fig. 3a), while the internal structures were alike except for the thin prosenchymatous fungal mantles and absence of clamp connections on the projecting hyphae (plate X, fig. 3b). This thin prosenchymatous fungal mantle is in accord with the ones found upon the same species growing in the Spruce-Fir zone at an elevation of 10,500 feet. The habitat, external form and internal structure of mycorrhizae on four new host plants are described in Table I. SUMMARY Ectotrophic mycorrhizae were found on eight of the fifteen different plant rootlets collected in the Uinta Basin, Utah. Five of the ectotro- phic hosts were growing above the Aspen zone where soil is neutral or 1936 Henry: Mycorrhiz^ from the Uinta Basin 67 slightly acid, while the remaining three occurred in the Sub-montane Shrub zone but in locations where the soil conditions were not that of a typical alkaline desert. All but one of the non-mycorrhizal species were located in the desert-shrub and desert-woodlands where the soil was distinctly alkaline. The dwarfed trees of Abies lasiocarpa and Picea Engelmanni growing in scanty soil among the rocks in the Alpine zone were excellent mycorrhizal hosts. These findings would seem to indicate that the ectotrophic mycorrhizae were able to develop and flourish in sand and sand-humus mixtures as long as the soil re- mained in neutral or acid condition, but were unable to live under the alkaline soil conditions of the desert. Betula fontinalis, Pinus brachyptera, Pinus edulis and Populus aurea are new additions to the mycorrhizal host list. Data for New Mycorrhizal Host Plants 68 Annals of the Carnegie Museum VOL. XXV o-B (D O Q c u A T3 S-i a; tr! a ^ 'o § „ X 03 oj '^ -- - o3 O M 'O • r5 d ci3 t: m 5 ’S g ^ Td '5 X d 1-^ •d dl 03 03 > d d '-M n ) o .a 03 d d -LS d d ° }-i ro a 03 8 .• d M o d d M d -•-> 1=^ a 0 d 1 a d a d 2 'a -M d CJ o» 03 !-i d d d a O O o d a -B >. CJ d 03 ^2 Q. CO ^ a d >> •d d d o d l^s 2 >> -d 03 -d d -d a o d o a 03 a ^ CJ ir d Bd 8 d M o3 a. as ^ ^ d 2 CJ d S 2 S § a d a 2 jz 'd. .2 P d a ^ B SI'S S) a d d g B bJO . g.a B d -d d ^■2b 03 a d S g 8 'gad S d .a o ^ 7B ^ CO 8>< 03 d ti o B „ d- d ^ d o CJ d o 8 g: T3 -d 2 B (Z) ^ B ^ a d Co 2 B 1 ^ tB ® 03 B d B B g CO P d! ’-' a B Bb ^ a id cli —all ig. la B B o ^ d d B B M • B - S B B O CO dx B CO (N B CO ^ B Q d l-l B o d (A B cd d 03 d d M ^ o d "d d O u U U U 03 d 70 Annals of the Carnegie Museum VOL. XXV EXPLANATION OF PLATE IX Fig. la. Coralloid cluster of ectotrophic mycorrhizae from Betula fontinalis X 14. Fig. ib. Cross section of ectotrophic mycorrhiza from Betula fontinalis X 156. Fig. 2a. Mycorrhizal short roots from Pinus brachyptera X 14. Fig. 2b. Cross section of ectotrophic mycorrhiza from Pinus brachyptera X 156. Fig. 3a. Coralloid cluster of ectotrophic mycorrhizae from Pinus edulis X 14. Fig. 3b. Cross section of ectotrophic mycorrhiza from Pinus edulis X 156. ANNALS CARNEGIE MUSEUM, Vol. XXV, Plate IX. 3a 3b ] r. t 72 Annals of the Carnegie Museum VOL. XXV EXPLANATION OF PLATE X Fig. la. Coralloid cluster of ectotrophic mycorrhizae from Populus aurea X 14. Fig. ib. Cross section of ectotrophic mycorrhiza from Populus aurea X 156. Fig. 2a. Mycorrhizal short roots, one dichotomously branched, from Abies lasio- carpa (dwarf tree) X 14. Fig. 2b. Cross section of ectotrophic mycorrhiza from Abies lasiocarpa X 156. Fig. 3a. Mycorrhizal short roots from Picea Engelmanni (dwarf tree) X 14. Fig. 3b. Cross section of ectotrophic mycorrhiza from Picea Engelmanni X 156. ANNALS CARNEGIE MUSEUM, Vol. XXV. Plate X. lb 3a ■t i I ^'Z^ ^i~t^'f!> -Z:’'i})^^ lU _ r< ‘ ?’^ l\ ^ r ,' ^ ^ ^ ■ V!\’ ^ . r ^ 'i^-' ’j s \ ''!| ^ z' V' //V / ^ ,^^'J X ''^ ‘\ ^ ^ , W X ''X-\7'^' >-'X \ ^ ’ \ “'^‘^ " /'’ 'i ; :.-^:'>Xv.X :,x >4x7X7 , -V X ■^\z^ ^' . r I z rx \ ■x ^ ^ i w ' .-h>'' vX'- .Y”;X''''.J:' ‘ '•-.' ;X Xv'^YX' ' X’' r . ; ;^x. ■■ _\.. «■! ^ ' V ^ A ‘ .^x '• s ^. -V^Y ^•■- ’''.vXx.^Ji ’V'\- "• ■ -sV 4 ' > . . X ■’ ' ' L X " X / Xx . ,., m - ■ j-T" ' ('■■-' -• ' vv S' Y '. ' f . ..'X;^:./' • .-• :/ ''H- X .w Y/ '*'2 ' k X- . V X '^x„ XX"^ 'Y ;/ 7,7: Yx^'' /-'■■ -VXI'- » / ; . L"'.' 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V"‘YXo,-7'^'^ < , / 'Y X/Y Y7Y-.VV’' 'X- 5 ;’7.77777/j.7 ,/,..r7. x- 777.7-4, ■„ 7.,''^ . :« ;.' , J'7 7',4f 7 -7 77-: 77 -xt 7 .;■ , ;, ■ i-! '/. 7^7 A V -77. = 7 'XV7 ■:: -7-'Y^4-7''7,v^^..^:Y- 7 VYY7>7’'''-:A7;.-':7 ■(> :V77'’'V7;V> X,74s'Y>‘'.: 5'T'V .''-77, . i: H V' 7VY7'f:,"4x- 'A ; a' 'V'-^ “ '' ' ^ YVV'Y YX.-iy 3 ’XY ':.-^ A -717- •"- 7,v""- 7 N)- 74 .''/7.,j44^.' y.'AVA, *'' ••'y'!: ^ 7.. Y ‘ ’A ^ ' v A v ■>' - kdY ■y-. - W'5.7-.;:'' ^-77 ,7.73-; -A:>7v7 -1.-- ,', '■"*7'.' ''7 i>-^ '/'' ■■>}Z^' P\.' ■ ■' -’ ’ Y Y' , A 7"^ Y X Y , :Y : X V -v " ^ ” Y ,7 vv- r 7 ’:■ 'Z-zp YaV ?'"-■ y 'Xav-- rzz, [ zp ^ .:X:XXX:-.7.-7'^. -7,,..7‘,V7;-;J-' -,(<-7 .' v-jt' -vAV 7' '\, ■' ■'^’-' .XY V?Y^ 7: "’YxC..,. . Cr:- y Y U ■, )7 Y A S'. A NEW SCOLECODONT GENUS, ILDRAITES, FROM ^ THE UPPER DEVONIAN OF NEW YORK I J- BY E. R. Eller ,4. : ' Reprinted from the Annals of the Carnegie Museum Vol. XXV, p. 73-76, 1936 Issued May 1, 1936 '•' ' T ) ^ f- ' ' i< . ' ) '■' 'r r ' ' -, ^ '' ' ' ; ' , T' 4- ;V'K^. ’ -'A. ■ 'y.^^4C - - ; ’' ''0- ^ ■' ■ ’ A A }A -aa^-:AsA^^:a-- iA>J . i - ^ ' .,...i .-.fe.r - - ... - \ '''.'“ ■ 'X i -4- '-•', i" j ' «■''.' ' I,' , '^‘ ^' ' : ^ '.V i I '"A •'■ ' : ■X 4xzAA X-XAAA'AAAAA'MyX >A -'fi'f ■ A'A'A'A ■^>:‘:A ■ v^'f ■ :- ZxZ4X c ■' 4A‘ApAA:^9 9Z:i mzk - '■ a -.; A^ZAAx :■ Xy ^ - : ., ^ ^ ^ '" AA'XZ-’- x4y' ';.' ':' X::4<^i- a\i .- :■ V - ~~try. . . . N .-i - .-...V - . ■ ^ ■ '■' 'kJ \A X'-'~yA4'^^ ■' '''"■'V ^ :' 'Ma feSSaa'A*;- . v , XfAMSAAVy. 'a. ’ zkyAy'-iy ^Xy yX^ ry^-^' l;AAy:Z,xy Jy- ffyzyz - ; 1 V- .a- ' , p' r .>t' - 4' 4 .*. -zj z ^ z. , -r{ a,. /;c. Vz. ' ' ' ' -:, zzyz ■' .) . A'" ~ ?',,>- ■'* Ai'a ' ’ .' -' '- '! 'o 7'-' z '- ' a-. : a ; --L '■. a ’;^,-.aa.r‘> . ."a~ ;''Aiaa Ay^ y, ' ; r..v :r\ .' ■v./a-'V..-:':' ■.a'/,-,; A,„aava.aj). y^x X v"aAc:Aa ART. VIII. A NEW SCOLECODONT GENUS, ILDRAITES, FROM THE UPPER DEVONIAN OF NEW YORK By E. R. Eller In a paper published in 1934^ the writer described among other genera four species of Arahellites, polychaete annelid jaws or scole- codonts. Three of these species, Arahellites spatiosioris, Arahellites latiis, and Arahellites alfredensis resembled each other quite closely, differing only in size, sculpture of the surface, and character of the denticles. One species, Arahellites hipennis, was quite unlike the other three. In this description the writer remarked that the jaws did not correspond (since this posterior margin is incurved) to any figured by other authors, except possibly a species, Arahellites spicatus figured by Hinde,2 but that the anterior portion including the hook and denticles compared well with Arahellites alfredensis. In the collecting season of 1935, the writer had the good fortune to find an articulated annelid jaw assemblage, plate XI, fig. i, at Alfred Station, New York, in the Alfred Shale, a local facies of the Gowanda beds, Canadaway Group of recent workers, and historically known as the Chemung of the Upper Devonian. The maxillae I of this new articulated jaw apparatus corresponds to Arahellites hipennis, plate XXIII, figs. 8-10, and maxillae II to Eunicites anchoralis, plate XXII, figs. 1-5 of the former paper. This new specimen again brought to the mind of the writer the question as to whether Arahellites hi- pennis and Arahellites spicatus Hinde should not be considered as generically different from the genus Arahellites. With this in mind the writer reviewed the various material and has decided to place the jaws of this kind in a new genus. Genus Ildraites, gen. nov. Genotype Arahellites hipennis Eller The anterior extremity of the jaws of maxillae I has an extremely prominent pointed hook and a row of several usually acute denticles lAnn. Car. Mus., Vol. XXII. pp. 303-316, pis. XXII, XXIII, 1934. 2Bihang till k. Svensk. Handl., Vol. 7, N:05, p. 18, pi. 2, figs. 47-49, 1882. 73 Issued May i, 1936. 74 Annals of the Carnegie Museum VOL. XXV along the nearly straight inner lateral margin. The anterior part is very similar to Arabellites. The posterior portion is sickle-shaped by a deep crescent-shaped bight while the posterior end of Arabellites is obliquely truncate. The reason for erecting a new genus is based primarily on this difference in the posterior portion of the jaw (maxillae I). This structural difference probably depends on a type of muscular attachment in the jaws which was different from that of other genera and it might be also reflected in the general morphology of this annelid. In other words, this structure suggests such alterations in the structure of the animal that generic separation of corresponding forms becomes, in the opinion of the writer, quite necessary. Ildraites bipennis (Eller) Maxilla I, II, and III or IV (?), Mandible (?) Arabellites bipennis Eller. Ann. Car. Mus., 1934, Vol. XXII, p. 311, pi. XXIII, figs. 8, 9, 10. Eunicites anchoralis Eller. Ann. Car. Mus., 1934, Vol. XXII, p. 307, pi. XXII, figs. 1-5. Maxilla I The paired jaws of maxillae I were lying on the top of the jaws of maxillae II especially along the inner lateral margin. During the clean- ing off of the outer lateral margin of maxillae II the part covering maxillae I was destroyed and now we have only a partial impression of the latter. However, enough of the maxillae I is preserved to give a clear idea about its form. The ends of both hooks are absent, not being preserved even as impressions. On the right jaw the impression of several denticles is discernible. Maxill.e II The jaws of maxillae II resemble Eunicites anchoralis of the former paper. The jaws under consideration in the present paper are com- plete except that the ends of the hooks and two of the denticles of the left jaw are broken off. The jaws are longer and they bear several more denticles than those figured before. The anterior part bears a bight corresponding in its general form to that of maxillae I. The denticles are asymmetrical and have a well-worn appearance. Some of the denticles appear to have a groove in their middle. All denticles 1936 Eller: A Scolecodont from the Upper Devonian 75 are bent upwards and inwards at more than 45° which is probably due to compression since in other specimens of this species the denticles lay in the same plane as the jaw. Under high magnification the surface of the jaws seems to be slightly pitted and also covered with minute tubercles. Maxilla III or IV Near the left jaw of maxillae II and partly under it is seen a jaw of maxillae III or IV. The jaw has a row of blunt denticles along its inner margin. So little of the jaw is visible that it is not possible to describe it in any detail. However, due to its large size the jaw probably belongs to maxillae III, although it is not possible to be positive about it. Mandible (?) In front of the jaw apparatus is seen an extremely thin, chitinous- like plate that extends back under the left jaws of maxillae I and II. It was impossible to remove any of the matrix without breaking the jaw. The size and character of this jaw part suggests the possibility that it may be a mandible. In examining the jaw apparatus no apertures or irregularities on the surface for the attachment of muscles have been found. It is interesting to note that a similar bighted structure is present on both the jaws of maxillae I and maxillae II. On maxillae I it is a deep in- dentation in the posterior area with two shanks. In the jaws of maxillae II the anterior margin is extended into a shank which forms the characteristic bight. These shanks and bights were probably for the attachment of the muscular system. In the modern family LeodicidcB many of the maxilla II have also deep indentations in the posterior area. 76 Annals of the Carnegie Museum VOL. XXV EXPLANATION OF PLATE XI Figure magnified 45 times. Fig. I Ildraites bipennis (Eller) Alfred Station, New York. Maxillae I, impression, left and right jaws. Maxillae II, left and right jaws. Maxillae III or IV, left or right jaw, extends from under the left jaw of maxillae 11. Mandible (?), seen near the anterior end of the left jaws of maxillae I and 11. The type is in the Carnegie Museum, number 8049, Section of Invertebrate Paleontology. ANNALS CARNEGIE MUSEUM, Vol. XXV. 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'0 0r-r::\2^ 0k0u0y’m,p0^\0p,i0-- t7-;« ”’ m20m0B0m0^0)400m!ik 0p -'0'f:":00^'i'0M0 '. 7f4 4'' v-4 7 ':4' 47 y3>44y:-.’-,,7”. . o.Y\ ‘Hi’ 40f-0P00:p00.).0:A,j0^0. yp: :0000:i0000.yp. 000:l}00 0 .v'^ f ■ , ^ ’.'v: '’?:dvi'7, „ ^4"' ^ 7f7\ ^ -im0 7,-7' ,/ ■ -.47' Am.7AA'^-44AA' v V 7 '.'.’^ ■ . 7, ^‘>-. ^ ^4-' ■ 7"7, ' /^ 02^^ ii -00 47 ' /.u'7..Y.,/--:^7.474:4-7 -..i;y74- 0:i .;•<- ••, I -■ .f-^ 44 00- f 4 7744 -4 '"4 -7 "4* 7, 4-- .. ; <’■ 74.,- 7 -k-J'- 00-1' -I ^4:4 74^'^7.. .. .0 -2p.0-J‘ 20-1' ' ‘ AIyY,. (V ^4 74. ■ '/)'!' ’,■ ., ■ V 0P 'r,i -S 2U0X THE CHAIN SlSfAKE, LAMPROPELTIS GETULUS GETULUS (L.) IN WEST VIRGINIA AND PENNSYLVANIA , ■ '■ a.: - ’ i By m. Graham Netting ; - 'U ! ^ i' *v • ' . . F ■'-V' ■ f ■. ^ 1) , 1 I 'F Reprinted from the Annals of the Carnegie Museum Vol. XXV, p. 77-82, 1936 Issued June 25, 1936 ml ART. IX. THE CHAIN SNAKE, LAMPROPELTIS GETULUS GETULUS (L.), IN WEST VIRGINIA AND PENNSYLVANIA* By M. Graham Netting The Carnegie Museum recently received a Chain Snake from West Virginia which appears to be the first of its species from the state to reach a museum collection. This donation has stimulated me to consider the distribution of this snake in the northern half of its range, and its habitat preferences. In the summer of 1931, I had the pleasure of spending a week in Pocahontas County, W. Va., as a member of the staff of the Oglebay Nature Training School. Many of my students proved to be excel- lent collectors and observers, and I have frequently had cause to be thankful for the friendships formed at this time, and renewed in later years, since many of these students have been most energetic in add- ing to the West Virginian collections of the Carnegie Museum. One of my students that year, and one of our most faithful contributors since then, Mr. Chester M. Shaffer, of Dorcas, W. Va., wrote to me, under date of May 15, 1932, as follows: “This afternoon I found a Chain king snake on the highway two miles north of Petersburg. It is the first one I have seen in West Virginia. The snake had just been killed by a passer-by. It was about 3 ft. in length, almost black with narrow white bands across the back about an inch or so apart. These bands widened to large white spots on the sides. The belly was largely white varying from white to yellowish white spotted.” This description, and Mr. Shaffer’s familiarity with the species in Florida, led me to accept his statement as the first valid report of the occurrence of Lampropeltis getulus getulus (L.) in West Virginia. Unfortunately, the specimen was not preserved. *I am especially indebted to Dr. Donald A. Cadzow, of the Pennsylvania Historical Commission, who sent me a print of the section of his film which showed a snake captured near Safe Harbor. I am also indebted to Mr. Neil D. Richmond, Dr. F. N. Blanchard, Mr. Roger Conant, Mr. W. Stuart Cramer, Dr. E. R. Dunn, Dr, George F. Johnson, and Mr. Carl F. Kauffeld, for pertinent information or assistance. MAR 25 77 Issued June 25, 1936. 78 Annals of the Carnegie Museum VOL. XXV More recently I learned from Mr. Neil D. Richmond, of Fairmont, W. Va., that a Chain Snake was secured in the South Branch Valley, near Franklin, Pendleton Co., W. Va., during 1931. This specimen was kept alive in the Biology Department of the State Teachers Col- lege at Fairmont until 1934, when it died and was discarded, at a time when Mr. Richmond was away. At Franklin, the South Branch of the Potomac River is at an altitude of about 1600 feet, and its valley is here rather narrow since it is hemmed in by mountain ridges. Last October Mr. Shaffer donated to the Carnegie Museum a well- preserved Chain Snake which he found on Sept. 14, 1935 freshly killed on the highway two miles north of Petersburg, Grant Co., W. Va. This specimen (CM 8719) was found less than fifty yards from the place where he saw the 1932 specimen. Petersburg itself is in the South Branch Valley, but the point where the specimens were found is located at an elevation of about 950 feet in the broad valley of Lunice Creek, which empties into the South Branch at Petersburg. The specimen has the lower jaw torn away, but except for this it is in excellent condition. It measures 1010 mm. in total length, which is probably less than was its length in life, since it is too tightly coiled to permit accurate measurement. Each character falls well within the limits of variation listed by Blanchard (1921: 55-58) for northern specimens of this species. Surface (1906: 174) lists the Chain Snake as of possible occurrence in Pennsylvania, but states ‘‘we have not collected nor received speci- mens in the State.” Surface’s (Ibid: opp. 176) plate XXXI, which is labeled “Milk Snake or Mouse Snake {Lampropeltis doliatus tri- angulus)," is obviously L. getulus getulus. It is improbable that this plate was based upon a Pennsylvanian specimen for the original photograph was made by Wm. H. Fisher, of Baltimore, and pre- sumably represents a Maryland specimen. There is a distinct possi- bility, however, that Surface, failing to distinguish the two species, may have had specimens of both listed under L. d. triangulus. To the shame of the Commonwealth of Pennsylvania, Surface’s valuable collections have been allowed to disintegrate completely in less than twenty-five years and, consequently, in questionable cases of this kind we can only indulge in idle speculation. Last year Mr. W. Stuart Cramer, of the Philadelphia Zoological Garden, examined, at my request, certain specimens in the collec- tion of the Franklin and Marshall College Museum, at Lancaster, Pa. 1936 Netting: Chain Snake in W. Va. and Pa. 79 Mr. Cramer was formerly located at Lancaster, and he is familiar with collecting conditions in Lancaster County, and with the work of all of the recent collectors in that area. This knowledge enabled him to determine definitely that the only specimen of L. g. getuliis in the Franklin and Marshall Museum actually came from just north of Port Deposit, Maryland (approximately eight miles south of the Pennsylvania line) even though it is erroneously labeled "Southern Lancaster County along the Susquehanna, May 25, 1929, Roy Palm- er.” In the absence of a preserved specimen, and in view of the fact that Dr. Roddy (1928: 41) reprinted Surface’s plate without correcting the title. Dr. Roddy’s (Ibid: 40) statement "This interest- ing snake of the Carolinian zone has been observed and taken several times in the Susquehanna Valley below the mouth of the Conestoga Creek” cannot be accepted as positive proof that the species occurs in Pennsylvania. A few years ago. Dr. E. R. Dunn, of Haverford College, viewed a motion picture record of archaeological work in the vicinity of Safe Harbor, Lancaster Co., Pa., which had been directed by Dr. Donald A. Cadzow of the Pennsylvania Historical Commission. In one sec- tion of the picture a student was shown holding a large snake which the narrator described as having been secured in the vicinity. Dr. Dunn wrote me that the snake shown so fleetingly in the film appeared to be a Chain Snake. In recent correspondence Dr. Cadzow has in- formed me that the specimen photographed was found on Grubb Creek near Shenk’s Ferry, about a mile and a half below the mouth of Conestoga Creek, Lancaster Co., Pa., in the summer of 1931, and that it was later released. Dr. Cadzow also sent me a print of the section of the 16 mm. film on which the snake occurs. I have had this bit of film enlarged, and I have also tried projecting it. In both instances, the grain prevented clear definition of the markings of the specimen. However, narrow bars are plainly visible in the enlargement and I am personally convinced that the specimen photographed was a Chain Snake. This evidence, although not conclusive, indicates that L. g. getuliis should be tentatively included in the herpetofauna of Pennsylvania, even though there appears to be no preserved Penn- sylvanian specimen in any collection. Although there are numerous records of the occurrence of the species in Maryland, the Port Deposit specimen, referred to above, appears to be the only record for the Susquehanna Valley in this 80 Annals of the Carnegie Museum VOL. XXV state. There are several additional records for the Chesapeake drain- age, and a number of records for the Potomac drainage, but I have not been able to secure any records for Maryland west of eastern Montgomery County. When the distributional records of the Chain Snake are plotted on a map, it is at once apparent that this species is common in the large river valleys in the Coastal Plain and Piedmont Provinces of Virginia, but that the Blue Ridge Mountains have prevented it from spreading westward. However, the West Virginia records listed in this paper prove that this wall is penetrable at one spot where the Potomac Valley has served as a ‘‘gateway to the west.” This evidence that the species has succeeded in passing the mountain barrier, and has ex- tended its range as far as the South Branch Valley makes its absence in the Shenandoah Valley both surprising and questionable. I feel assured that the present lack of records for western Maryland and the Shenandoah will be corrected by more collecting in those areas. In Pennsylvania the species may eventually be found to be a rare inhabi- tant of the Potomac and lower Susquehanna drainage areas, from Bedford County east to Lancaster County. Ditmars (1907: 362) describes the habitat of the Chain Snake as follows: “Specimens captured by the writer were in rather dry patches of timber; some were taken while basking in the sun of small glades in the forest; others were found hiding under fallen tree trunks.” Wright and Bishop (1915: 169) point out that the species prefers the drier parts of the Okefinokee Swamp. Corrington (1929: 74) states, “taken in the coastal plain only. The principal collections were from marshy situations, less often in drier woods.” The number of some- what conflicting statements concerning the habitat of this species could be multiplied by citing additional authors. Mr. Roger Conant informs me that captive specimens of the Chain Snake spend much of their time lying in their water pans. Mr. W. Stuart Cramer once col- lected a specimen in the water in a brackish swamp at Heislerville, near Cape May, Cumberland Co., N. J., which disgorged a large Matrix sipedon sipedon. The published statements of Wright and Bishop, and Corrington, and the experiences of Conant and Cramer indicate that this species is more moisture-loving than Ditmars’ statement indicates. The preponderance of records along rivers, coupled with the almost complete absence of records in upland situations at any considerable 1936 Netting: Chain Snake in W. Va, and Pa. 81 distance from streams, is additional evidence that L. g. getulus is limited to the vicinity of moderate to large-sized bodies of water which provide broad valleys, marshes, or moderately open and gently sloping terrain. Turtles flourish under identical conditions, and the extreme fondness of the Chain Snake for turtle eggs (Wright and Bishop, 1915: 170) lends support to the view that the snake is normally associated with turtles and is relatively common in habitats which turtles select for egg-laying. Although the species is practically re- stricted to the type of environment outlined above, the citations indicate that it frequently selects the drier situations of this environ- ment. In the northern half of its range, at least, the Chain Snake may be said to occur in, or within a daily range of, fluvial habitats within the Piedmont and Coastal Plain Provinces. I doubt whether the species may properly be considered an inhabitant of the Atlantic Coast littoral, for the many coastal records are all located near river mouths. Beyond the Blue Ridge, within the Valley and Ridge Province, it is to be expected only in the valleys of Atlantic drainage streams which have provided water gaps for convenient penetration of the mountains. I do not believe that stream capture at the head- waters of mountain streams has ever affected the distribution of this form. It should be noted that Stejneger and Barbour in the third edition of the Check List (1933: 108) add West Virginia to the range of LampropelHs getulus nigra (Yarrow). I am not aware of the evidence upon which this range extension is based, but the occurrence of the form at Hanging Rock, Ohio (Blanchard, 1921 : 48) which is only a few miles down the Ohio River from Huntingdon, W. Va., has led me to expect it in West Virginia. The very region — along the Kentucky border — where it should be collected is zoologically the least known area of West Virginia. Residents along the Big Sandy and Guyandot Rivers should be encouraged to search for this interesting snake. SUMMARY 1. Lampropeltis getulus getulus is recorded from West Virginia for the first time on the basis of one existing specimen, and two additional records of specimens which were not preserved. 2. The former published records for Pennsylvania are considered insufficient to establish the occurrence of the species in the state, but 82 Annals of the Carnegie Museum VOL. XXV it is admitted to the state fauna on the basis of a single Lancaster County specimen which was captured, photographed, and released. 3. The habitat, and the distribution of the Chain Snake in the northern part of its range are discussed. It is concluded that the species inhabits large valleys of Atlantic drainage streams, and that it has penetrated west of the Blue Ridge only where water gaps have provided entry to the Valley and Ridge Province. 4. The need of specimens of Lampropeltis getulus nigra from West Virginia is mentioned, and the area where these may be found is out- lined. LITERATURE CITED Blanchard, F. N. 1921. A Revision of the King Snakes: Genus Lampropeltis, U. S. N. M. Bull. 1 14: vi-|-26o, 78 figs. CORRINGTON, JULIAN D. 1929. Herpetology of the Columbia, South Carolina, Region. Copeia, no. 172: 58-83. Ditmars, R. L. 1907. The Reptile Book. Doubleday, Page, & Co., New York: xxxii-l-472, pis. 1-136. Roddy, H. J. 1928. Reptiles of Lancaster County and the State of Penn- sylvania, Publ. Dept. Nat. Hist, of Franklin & Marshall Col- lege, Science Press, Lancaster: 1-16, illus. Stejneger, L. & Barbour, T. 1933. A Check List of North American Amphibians and Reptiles, 3rd ed. Harvard Univ. Press, Cambridge: xiv-l-185. Surface, H. A. 1906. The Serpents of Pennsylvania. Monthly Bull., Div. Zook, Pa. Dept. Agri., 4, nos. 4-5; 113-208, pis. 15-42. Wright, A. H. & Bishop, S. C. 1915. A Biological Reconnaissance of the Okefinokee Swamp in Georgia. H. Snakes. Proc. Acad. Nat. Sci. Philadelphia, 67: 139-192, pi. 1-3. i- V .y y'^'c^y i :• . ■^"Vi '- ?''r^: iy%yv-)((ry^"'^ y ^y -■ ^ y^'yy'h', yy yiiydiy^y'^^y y^'-y-^ y'-y^yr^'. „ V' >r. . i'"-': ^t^v4 ' " \lr ^ /t-. ^„' ; '' 'V ' t X 1 > •''' ^ A*';,' ..‘.'fa , f " y "x^" •.’r’ /;-vr% ^ r-. ' y X, hy \ y y'^ i yy y'/ . 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Xx" 44;y>' 7)' 444’^ 'Xxk k " ' ^^^.4 k #y444#y 44’^^ xyA'--- JC>^ THE LAND AND FRESHWATER MOLLUSCA OF NEWFOUNDLAND X: BY S. T. Brooks r r - I Reprinted from the Annals of the Carnegie Museum Vol. XXV, p. 83-108, 1936 ^ Issued June 20, 1936 S> .1 f. t j I ART. X. THE LAND AND FRESHWATER MOLLUSCA ■ OF NEWFOUNDLAND I By S. T. Brooks ' (Plates XII and XIII) Newfoundland was visited by the author during the summer of ' 1934 to obtain material for a further study of the early distribution of the molluscan species in North America. Collections in this area were sparse but the records contained below seem to be of sufficient value to warrant publication. Many of these are from specimens in the collections of the Carnegie Museum and the ones included from the records of other institutions are so designated. I wish to acknowledge the kindness of the following individuals and institutions who have given me such great aid in this work: Dr. A. Avinoff and the Trustees of this museum who made the trip possible; Dr. F. C. Baker of the Museum of the University of Illinois for his study of the Lymnmdce; Mr. Calvin Goodrich and Dr. Van der Schalie of the Museum of the University of Michigan for their identification of the Naiades; Mr. W. J. Clench for his designations of the PhysidcB and for the several records^ from the Museum of Compara- tive Zoology; Dr. H. A. Pilsbry and Air. E. G. Vanatta for their assistance and for the use of the many records^ published by Mr. Vanatta, and to Dr. Paul Bartsch and Dr. Harald Rehder for their identification of the SuccineidcB and for the records^ from the United States National Museum. Paratypes of the species herein described have been deposited by Dr. Baker and myself with the following: United States National Museum; Museum of Comparative Zoology; Museum of the University of Michigan; Museum of the University of Illinois; the Academy of Natural Sciences of Philadelphia; and with Dr. S. S. Berry of California. Newfoundland is the tenth largest island and within the last year or so it has reverted to its original status as a British Colony.^ Not ^Indicated in text as M. C. Z. ^Indicated in text as A. N. S. P. ^Indicated in text as U. S. N. M. ^The first British Colony and the beginning of the present British Empire. 83 MAR 25 1940 Issued June 20, 1936. 84 Annals of the Carnegie Museum vol. xxv CHART OF SPECIES SHOWING THEIR DISTRIBUTION Species Helicigona arbustorum Helix hortensis Zonitoides arborea Hawaii minuscula* Striatura exiguua* “ milium* Euconulus fulvus Retinella electrina Vitrina limpida Deroceras agrestris “ Iseve Gonyodiscus cronkhitei “ “ anthonyi Helicodiscus parallelus Punctum pygmaeum minutissimum* . . . . Arion ater “ hortensis “ fasciatus Succinea ovalis “ a vara “ peoriensis* “ groenlandica* “ verilli Pupilla muscorum Vertigo modesta “ “ parietalis “ “ castanea “ gouldi paradoxa St ‘ elatior Columella edentula Cochlicopa lubrica Vallonia albula Zoogenites harpa Planogyra asteriscus* Carychium exiguum* Stagnicola palustris “ “ papyracea** “ “ perpalustris** “ newfoundlandensis** Fossaria obrussa* “ “ brooksi** “ umbilicata* Radix pereger geisericola Gyraulus parvus “ hornensis* Helisoma companulata davisii* “ “ minor* Physa gyrina “ heterostropha Valvata lewisii Valvata sincera n3danderi* Amnicola limosa porata*? Distribution West Central East X Importation X X X X X X X X X X X X X X X X X Importation? X X X X X X X X X X Importation X X ’ * X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X Margaritana margaritifera X X X Anodonta marginata X ■ . X “ implicata* . . X *new records. **new to science. 1936 Brooks: Mollusca of Newfoundland 85 that this political change has affected the molluscan kingdom, but at the present time a greater interest is manifest in Newfoundland about its animal and plant life than ever before. A friendly coopera- tion will meet the collector upon the island whether he is in search of plants, insects, or just plain “wrinkles” (snails) and “pussels” (mussels). Two months were spent in traversing this region and the three major areas with their collections are indicated upon the accompanying chart. As shown here the greatest number of forms are western in distribution. This is in the region of which parts were untouched by the ice of the later Pleistocene Period. However, as it will undoubt- edly be proven that the western portion does have the greatest number of native species, it is also necessary to point out that the present disparity is due mainly to the greater number of collections having been made on the west coast. It is necessary that more work be done around and west of Shoal Harbor, on the east coast, in order that the present distribution be properly known. At the present writing it seems that, since the Pleistocene, the dispersal of forms in New- foundland has been generally eastward but a final picture of this phenomenon must await more collections in the eastern, southern, and central areas. Not surprising, but quite instructive, was the finding that the great group of land snails have, through this long period of isolation, shown such a basic genetic strength in that they have retained specifically the features of their kind now living in northern Asia, Europe, and America. In no case are there any differences strikingly separating them from their other northern relatives. This lack of change is per- haps due to the similarity of the climate and vegetation of New- foundland to that of the mainland. On the other hand, this long period of isolation has caused a greater change among the plants of the island. Many differing from those of the mainland clothe the hills and valleys and point to a long period of differentiation since, as well as prior to, the last glaciation. This paper, then, is not so much a statement of new facts as a restatement of facts and assumptions derived from earlier studies of the northern faunal areas of the earth. It is, however, as complete a resume of the present knowledge of the molluscan fauna of Newfoundland as it is possible to make on the basis of the scanty collections. 86 Annals of the Carnegie Museum VOL. XXV DISCUSSION OF DISTRIBUTION OF THE FAUNA A discussion of distribution, even in its most recent phases, is ever fraught with danger, and necessitates a certain form of mental gym- nastics and credence well flavored with doubt. In the first place we know from other studies that there are certain circumpolar species which have been distributed to the three continents through what we might call the northern door; a sort of circumpolar ‘'Garden of Eden” (Plate XIII). A study of these species will bring up in our imagina- tions a picture that shows a former continuity of the land masses. By means of this picture we can easily see how the various forms were dispersed and how through their migrations they advanced into each land, bounded only by the various ecological conditions governing their existence. Some were hardy and there were few limits to their movements, others were slower and more sensitive and more hedged in by their environment. After these ancient lands had become populated, a series of vacilla- tions occurred and the contiguous masses were separated. New- foundland, which had formerly been a part of the ancient land includ- ing the maritime provinces of Canada and New England, was separated from the mainland during the late pleistocene and the Straits of Belle Isle were formed. This stretch of water was an effective barrier to any further dispersal of molluscan land life and probably was, later, an effective barrier to the glaciers. Even during the greatest glacia- tion it is probable that the island was not completely covered.^ Cer- ^Recently, in letters to the author, Dr. Carl O. Dunbar, Curator of Inverte- brate Paleontology in the Peabody Museum, Yale University, and an authority on the glacial history of Newfoundland, has proven beyond doubt that the Long Range was, at one time during the Pleistocene, completely glaciated. Fernald believes that, amid the ice and snow, there were islands which gave shelter to the present flora. To me it is obvious that the present distribution of the land snails indicates that they did live through the vicissitudes of glaciation and the climatic changes existing after the separation of Newfoundland from the mainland. A complete glaciation of the entire island of Newfoundland, after its separation from the mainland, is precluded by the presence of the many species necessarily spread by a migration from the mainland. No human agency, no flight of birds, no importa- tion of food-stuffs by the Red Indians or Beothucks, could explain the present population of the land snails, many of which are so minute as to escape notice by any but the specialist studying them. From my standpoint it is then necessary to explain the evidence as follows. Following the complete glaciation of Newfound- land and the probable destruction of the fauna of that region, the warm interglacial 1936 Brooks: Mollusca of Newfoundland 87 tain peaks, certain areas, rose above the ice and formed habitable “glacial islands” upon which the plants and animals were enabled to exist through this hazardous period. These islands are indicated by the presence of a great number of plants (Fernald, M. L. cf Bibliog- raphy), and the present day, endemic molluscan fauna. With the dissipation of the ice the animals and plants again populated the sur- rounding areas and the present distribution is the result. It is much too early in the study, or one might better say, in the collection of the Newfoundland fauna, to indicate whether or not any area outside of that of the Long Range was unglaciated. At first glance the accompanying chart suggests an eastern concentration of species. Disregarding the ones introduced through commerce, we have only one species of land snail which is not found either in the central or in the western portion of the island. The relatively few collections enable this to be explained as merely an oversight. So, from the study of the land shells, there is no reason to believe other- wise than that the natural dispersal of forms following the recession of the ice accounts for the present distribution. The very large number of species occurring on the west coast not only indicates their parent region but indicates also the greater number of collecting stations. The eastern part of Newfoundland has been only sparsely collected. The water forms included here have a much more doubtful history. I believe that the majority of the forms listed are the relicts of an earlier distribution, probably contemporaneous with the land snails. This seems to be easily acceptable in view of the present distribution of Margaritana margaritifera, Stagnicola palustris and varieties, Fossaria ohrussa and forms, the doubtful (in distribution) Radix pereger geiseri- period allowed a re-population of Newfoundland from the mainland. The formation and flooding of the Bay of St. Lawrence followed, isolating the fauna and flora of this new island. Subsequent glaciations were local and sporadic, allowing refuge for the hardy northern species. 1 believe that this explanation, weak though it is, will be upheld by future studies of the marl deposits and other subfossil-bearing beds of Newfoundland. If not, and if the glacial studies carried on by Dr. Dunbar and his colleagues still conspire to wipe out the fauna and flora by a complete glaciation after the formation of the Bay of St. Lawrence, it will truly seem that the fauna of Newfoundland has found some means of dispersal yet unknown to the “distri- butionalist,” and yet to be illuminated through observation and research. If, however, the land snails did come to Newfoundland through outside agencies, it was comparatively simple for them to attain their present distribution throughout Newfoundland during the many thousands of years that have followed. 88 Annals of the Carnegie Museum VOL. XXV cola, and the members of Gyrauliis, Physa, Valvata, and Amnicola. They are, in other words, a parcel of the general distribution populat- ing the early North American continent. With my sparse collections no more explicit explanation may be made. It is, however, extremely doubtful that any distribution of water forms into Newfoundland, since the ice age, has occurred. In the event of such an invasion, the entire island would necessarily have been surrounded with fresh water as the majority of the streams bearing these forms empty along the northern coast and are shut off from the mainland by the extensive northern peninsula. LIST OF SPECIES Eamily HELICID^ Helix hortensis Muller This common species so exhaustively treated in the past is, as one can see from the following list of localities, found mainly on the west coast of Newfoundland. In the Carnegie Museum it is also repre- sented by specimens from Terra Nova (collected by Mrs. G. H. Durgin in the George H. Clapp collection) and from Shoal Harbor, both localities on the east coast. From questioning fellow travelers on the train I found that it was the common form to be found in their gardens (also eastern). From my collections it seems that this species will be found to be distributed generally across the northern portion of the island, although there is a very slight possibility that the col- lections at Shoal Harbor may point to an area which the glaciation left unharmed and which is thus supporting its own relict fauna. The locality at which I made the collection at Shoal Harbor is between six and eight miles inland from the town, on the slopes of a hill surrounded by spruce forests, dense undergrowth of alders, etc., and acres of boggy land. There is no possibility of this colony having been planted either intentionally or accidentally due to its inaccessibility. It is also pecu- liar that none of the townspeople, except my guide, Mr. Leslie Tuck, had seen one of these shells in the vicinity. I found them only after several discouraging days when Mr. Tuck finally remembered having seen “shells” at that place several years previous. The specimens at hand, with one exception, are banded. The Shoal Harbor ones are typically banded and dark; the ones from Lomond, Bonne Pay, show nearly a complete coalescence of the bands 1936 Brooks: Mollusca of Newfoundland 89 (123) (45), as do those from Gros Morne Mountain, Long Range, Bonne Bay. In the latter the bands 4 and 5 are separate but may or may not be joined at the aperture. The one specimen labelled “Bay of Islands” has distinct bands but shows the same tendency toward fusion. Three of the shells from Terra Nova are bandless while four have the formula (12345) and one the formula 1(1234)5. Some specimens received only recently from Dr. Carl O. Dunbar, Curator of Invertebrate Paleontology, Peabody Museum of Natural History, Yale University, show the following characteristics: three specimens from the mouth of Romaine Brook, 2 miles east of Port au Port, are all typically banded; 12345. Four specimens from the dune sand between Port Saunders and Pointe Riche differ from the others; one is bandless and exhibits the typical canary yellow color while the others are respectively 12345, (123) (45), and (12)3(45). From my own findings and the findings of others there seems to be no reason to believe that this species came to Newfoundland and eastern America through any agency of man. They have, like the majority of forms taken up here, come into these regions through the natural growth of a species out from its center of origin. In this case, due to the destruction of the ancient distributional channels, we must accept Europe as the center of their dispersal. In view of the scanty area occupied in North America I feel that here we have the outer distributional fringe, or “dispersal fringe,” demarked by their presence. Helix hortensis, in contrast with the more hardy Cochlicopa lubrica, is an example of the slow and more sensitive migrant. The distribution of this species in North America readily shows that it is hampered in its progress by temperature. The rigorous winters of our inland regions prevent it from spreading very far from the tempered climate of the sea coast. I would not say that this is the only barrier to this group, but it seems the obvious one in view of the fact of the greater dispersal of other circumboreal forms. This condition has been indicated in the writings of C. W. Johnson (1906), O. O. Nylander (1908), W. W. Winkley (1904), and others. Attempts to start colonies of this species in the comparative mildness of the Pittsburgh (Penn- sylvania) region by Dr. George H. Clapp and the author have all ended in failure. Dr. Clapp believes that rodents are the main cause of their disappearance although I have been successful in keeping a colony alive and fairly intact without protection until cold weather. You will perhaps think of the cold winters of Newfoundland and the 90 Annals of the Carnegie Museum VOL. XXV St. Lawrence valley. It is true that the populated areas have cold winters, but it is also true that early and deep snowfalls blanket the hills and valleys until the mild days of spring thus protecting the ani- mals from extremes of temperature. In our region (Pittsburgh) very little snow falls to protect them from the frequent freezes. Western Stations: Bonne Bay; Tuckers Head, Lord and Lady Cove, Killdevil Mountain, Main Arm, Southern Arm, Deer Arm, Beach at Lomond, Gros Morne Mountain, (A. N. S. P. and C. M.). Bay of Islands; Hannah’s Head, French Island, Lark Island, “Bay of Is- lands.” Straits of Belle Isle; Doctor Hill, Bard Harbor Hill, High- land of St. John, St. John’s Bay (A. N. S. P.). West Coast and south; Robinson’s River, Serpentine River, Little Codroy and Great Codroy Valley, East River, Hawkes Bay, Ingornachoix Bay at Pointe Riche (A. N. S. P. and C. M.), Lewis Hills near Port au Port (M. C. Z.). Easteryi Stations: Shoal Harbor, Terra Nova (C. M.). Helicigona arbustorum (Linnaeus) This species was reported by J. F. Whiteaves (1863) from St. Johns. The specimens were found on the “grassy slopes facing the sea, near the narrows of St. Johns’ Harbor” by Robert Bell in July, 1885. I was over this area in June, 1934, but found nothing. As indicated in the note this is probably an introduced form as it was found near the scene of several wrecks. A patch of heather is nearby and it is thought that this was from the discarded stuffing of bedding brought by the immigrants; they discarded the bedding when it become spoiled from moisture during their escape from their wrecked vessel. Family ZONITIDZE Zonitoides arborea (Say) This species, common to Asia and America, has a wide distribution on this island, occurring from the eastern coast to the Long Range and north to the very tip of the peninsula. Its wide distribution in North America is well known and from its appearance in Newfoundland it seems to have populated this area at an early date. Although it is a common form it is never collected in great numbers. At Grand Falls it occurred with Cochlicopa lubrica under damp logs near pools along a wood road. Back from the river it was not found, due, perhaps, to 1936 Brooks: Mollusca of Newfoundland 91 the acid bogs although the “flies” made the search for it in these sheltered places a veritable nightmare. Western Stations: Bonne Bay; Gros Morne Mountain, Beechy Point, Stanleyville, Lomond. Bay of Islands; Hannah’s Head. Straits of Belle Isle; Bard Harbor Hill, Doctor Hill, Burnt Cape at Pistolet Bay, Pointe Riche. (A. N. S. P. and C. M.). Central Stations: Banks of the Exploits River at Grand Falls (C. M.) Eastern Stations: Shoal Harbor (C. M.). Hawaii minuscula (Binney) A new record for Newfoundland and a common circumpolar species of Asia and America. This species was found in the siftings of forest loam from Lorhond, Bonne Bay, west coast. Striatura exigua (Stimpson) This species was collected at Lumber Camp No. 31, near Lomond, and at Gros Morne Mountain, Bonne Bay. It is new to the fauna of this island but in America it is found from Maine and Quebec west to the Rocky Mountains. Striatura milium (Morse) A new record for Newfoundland and collected at Shoal Harbor on the eastern coast. Euconulus fulvus (Muller) This species, common to three continents, is wide spread in its distribution in Newfoundland. Western Stations: Bonne Bay; Tucker’s Head. Bay of Islands; Hannah’s Head. Straits of Belle Isle; Cape Norman, Savage Cove, Ha Ha Cape, Bard Harbor Hill, Doctor Hill, Schooner Island, Anse aux Sauvages (A. N. S. P.). Central Stations: Red Indian Lake and Banks of the Exploits River, Grand Falls (C. M.). Eastern Stations: Shoal Harbor (C. M.). 92 Annals of the Carnegie Museum VOL. XXV Retinella electrina (Gould) This typically American species is found from the eastern coast of Newfoundland to Alaska. Western Stations: Bonne Bay; Tucker’s Head, Lomond. Bay of Islands; French Island, Middle Arm, Hannah’s Head. West coast and north; Brig Bay, Schooner Island, Anse aux Sauvages, Cape Norman, Ha Ha Cape, Bard Harbor Hill, Penguin Head (A. N. S. P. and C. M.). Central Statiojis: Banks of Exploits River, Grand Falls (C. M.). Eastern Stations: Shoal Harbor (C. M.). Vitrina limpida Gould So far, this species has been found only on the west coast (A. N. S. P.) near Pointe Riche. In America it is a species confined to the northeastern area. Its most southern range is along the Ohio River in Allegheny County, Pennsylvania. It had been distributed by floods originating in western New York state where it is a fairly common species. Family LIMACIDT: Deroceras agrestris (Linnaeus) The consensus of opinion would place this species among those of the introduced forms. However, in Newfoundland it has been col- lected from two widely separated localities; Bard Harbor Hill and Doctor Hill (A. N. S. P.) on the west coast and on the banks of Manuels River at Manuels, near St. Johns (C. M.). Deroceras laeve (Muller) This species dilTers from the above in being an endemic form. It occurs in both Europe and America and in central and western New- foundland. It is undoubtedly a species of the “original distribution.’’ More collections will greatly extend its range on this island. Western Stations: Mauve Bay near Cape Bauld, Lark Harbor, Savage Cove, Ha Ha Cape, Yankee Point, Flower’s Cove, Doctor Hill (A. N. S. P.). Central Stations: Banks of the Exploits River, Grand Falls (C. M.). 1936 Brooks: Mollusca of Newfoundland 93 Family ENDODONTID^ ^ Helicodiscus parallelus (Say) This species is distributed over the entire eastern United States and southern Canada. In Newfoundland, it has been found on the west coast at Hannah’s Head (A. N. S. P.), Lomond (C. M.), and in the central area at Grand Falls on the banks of the Exploits River (C. M.). Punctum pygmaeum minutissimum (Lea) In the United States, this species has a range from Maine south to Virginia and west to Arizona. In Newfoundland, it has been col- lected at Tucker’s Head, Hannah’s Head, Bard Harbor Hill on the west coast (A. N. S. P.) and from Shoal Harbor on the east (C. M.). Gonyodiscus cronkhitei (Newcomb) This form of western America is wide spread in Newfoundland. No records at hand show that it has been collected at any place between Newfoundland and the Rocky Mountains. In the collections of the A. N. S. P., it is listed from Tucker’s Head, Hannah’s Head, Anse aux Sauvages, Yankee Point, and in our collections from Beechy Point, Lomond, and Gros Morne Mountain on the west coast, at Grand Falls in the central region, and from Shoal Harbor on the eastern seaboard. Gonyodiscus cronkhitei anthonyi (Pilsbry) Although my collections contained none of this form it has been collected (A. N. S. P.) in the western part of the island; Bard Harbor Hill, Savage Cove, Brig Bay, Pointe Riche, Cape Norman, Ha Ha Cape, Grassy Island, St. John’s Bay and Doctor Hill. Family ARIONIDT: Arion ater (Linnaeus) Undoubtedly an introduced form, collected at Bay Bulls, east coast (A. N. S. P.). It has been found only twice before on the American Continent. 94 Annals of the Carnegie Museum VOL. XXV • Arion hortensis Ferrusac Another introduced species of a very localized distribution in America. Collected in the western part of Newfoundland at Beechy Point, near Lomond, Bonne Bay (C. M.), Lark Harbor, Bay of Is- lands (A. N. S. P.), and on the eastern side at Manuels on the banks of the Manuels River (C. M.). Arion fasciatus Nils Introduced: Reported from Trepassy and Whitbourne (A. N. S. P.), both in the eastern part of the island. Family SUCCINEID.F: Succinea ovalis Say Western Stations: Lark Harbor, Savage Cove, Flower’s Cove, Eddies Cove, St. John’s Island, Bard Harbor Hill, Doctor Hill. Bonne Bay; Tucker’s Head (A. N. S. P.), Lomond, Camp 31, Stanleyville, Beechy Point, Gros Morne Mountain, “Bay of Islands’’ (C. M.). Deer Lake, Nicholsville, Hawkes Bay (M. C. Z.). Baleana Bay (No. 180190 U. S. N. M.). Central Stations: Banks of the Exploits River at Grand Falls (C. M.). Eastern Stations: Shoal Harbor and Terra Nova (C. M.). Succinea avara Say Western Stations: Sandy Cove (Bay?), Cape Norman, Yankee Point, Four Mile Cove, Big Brook, St. John’s Island, Flower’s Cove, Savage Cove, Savage Point, Pointe Riche, Brig Bay. Bonne Bay; Tucker’s Head (A. N. S. P.), Lomond (C. M.). Succinea verrilli Bland Anchor Cove, Flower’s Cove, Bard Harbor Hill, St. John’s Island, Old Port au Choix, St. John’s Bay, Main River (A. N. S. P.), all west coast. Succinea groenlandica (Beck) M oiler A surprising species to find and a new record for Newfoundland. Collected at Lomond, Bonne Bay (C. M.). 1936 Brooks :’Mollusca of Newfoundland 95 Succinea peoriensis “Wolf” Walker Another new record from Lomond Depot, Camp 31, and Stanley- ville, all on western coast (C. M.). Dr. Rehder says that these are not all typical of the species and somewhat “similar to S. peoriensis fultonensis F. C. Baker.” Family PUPILLID^ Pupilla muscorum (Linnaeus) Western Stations: Bonne Bay; Tucker’s Head. Straits of Belle Isle and north; Doctor Hill, Pointe Riche, Ha Ha Cape, Ha Ha Bay, Cape Norman, Anse aux Sauvages (A. N. S. P.). Vertigo modesta (Say) Western Stations: Tucker’s Head, Pointe Riche, Anse aux Sauvages, Great (?) Sacred Island, Savage Cove, “Sandy Cove = Poverty Cove” = Sandy Bay?, Bard Harbor Hill, Doctor Hill, Flower’s Cove, Ha Ha Cape, Brig Bay, Yankee Point, Savage Point; there are two by this name, this may be the more northern one in Cremaillere Harbor and not the Savage Point at Great Goose Harbor (A. N. S. P.). Vertigo modesta parietalis Ancey Western Stations: Anse aux Sauvages, Great Sacred Island, Ha Ha Cape (A. N. S. P.). Vertigo modesta castanea Sterki Western Stations: Schooner Island, Pistolet Bay, Cape Norman, Flower’s Cove (A. N. S. P.). Vertigo gouldi paradoxa Sterki Western Stations: Bonne Bay; Tucker’s Head. Bay of Islands; Hannah’s Head. Straits of Belle Isle; Ha Ha Cape, Bard Harbor Hill (A. N. S. P.). Mr. Vanatta writes that V. coloradensis and V. coloradensis hasidens are now considered as the above species. 96 Annals of the Carnegie Museum VOL. XXV Vertigo elatior Sterki Schooner Island (A. N. S. P.) and Lomond, Bonne Bay (C. M.), both on the west coast. Columella edentula (Draparnaud) Western Stations: Anse aux Sauvages, Sacred Island, Schooner Island, Savage Point, Yankee Point, Ha Ha Cape, Doctor Hill. Bonne Bay; Tucker’s Head (A. N. S. P.). Family COCHLICOPID^ Cochlicopa lubrica (Muller) This species is probably the most common of any of the shelled snails in Newfoundland. It is surprising not to find it in the eastern part of the island. The absence of the species at Shoal Harbor is probably due entirely to oversight. Western Stations: Bonne Bay; Beechy Point, Stanleyville, Lomond (C. M.), Tucker’s Head. Bay of Islands; Hannah’s Head, Tweed Island, Penguin Island (Bay of Islands?). Straits of Belle Isle; Schooner Island, Ha Ha Cape, Eddies Cove, Eddies Cove Brook, Bard Harbor Hill, Doctor Hill (A. N. S. P.). Central Stations: Banks of the Exploits River, Grand Falls (C. M.). Family VALLONIIDT: Vallonia albula Sterki From the records of the A. N. S. P. ; Tucker’s Head in Bonne Bay, Penguin Head, and at Pointe Riche on the northwestern coast. Zoogenites harpa Morse From the records of the A. N. S. P. ; Ship Cove, Sacred Bay, Anse aux Sauvages, Deer Pond Brook in Bard Harbor, and collected by me at Shoal Harbor. Planogyra asteriscus Morse A new record for this species; collected at Lomond (C. M.). 1936 Brooks: Mollusca of Newfoundland 97 Family CARYCHIID^ Carychium exiguum (Say) A new record from Lomond (C. M.). The Aquatic Gastropoda Family LYMN^ID^ Stagnicola palustris (Muller) The following collections from the records of the A. N. S. P. may be found to agree* with one of the varieties described by Dr. Baker (1935)- Until this has been determined I shall list the localities separately. Western Stations: Straits of Belle Isle; Cook’s Point, Pistolet Bay, Schooner Island, Sandy Cove, Flower’s Cove, Otter Pond near Plumb Point, Brig Bay, Eddies Cove, Boat Harbor, Big Brook, Savage Point, St. John’s Island and Bay of St. John. Stagnicola palustris papyracea Baker and Brooks This new variety was collected in Rocky Pond, near Whitbourne. The descriptions as published in The Nautilus (1935) will be repeated here for convenience sake. “Shell differing from S. p. ungava F. C. Baker in being more ovate with spire and aperture about equal in length, the whorls of the spire not as high as those of ungava, the sutures less deeply indented; inner lip narrow, the columellar plait quite distinct and the umbilical chink small or absent; color light horn, interior of aperture light or dark chocolate color; sculpture of finer lines than in ungava. All apices are decollate and but four whorls remain, but there are ap- parently six whorls in mature examples. Length 18.6 diam. 10.5 aper. length 10 . 5 width 5.7 mm. Holotype ” 18. 1 ” 10. o ” ” 10.5 ” 5.7 ” Paratype ” 17. 1 ” 9.9 ” ” 9.5 ” 5.1 ” This race of palustris at once suggests S. p. ungava, differing in the shorter spire, narrower inner lip and distinct columellar plait. It differs from S. p. elodes in its shorter, less acute and obese body whorl. The shells are very thin which suggests the varietal name. The race is very constant in form and does not appear to vary to any extent toward typical palustris or its known varieties.’’ *Mr. Vanatta writes: “Could call all Lymnaea palustris from Newfoundland, var. newfoundlandensis B. and B.’’ 98 Annals of the Carnegie Museum VOL. XXV Stagnicola palustris perpalustris Baker and Brooks A new variety found inhabiting the pools along the Exploits River at Grand Falls. “Shell obese, with large ovate aperture and short, dome-shaped spire, the latter shorter than the aperture; sutures well impressed; outer lip thin with only a slight variceal thickening within the edge; inner lip rather broad, appressed tightly to the columella leaving only a slight umbilical chink; columella plait heavy, parietal wall with distinct callus; color dark horn, interior of aperture dark chocolate colored; sculpture of heavy growth lines and impressed spiral lines; surface often malleated; only four whorls visible in mature shells, all spires decollated. Young shells indicate that there are probably six whorls in fully matured shells. Length 19. i diam. 11.3 aper. length 12 . o width 6.6 mm. Holotype ” 17.6 ” 10.4 ” ” ii.i ” 6.0 ” Paratype ” 17. I ” 10.5 ” ” 10. 5 ” 5.8 ” This race of palustris is at once recognized by its broad form, de- pressed spire and large aperture. It does not closely resemble any of the described forms of this protean species, approaching most closely to some short spired forms of S. p. nuttaliana from the Rocky Moun- tain area.” Stagnicola newfoundlandensis Baker and Brooks This species inhabited the “steady” near camp 31, eight miles from Lomond, Bonne Bay. “Shell elongated with acute somewhat turreted spire as long as or longer than the aperture; spire whorls rounded with well impressed sutures; body whorl well rounded; aperture ovate, outer lip thin, inner lip flattened and reflected over the umbilical region leaving a small chink; the callus on the parietal wall is thin or absent; columellar plait absent or but slightly developed; color dark horn, aperture coffee colored within; sculpture of coarse growth lines and well developed spiral lines; there are six whorls. Length 20 . 0 diam. 9.9 aper. length 10. 0 width 5.3 mm. Holotype ” 22.0 ” 10. 0 ” ” 10. 0 5.1 ” Paratype ” 18.8 9.2 ” ” 9.0 4.7 ” ” 17.0 ” 90 ” 9.2 5.1 ” This lymnaeid resembles some of the elongate forms of the Stag- nicola emarginata complex, especially canadensis and ontariensis. The color of the shell and aperture are different from canadensis and the inner lip is not turned back to form so flat a projection over 1936 Brooks: Mollusca of Newfoundland 99 the umbilical region. Compared with specimens of ontariensis from the St. Lawrence River below Quebec the shell is more elongated with longer, narrower, more acute spire, the inner lip is not flattened and the color is much darker. A few specimens of the new form resemble certain forms of S. p. elodes but the typical forms have a different and heavier sculpture, a more rounded body whorl, a more acute spire, and the columellar lip is wider at the lower part and lacks the heavy, twisted plait of typical elodes. There is a great A^ariation in the height of spire and width of shell but the greatest number of specimens are uniform and it seems best to recognize this form as a species distinct from either the palustris or emarginata complex.” Fossaria obrussa (Say) This species was collected at the same locality as the above. The range of this form as given by Baker (1911) is “from the Atlantic to the Pacific Ocean and from Mackenzie Territory and Quebec south to Arizona and northern Mexico,” so it is not surprising that this species is found in Newfoundland. However, this is the first time that this species has been recorded from there. It was collected on exposed sticks and stones and in the surface film covering deep muck of vegetable origin, near the “steady” below the “little pond.” Within a short distance there was a large bed of Margaritana margaritifera, and in the shoal waters were the Helisoma and Gyraiilus mentioned below. It was unusual to find areas supporting as many forms as did this one. Fossaria obrussa brooksi F. C. Baker The type locality for this form is the same as that of newfotmd- landensis. These were collected from the surface of the mud around the swampy areas of the “steady.” As near as I could determine the term “steady” applies to any enlarged, lake-like portion of the creek or river. This also happened to be one of the favorite “salmon holes” of the region. “Shell differing from Fossaria obrussa decampi in having a longer, more acute and turreted spire, a shorter, more obese body whorl, a smaller, rounded aperture and a larger umbilicus; whorls shouldered; color light or dark horn, the aperture chocolate or coffee colored within; there are six whorls. Length 11.5 diam. 6.0 aper. length 5 . i width 2 . 9 mm. Holotype ” 91 4-7 ” 4.6 ” 2.2 ” Paratype 8.4 4.0 ” 3-8 ” 2.0 ” ” 100 Annals of the Carnegie Museum VOL. XXV This distinct little lymnaeid is related to obrussa, approaching most nearly to the race decampi. Its long scalariform spire, short, rounded body whorl, and small rounded aperture, will at once distinguish it from decampi." Fossaria umbilicata (C. B. Adams) Baker’s (1911) intimation that this form was an inhabitant of the more northern regions (“of the Canadian and Nova Scotian regions”), finds its proof in the discovery of this species in Newfoundland. It was collected along the logging road to Deer Lake not far from Lom- ond, Bonne Bay. It was found on damp sticks and stones in the marshy drainage of a roadside spring. Radix pereger geisericola (Beck) This species was reported from Junction Pond, Whitbourne, by Vanatta. My specimens came from Rocky Pond which is only a few minutes down the railroad tracks from the former locality. This species was originally described as living in “the hot waters of the geysers of Iceland (vide Morch)”, according to Baker (l.c.), and it is indeed a mystery how they came to the cold Newfoundland lakes. Family PLANORBID^ Gyraulus parvus (Say) Vanatta lists this species as occurring on the west coast; near the Straits of Belle Isle, Otter Pond near Plumb Point, East Brook (? East River on my map), at St. Barbe Bay, and Eddies Cove Brook in Eddies Cove. Gyraulus hornensis F. C. Baker Collected in the “steady” near camp 31. This species has only recently been described from Canada, and as Dr. Baker says, “This is a long way from home but is certainly this species.” I shall repeat the description as he gives it (1934). “Shell depressed, the periphery rounded; color light corneous, surface shining; sculpture of fine oblique lines of growth and with very fine spiral lines, more or less conspicuous; nuclear whorls small, rounded, spirally striate in sculpture; whorls about four, rapidly en- 1936 Brooks: Mollusca of Newfoundland 101 larging, the last somewhat expanded near the aperture, roundly angled at the periphery of the last whorl, the upper part of the body whorl slightly flattened; spire flat, the whorls coiled in the same plane; the body whorl may be nearly in line with the spiral turns or it may be deflected about a third of the distance from the aperture; sutures deeply channeled; base concave exhibiting all of the whorls, the umbilical region wide, but the body whorl well rounded, not flattened or having a reamed-out appearance; aperture obliquely, ovately rounded; lip thin, sharp, simple, or slightly thickened with a callus deposit; parietal wall with a wide callus.” Greatest height 2.0, greatest diameter 4.6 mm. Described from Birch Lake, Horn River, 75 miles^above the Mackenzie River, Mac- kenzie District, Canada. Collected by E. J. Whittaker in 1921. This species ranges from Wisconsin and North Dakota north to the type locality and east to Newfoundland. Helisoma campanulata davisii (Winslow) Collected along the road near camp 31, Lomond, Bonne Bay. This species was described by Miss Minna Winslow (1926) from the Pinnebog River, Huron County, Michigan. This variety dilTers, according to Baker from rudentis (Dali) “in its -smaller size, much less axial height, and particularly in the form of the umbilicus, which in rudentis is distinctly cone-shaped or funnel-shaped.” This is the first record of this variety from so easterly a region. Is it a case of distribution by natural means or is it a convergent development due to the similarity of habitat and climate? The latter seems the most feasible. Helisoma campanulata minor (Dunker) This form of campanulata was collected in Stony Brook, across the Exploits River at Grand Ealls. It is figured and commented upon by Miss Winslow (l.c.), from Crooked Lake, Montmorency County, Michigan. Family PHYSID^ Physa gyrina Say Mr. Vanatta lists this from “Otter Pond, near Plumb Point” and from “Brig Bay,” west coast. 102 Annals of the Carnegie Museum vol. xxv Physa heterostropha Say In the collections of the Museum of Comparative Zoology from Nicholsville, Humber River, and in my collection from camp 31, Lomond, Bonne Bay, on the west coast. In the central area I col- lected it at Stony Brook, across the Exploits River from Grand Falls, and in pools along Rushy Pond Road at Grand Falls. Family VALVATID^ Valvata lewisii Currier Vanatta lists this from Eddies Cove Brook, west coast, and I col- lected it from the “steady,” camp 31, Lomond, Bonne Bay. Valvata sincera nylanderi Dali Collected by me in Rocky Pond, VVhitbourne, in the eastern area. Family AMNICOLID/E Amnicola limosa porata Say Or near this species. Collected by me in Rocky Pond, Whit- bourne, and in Stony Brook, across the Exploits River from Grand Falls. The Fresh Water Pelycypoda Margaritana margaritifera (Linn sens) Southwest River, Belvoir Bay, in Hare Bay (C. M.). Southwest River (M. C. Z.), probably same locality; both southwestern. Dr. Bryant Walker (1910) records it from St. Barbe Island, Birchy Brook and Sandy Pond; probably western. Vanatta lists it from Junction Pond, Whitbourne; eastern, and I collected the same from “Chitman’s Water,” and Rocky Pond, Whitbourne; both probably of the same drainage, and from the Exploits River, Grand Falls; central. I also found it near camp 31, Lomond, Bonne Bay. Dr. Bartsch writes that they have two sets. Nos. 86292 and 86288, U. S. N. M., labelled “Newfoundland.” From the above stations we can see that this species is wide-spread over the island, occurring from the eastern portion at Whitbourne 1936 Brooks: Mollusca of Newfoundland 103 west to Lomond. It will probably be found to be distributed over the entire island. Various Newfoundlanders, that I talked to, told me of taking “pussels” during the spring and boiling and eating them. I questioned them to determine whether these were the fresh water forms, and they assured me that they were. I presumed that they were speaking of this species; however, I marvelled at their gastronomic prowess. Anodonta marginata Say Reported by Vanatta from Otter Pond near Plumb Point, and from Junction Pond, Whitbourne. Anodonta implicata Say Collected by the son of Peter Pettipas for me at Whitbourne in “Well’s Gully.’’ This is the first record of this species from New- foundland. Its presence here supports Whiteaves’ findings (1863) in the St. Lawrence River, and Marshall’s findings (1895) in the Ottawa River, both findings questioned by Ortmann in his catalogue. BIBLIOGRAPHY Baker, F. C. 1911. Lymnseidse of North and Middle America, Recent and Fossil. Chicago Academy of Sciences, Special Publication No. 3, pp. 1-539. 1928. The Fresh Water Mollusca of Wisconsin, Part 1. Wis- consin Academy of Science, Arts and Letters, pp. 345-350. 1934. A New Species of Gyraulus from Canada. The Canadian Field Naturalist, Vol. 48, p. 135. Baker, F. C., and Brooks, S. T. 1935- New Species and Races of Lymnaeidse from Newfoundland. The Nautilus, Vol. 49, No. i, pp. 10-13. Binney, Amos. 1851. The Terrestrial Air-breathing Molluscs of the United States, Part II, p. 112. (H. hortensis in St. Pierre). 104 Annals of the Carnegie Museum VOL. XXV Clapp, George H. 1900. Helix hortensis in Newfoundland. The Nautilus, Vol. 14, p. 72. 1907. Helix hortensis on Magdalen Island. The Nautilus, Vol. 20, p. 105. Clench, W. J. 1929. Arion ater ater (Linne) in Maine. The Nautilus, Vol. 42, p. 104. Cockerell, T. D. A. 1890. Helix hortensis in America. The Nautilus, Vol. 3, pp. 139-140. 1907. Helix hortensis in Newfoundland. The Nautilus, Vol. 20, p. 94. Dall, W. H. 1905. The Harriman Alaskan Expedition. Doubleday, Page and Company, New York, pp. 1-171. Fernald, M. L. 1926-27. Two Summers of Botanizing in Newfoundland. Rho- dora, Vol. 28, pp. 49-241. 1930. Unglaciated Western Newfoundland. Harvard Alumni Bulletin for January. Henderson, Junius. 1930. Newfoundland a Promising Conchological Field. The Nautilus, Vol. 44, pp. 9-10. Hooker, Joseph D. i860. Outlines of the Distribution of Arctic Plants. Transac- tions of the Linnean Society of London, Vol. 23, pp. 251-348. Johnson, C. W. 1906. On the Distribution of Helix hortensis Miill., in North America. The Nautilus, Vol. 20, p. 73. 1908. A Summary of Robert Bell’s Paper from the Canadian Naturalist and Geologist (Vol. 4, 1859, p. 215), The Nautilus, Vol. 21, pp. 130-131- Marshall, W. B. 1895. The Geographical Distribution of New York Unionidae. 48th Annual Report of the New York State Museum, pp. 47-99. 1936 Brooks: Mollusca of Newfoundland 105 Nylander, O. O. 1908. A Note on Helix hortensis. The Nautilus, Vol. 22, pp. 30-31- Vanatta, E. G. 1925. Newfoundland Shells. The Nautilus, Vol. 38, p. 92. 1927. Land and Freshwater Shells from Newfoundland. The Nautilus, Vol. 40, p. 113. 1933. Newfoundland Shells. The Nautilus, Vol. 43, p. 133. Walker, Bryant. 1910. The Distribution of Margaritana margaritifera (Linne) in North America. Proceedings of the Malacological Society of London, Vol. 9, pp. 126-145. 1918. Foreign Land Snails in Michigan. Occasional Papers of the Museum of Michigan, No. 58, pp. 1-3. Whiteaves, J. F. 1863. On the Land and Fresh-water Mollusca of Lower Canada. The Canadian Naturalist and Geologist, Vol. 8, pp. 98-107. WiNKLEY, Henry W. 1904. Helix hortensis in New England. The Nautilus, Vol. 17, pp. 121-122. Winslow, Minna. 1926. The Varieties of Planorbis campanulatus Say. Occasional Papers of the Museum of the University of Michigan, No. 180, pp. 1-9. 106 Annals of the Carnegie Museum VOL. XXV EXPLANATION OF PLATE XII 1. Stagnicola newfoundlandensis Baker and Brooks. 2. Fossaria ohrussa brooksi F. C. Baker. 3. Stagnicola palustris perpalustris Baker and Brooks. 4. Stagnicola palustris papyracea Baker and Brooks. Photographed on millimeter ruler. ANNALS CARNEGIE MUSEUM, Vol. XXV. Plate XII. 108 Annals of the Carnegie Museum VOL. XXV EXPLANATION OF PLATE XIII Map showing the Polar Regions and the submerged (stippled) continental shelf and shallows; the depths cross-lined. It is evident from the studies of Fernald and others that this circumpolar area was not only a “pathway” but the dispersal area for many of our present plants and animals of northern affinities. This back- door “Garden of Eden” has probably been utilized from the Carboniferous to late Pleistocene time, and during the long and warm interglacial periods. ANNALS CARNEGIE MUSEUM, Vol. XXV. Plate XIIL REMARKS ON A SKULL CAP OF THE GENUS TROODON f? - :■ ■ By Charles W. Gilmore Rejirinted from the Annals of the Carnegie Museum f f Vol. XXV, p. 109-112, 1936 L ■ , ■ ■■ - ■, ('■ ' Issued June 20, 1936 .y- )'-: , : 'x;-; - --.'V-,,-4:v-^v^S''^:' \' ■ ■ . ^ ■ t;.^ '-rv'- 7:''. ■'i ;> ' V A / . '■ i . ■■ ■ A -A^tN I- A"^ ' -A v'AV>- I' J -A pm '■’"r ^ < -?V '"A. A' 'AA?'> - - • A- A ■' ■ r"\ ivAAA-'''' . . ■ ‘ - ■- AAA..'^“- ' '-.V . ■'i^m ‘ . -^ ‘ •, . ^• ••- *'• '■^' •- . 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^•' ...xXY yX X rXv>^,i;7 XX * s ^ V’ A Y""-7AAb 1 ,7Xb XX 7a 1 . ; : " y-' v>> s ' ■ A.:'.-,- Y-,fe • -:'.. , ■• 7.- :. ' ' Xa YXX'-Y YXyXX ■V ' : A, •■- Y'' Y7X:-X ■.■■■ -;, 7.7., YA Y'-Y X A ■ yXy a a YrAxA .AA 'YAJk •Y: ■• f ■X-A^^X / ■ A- -7/ 7A" 7 'S',. 'v.'’.-YiS'.'',.i' ART. XI. REMARKS ON A SKULL CAP OF THE GENUS TROODON By Charles W. Gilmore Curator of Vertebrate Paleontology U. S. National Museum An incomplete skull in the paleontological collections of the Carnegie Museum is of interest in being the second recognizable specimen found of Troddon wyomingensis Gilmore.^ Since the entire top of the thickened skull cap is preserved, much of which was missing in the type, it contributes to a better under- standing of this curious, but little known dinosaur. This specimen was collected by the late W. H. Utterbeck, but unfortunately it is without data as to exact locality, though pre- sumably from the Lance formation in Wyoming. It consists of the greater portion of the dome-like part of the skull, see fig. i, and dis- plays for the first time the precise shape of the top of the head in this species. Fig. I. Skull of Troddon wyomingensis Gilmore, Cat. No. 3180 C. M. Viewed from the left side, P.f., postfrontal; Sq, squamosal. One-fourth natural size. The occipital border, and also the left side anterior to the top of the infratemporal fossa, is essentially complete. Anteriorly, immediately 'Gilmore, Charles W., Proc. U. S. Nat. Mus., vol. 79, 1931, Art. 9, pp. 1-4. 109 Issued June 20, 1936. MAR 2 5 no Annals of the Carnegie Museum VOL. XXV posterior to the junction of the frontals with the nasal bones, the skull is abruptly broken. The specimen differs from the type, with which it has been directly compared, in being a fourth smaller, in having a more ornate orna- mentation and a narrower median emargination of the occipital border. In all other respects insofar as these specimens can be com- pared they are in full accord. For the present, therefore, it seems safe to refer it to T. wyomingensis. The profile of the dome confirms the correctness of the restoration of the missing parts of the type as originally published. The dome surface throughout its whole area is smooth, being devoid of the foramina and markings that characterize the Troodon validus skulls. Viewed from above, see fig. 2, the posterior border is truncate with a narrow median emargination devoid of ornamentation. The supra- Fig. 2. Skull of Troodon wyomingensis Gilmore, Cat. No. 3180 C. M. Viewed from above. One-fourth natural size. temporal fossae as in the type specimen are entirely roofed over by bone. On either side of the smooth emarginated median area, are three parallel transverse rows of rounded node-like protuberances. These nodes alternate in the three rows and form an ornate sculpture for this portion of the skull. The tops of the most inferior row, how- ever, are flattened and they lie principally upon the inner branch of the squamosal, almost hiding it from a posterior view, see fig. i. On the left side a portion of the coalesced postfrontal and squamosal 1936 Gilmore: Skull Cap of Troodon 111 bones have their surfaces ornamented by low bosses and lines marking out scutal areas. Fig. 3. Skull of Troodon wyomingensis Gilmore. Cat. No. 3180 C. M. Viewed from below. O, orbital surface; S.t.f., supratemporal fossa; Sup.o., supra- occipital. One-fourth natural size. The ventral view is more broken and less complete than the type, and for that reason contributes nothing to our further knowledge of the structural details of this portion of the skull. COMPARATIVE MEASUREMENTS No. 3180 Type C. M. U. S. N. M. Greatest width of skull across the squamosals 240 310 Distance from center of orbital roof to rear of skull 190 244 Greatest width of dome mass 204 275 Greatest ventral thickness, about 125 180 The skull cap is all that is known of the skeletal structure of Troddo^i wyomingensis. In 1924, I published^ a description of a partial skeleton of the very much smaller Troodon validus (Lambe) and at that time a preliminary skeletal restoration was attempted. It was referred to the new family Troodontidae, which in turn was assigned to the Ornithopodous ^Gilmore, Charles W., Bull. No. i, University of Alberta, 1924, pp, 5-43, pis. 1-14. 112 Annals of the Carnegie Museum VOL. XXV dinosauria. This disposition of the animal aroused strong criticism, especially from the late Baron Franz Nopcsa,^ who at first contended that it pertained to the armored dinosauria and later adopted the more radical hypothesis that the skeleton was a composite, consisting of the skull of an armored dinosaur, the skeleton of an unnamed ornithopod, and some fish bones that formed the abdominal cuirass. RusselF has made a full and adequate reply to these criticisms, in which he gives support to Gilmore’s original contentions. After a lapse of more than ten years I am still of the opinion these materials were properly associated and properly classified. Mr. George F. Sternberg who collected the specimen assures me that all of the bones attributed to this skeleton were found in close association within a small area of comparatively barren strata. It seems most improbable, as pointed out by Russell, “that the skull of one animal, the partial skeleton of another, and some fish bones” should have been collected within this small area.” There was no duplication of parts, and the proportionate size of skull to skeleton was in harmony. Only by the discovery of supplemental and more complete materials can the questions raised be settled for all time, and it is therefore with great interest that we await the discovery of more complete specimens of Troddon, which will disclose the structural peculiarities of this interesting dinosaur. ^Nopcsa, F., Geol. Hungarica, Ser. Pal. fasc. 4, 1929, pp. 64, 65; Ann. and Mag. Nat. Hist., 1931, Ser. 10, vol. VIII, pp. 70-72. ^Russell, L. S., Annals and Magazine of Nat. History, 1932, Ser. 10, vol. IX, p. 334. 561. 73 NOTES ON A COLLECTION OF REPTILES FROM BARRO COLORADO ISLAND, PANAMA CANAL ZONE ^ ) - '>'3 ^ . ; BY M. GRAHAM NETTING Reprinted from the Annals of the Carnegie Museum Vol. XXV, p. 113-120, 1936 , Issued June 25, 1936 «7 €7g|sisf : ^'7 77; 7; 7 f 7 7;.* - ; ^3 -?^:7 :V«fe47 ;'3 » '/s .7'i7-'''7; ’■■■ 3-.' ■ ' ;;^ A >/ , ■- : •. , '•(. ' 3 - ;, - - . ; -r ^' - rtV ' / , 3 vr 7 - .; ^ ,7r .v^r- ''- ■■/ , . ‘‘ V ' ^ " :7 ' .■ ■ \ 'V-i'lv'' /^■'7'3N';V7; .7v7' ' ''' ' ' ',\ ' ' . ' ' 1 . 7' ■■'. ,7.7r'-;V'„,:;, , 7 .• - ' . , ' ': , ,: . -i\. _ ' V 7 I ’ .'" -7' ' , . ^ .' - ' '^'-'J''’ 'P ■' •' ,,' 7' 77 :Ai;J:rAy .7-^'' 77; ’>7:7-3-^ .r7^' .7:'7' , '.■■■> ^■■' - :J--^ - y-ii .V77, V i.<:77-7.',7y:..7..7... ..77-''7-:>77-7^7r";:7,7-'K A'v•^ ' ':a - . , , '.i , y .,,^ ,x.., ,._ . .» ■7.*i-:''" ' c --^.v '■*■,-;■■■', ' 7 ,..' -7 '-/'''V }>y^;v-7 'v A :,"'.A;;'--'7:r2' ■■''.r-’^-3"A ’: ^"^7.: /I i -.^7^ , - . ^ ■■777^7 ’’‘7 3 3 ^ • 7;7' "7> 7 .;S7*:7'3}^;'7 7":"77 ■; ' • ' . 7 ?■.,; -■^ ' , ,'7,'^ - ' ^7 , 37^ ,.3 . -. , , .7 ,.'7. '7f73..-_ .■7 ;v 77 '.7-,' ‘ 7'7'7 - ^7-':-\> "y-: 7 7 7 's.'7>' .7 ■ 7 ^- ■■ ' • .^yv • 7:U3r N ■7 ,. 'A';v.77 '■■' '■ -'7-7''. y'-' ■7- V 'Y ' ^ 7 : y ■' A'- A-A'-'- AA'^ ■'■'"' -7 -7 ' .-■ y ' '^ v " , Y \ f I 7, 7 7 ■ 7> . '■ < . -'7 7;- - ::y' ^ 77 l;77 . 7S 7-7 7 . /7.^y ,•7 7 7 7'f'7. ^ 7 '■' 7'. A > i A ^ ^ ^ y . ^ ^ 7; i'7y • y 7>-.‘- ;- 7 ^ AA 7 7- ;,737-y 7.-'- 7 , -,v ' f' ^ ' I :7f77 7-7. 'Ay V .. ' ' ’■ 7>'' A 7 -A 7 -S; .> . 7 3;'^777| 7.r7-^;' ; A7^.' r- 'A :■ . ' Ai,A ..■' 7 -• j.- ••. ' .' ;■ 7 7.:;7> ■ 4 77?.>. ' ./' ‘■'‘’'•7' A 77 y,; y 7 y-7 ART. XII. NOTES ON A COLLECTION OF REPTILES FROM BARRO COLORADO ISLAND, PANAMA CANAL ZONE* By M. Graham Netting (Plate XIV) During March 1934, Mrs. Netting and I spent twenty-six days on Barro Colorado Island, which is situated in Gatun Lake in the Panama Canal Zone. This island, which has been a natural reservation since 1922, is the seat of a Biological Station which is maintained under the able directorship of Dr. Thomas Barbour, under the auspices of the National Research Council. Every scientist who visits the island leaves with a deep appreciation of the foresightedness of Dr. Barbour, and the wisdom of the National Research Council, in initiating and sponsoring the only biological station in tropical jungle under the American flag. Similarly, every worker receives at the hands of Mr. James Zetek, the very helpful and energetic Resident Curator, more counsel and assistance than he can ever repay or adequately ac- knowledge. During our short stay I did not attempt to collect large series of specimens since the herpetological fauna of the island is well known from the previous work of Dr. Barbour, Dr. Dunn, Mrs. Gaige, and others. I did endeavor, however, to collect one specimen of each species at every spot on the island at which I encountered it. Since my efforts were concerned especially with amphibians with a view toward determining accurately the distribution of the various species, most of my collecting was done at night, and the collecting of reptiles was secondary to the main purpose. A large collection of frogs, in- cluding twenty-two of the thirty-one island species, and representing *I am indebted to my good friend and mentor, Dr. E. R. Dunn, who critically determined the lizards and snakes, and who, from the time he first stirred my interest in Barro Colorado, has given unstintingly from his wide knowledge of tropical conditions and faunas. 114 Annals of the Carnegie Museum VOL. XXV many localities, was assembled. Analysis of this collection indicates that the streams of the island, on the bases of their distinctive frog populations, may be divided ecologically into five zones. Since the report on this collection is not yet ready for publication, it now ap- pears desirable to report briefly upon the smaller reptile collec- tion. Surprisingly enough the collection of amphibians added no species to the island faunal list whereas the collection of reptiles, which totaled only sixty-four specimens, proved to contain new records. Fifty-seven species of reptiles were previously known to occur on Barro Colorado. My collection includes only sixteen species, but of this number three of the nine species of snakes proved to be new to the island. These additions increase the known reptile fauna to sixty species, of which number exactly half are snakes. The high percentage of unreported snake species in my collection, may be attributed to night collecting since each of these additions was secured at night. The collection is far too small to justify any statements as to the distribution of most of the species involved, but definite stations for each species are included here in the hope that this action will stimulate future workers to carefully document their specimens so that those which are added to museum collections in the future will be available for distributional studies. Each of the existing trails on the island has been named, and each trail has been marked at lOO meter intervals, which greatly facilitates the location of collecting spots. ‘‘Stations,” as referred to below, indicate the number of such intervals from the origin of the trails named. It should be remarked, in this connection, that the naming of each stream on the island would be of inestimable benefit for future distributional work. When Barro Colorado Island is compared with Trinidad, with which the author is reasonably well acquainted, the extremely small lizard population, considering numbers of individuals, of the former is little short of amazing. Snakes are somewhat more common on thickly populated, and mongoose-ridden, Trinidad than on Barro Colorado. However, many habitats are not represented on Barro Colorado; important rodent-attracting crops are not grown on the island; and protection is accorded to all of the birds and mammals, many of which are reptile predators. 1936 Netting: Reptiles from Barro Colorado 115 TESTUDINATA Kinosternidae Kinosternon leucostomum Dumeril Three specimens, C.M. 7700-02, of this Atlantic slope form were found in a steep-sided, leafy-bottomed pool near the headwaters of the large stream which crosses Abram Conrad trail near station 2. These turtles were taken after dark on March 21 in less than ten minutes even though the water was murky and waist deep. I was surprised to find the species so numerous and so easy to capture in a stream in- habited by at least one large Caiman. LORICATA Crocodylidae Caiman fuscus (Cope) One young specimen was collected in the lowest pool of Allee Creek, near the laboratory dock, on March 9. This specimen, which was kept in a cage under our cottage, died while we were collecting at another part of the island. On our return our attention was called to its demise by circling vultures, which could not have seen the specimen. Furthermore, decomposition was not far advanced for the specimen preserved properly. Another small specimen was seen in the pool with the turtles mentioned above, and a specimen over five feet in length was observed about a hundred yards upstream from this pool. SQUAMATA Lacertilia Gekkonidae Sphaerodactylus lineolatus Lichtenstein One specimen, CM 7672, was found on the wall of a building at the laboratory on March ii at ii a.m. Several other specimens were observed in the daytime moving about on the inner walls of buildings in the vicinity. These geckos are more diurnal than Hemidactylus or Thecadactylus but far less tolerant of direct sunlight than are some species of Gonatodes. 116 Annals of the Carnegie Museum VOL. XXV Iguanidae Anolis frenatus Cope A sub-adult specimen, CM 4667, of this large Central American species was collected at Snyder-Molino trail, station 2, on the morn- ing of March 10. A newly-hatched specimen, apparently referable to this species, was found sleeping on a tree leaf on the bank of Lutz Creek on the night of March 23. This young specimen, which has a total length of 109 mm., was uniformly green in color. Dr. E. R. Dunn informs me that this name must be used for the B.C.I. speci- mens which have previously^ been referred to A. squamnlatus, A. longipes, and A. purpurescens. Anolis limifrons Cope Thirty-two specimens of this extremely common lizard were col- lected at ten stations in widely separated parts of the island. Many specimens were taken at night while they were sleeping on leaves of plants and trees. Basiliscus basiliscus (Linnaeus) The “moracho” probably occurs along the entire shoreline of the island, and inland along the larger streams as well. Although only the specimens listed below were collected, literally hundreds were observed. Two nests of hatched eggs were found, which Sylvestre Aviles, an excellent woodsman, insisted were those of this lizard. I was unable to prove the connection, but the nest situations indicated Basiliscus as the most probable species. The first nest, of seven eggs, was found on March 7 in a small cavity, in an eastwardly-facing vertical red clay bank, 56 inches above a dry stream bed and approximately ten yards from the lake shore. The eggs measured approximately 20x12 mm. and were white-shelled. The following day seven egg shells were found scattered on top of the ground beside the concrete steps leading from the dock to the laboratory, and only a few steps above the dock. 1 CM 7661 Laboratory dock, March 8. 2 CM 7676-77 shore at Barbour House, March 12. I CM 7679 Alice Creek, March 12. ^Barbour, T. 1934. Bull. M.C.Z., 77: 147. 1936 Netting: Reptiles from Barro Colorado 117 I CM 7698 Abram Conrad trail, near station 2, March 21. I CM 7722 dock at Fuertes House, March 24. Teiidae Ameiva festiva Lichtenstein This species was observed frequently on the edge of the laboratory clearing, near the beginning of Barbour Lathrop trail, but because of the proximity to the laboratory no specimens were shot. Ameiva undulata (Wiegmann) This species proved to be less common than I had expected it to be. Only two specimens were collected, one, CM 7665, at Snyder-Molino trail, station 4, on March 10, and the other, CM 7678, at Thomas Barbour trail, station 13, two days later. SERPENTES Colubridse Xenodontinae Leptocalamus sclateri Boulenger (Plate XIV, figure i) One specimen, CM 7683, was found in a litter of leaves and trash in a small depression in a steep bank about three feet above a wet ditch. This ditch was beside Barbour Lathrop trail between stations 3 and 4. The specimen was collected in the early evening of March 14 and would most probably have escaped observation completely had not the beams from our headlights picked up its striking white head. This specimen is the first to be taken on Barro Colorado Island. Urotheca dimidiata (Cope) The Lake trail is bisected by a large stream which falls rapidly to- ward the lake through a deep ravine with very steep clay banks and a rocky bed. One specimen, CM 7684, was captured about 8 p.m., March 15, as it was crawling over the leaves in a dry section of the stream bed. While I was engaged in grabbing and bagging this 118 Annals of the Carnegie Museum VOL. XXV specimen, I saw another about fifteen feet above the stream bed on one of the clay sides, and Mrs. Netting saw another high up on the opposite bank. Unfortunately, in spite of our best efforts to scramble up the slippery banks both of these specimens eluded capture. The alternating rings of blue-black and coral red of these examples were far too brilliant, in the light of our headlamps, for us to have any doubt as to their conspecificity with the specimen captured. This mention of the snakes ‘‘which got away” is included only to indicate that al- though the specimen at hand is the first to be taken on the island, the species cannot be considered rare. Colubrinae Dendrophidion dendrophis (Schlegel) One specimen, CM 7711, was collected on a dry stream bed near Fuertes House on the afternoon of March 24. Imantodes cenchoa (Linnaeus) Three specimens were found on bushes at night, as follows: 1 CM 7694 Donato trail, station 4, March 16. 2 CM 7720-21 along stream near Zetek House, March 29. Imantodes elegans (Boulenger) The first specimen to be secured on the island, CM 7723, was col- lected at night along Allee Creek above Pearson trail, on March 30. In addition to the difference in color, this species differs from the preceding in having smaller vertebral scales, and in having three labials in contact with the eye instead of two. Leptodeira annulata annulata (Linn^us) One specimen, CM 7682, was taken on the night of March 13 on a stream bank near Raymond C. Shannon trail between stations 7 and 9. A second, CM 7685, was collected on the evening of March 15 on the bank of the stream which bisects Lake trail. 1936 Netting: Reptiles from Barro Colorado 119 Phrynonax poecilonotus Shropshire! Barbour and Amaral This was the only snake which I observed about the laboratory. The specimen, CM 7662, was taken from an orange tree near the laboratory on the morning of March 9, and CM 7680 was found crawling through the grass in front of the laboratory on the morning of March 13. Both of these specimens were of the reddish phase. A third specimen, CM 7696, with yellow markings, was caught by Cristobal Marquinez on the tiny island of Slothia on March 21. This specimen was kept for some time in a cage with several examples of Kinosternon where it frequently exhibited the defense reaction of the species. At such times the neck was inflated for about eight inches, thus displaying the yellow-marked scales, the neck was also flattened vertically, the lower jaw was swung to one side, and the mouth was held half open, and every few minutes the snake hissed or struck at the moving turtle. Oxybelis acuminatus (Wied) (Plate XIV, figure 2) Four specimens were collected, between nine o’clock and noon, at the localities listed below. CM 7670, which measured 847 mm. in body length, contained a Basiliscus hasiliscus which was 275 mm. in length from the forelimbs (head digested) to the tip of the tail. I CM 7660 Lake trail, March 8. I CM 7670 Alice Creek below laboratory, March ii. I CM 7695 David Fairchild trail, station 7, March 17. I CM 7697 Wm. Morton Wheeler trail, station 8, March 20. Spilotes pullatus pullatus (Linnaeus) One “tigre,” CM 7668, was collected on the afternoon of March 10 on Donato trail. This snake was in very poor condition, possibly due to a heavy infestation of ticks, of which over 200 were counted on its body. 120 Annals of the Carnegie Museum VOL. XXV EXPLANATION OF PLATE XIV Photographs from life by M. Graham Netting Fig. I. Leptocalamus sclateri Boulenger, C. M. 7683. Fig. 2. Oxybelis acuminatus (Wied). ANNALS CARNEGIE MUSEUM, Vol. XXV Plate XIV. Figure 2. ! ART. XIII. VERTIGO CLAPPI, A NEW LAND SNAIL FROM WEST VIRGINIA By Stanley T. Brooks and G. R. Hunt Vertigo clappi sp. nov. Shell brown, shining, striated, striae forming slight riblets on the last two whorls; umbilicus open, deep; whorls five and one-half, the last strongly constricted behind the lip; aperture biarcuate, small, only slightly expanded; upper palatal fold long, blade-like, curved, springing from the heavy ridge formed by the intersection of the two arcs but beginning quite back of the lip; the two columellar lamellae are blade-like, rounded, similar in shape and lie horizontally and close together; infraparietal and angular lamellae prominent but due to the small size of the aperture are close together and closely approach the columellar and palatal margins of the aperture. Altitude of the shell 1.5, diameter 0.8 mm. Fig. I. Vertigo clappi Type locality: Renick, Greenbrier County, West Virginia; Carnegie Museum collection, holotype No. 62.28186. Collected by Mr. G. R. Hunt during the summer of 1935. The authors designate this new species as Vertigo clappi in honor of Dr. George H. Clapp, whose erudition and enthusiasm is a constant inspiration to his admirers and associates in his scientific work. 121 mar 2 5 C- Issued July 15, 1936. So 1, 1 1 // i, » ^ , '4- THE LAND SNAILS COLLECTED DURING THE 1936 VOYAGE OF THE “VAGABONDIA” WITH THE DESCRIPTIONS OF NEW SPECIES OF PLECTOSTYLUS J, r- '' ( I: I 15"' ^ “ -y- ./ .V Vi '"V ' ■ ' V (I jpy Stanley Truman Brooks ' ) , ■■ ! ) 'Pi:' , , y-‘ ); , ’,y ■ y- " 'I ’ _;v" - . . r , ' >y pUp , ■• ' ■ ^ ' Y r ' y= - ■ ' ' P', '■! V ■ y.'' :■ Reprinted from the Annals of the Carnegie Musem Vol. XXV, p. 123-126, 1936 Issued July 15, 1936 .-y- ■ S P'- 'V" ) pp. , • , - f ■; ^ '-A'' , ' ~ ‘ - ' I' ‘ ' ,, ' . ' ^ ^ . . ' / ' ■ ' ' , j , V" ' ,i , n , ,’ ' '-? ■..V, '/.!./ . - .-■.U . - .. ^ ' 'V, ’ "■V ^ ^ ' , / . * if' ' . ^ ' ' /’ . . - ’ J . ^ ^ ^ '■ ’. ■ , ' ' ,■'? ‘ ^ 4'" ^ \ . v ^ ’ " ' fc r I . ': , ^ ^ . 4 - , If -4/ '■'. ,;4 : ' V f’" ' ''^^ -'->4' : '4 '-4x^'', i- "'■'". <, - ■ ■ ^ * -; -■'’ ' - '"' y|,./- ■' ^ - - '/ 'I ‘ ■ " - i ' .-' r/'j' ' I- ■, \--'‘ -. • ' '■■' ' ■'■ ,{ ' , '■y V4 :,.. r.y - /v, ;, ',, ; ,y. I , > ,;•' ■■■« ! ' 'y y,- '■ ■; V';- i ./i -. . -fr: i. ■• ■ '-, '■ _ ' ," ('' ' - I _ ,■■ :,; ,- ■4 s', , ■■■■' 4' •,' 44 - ' 4''--'':^ Ts 4 .S4', .4' '■ 'i ■■■;,, '44/’;4i ’ ; , 4:''.s- 4^ .sl'-N / ■■,','"s4 > , ; 4] 4'S V N>, ■'■ fs4.s5 4 ^ . 1 .'\ I ^ .V ' f ^ ^ ‘4,1 V- ' ■ y : 4 ’■ ■' '."C M 'I '' .) -4 '-' ^ > 4 ^ . 44 ' -.1- ' ^ . ^'4 % ■' ^ Vt . 4 ,'’s.4-s!--.s ij ' ' . I ' : V ’■ ';,.V i -'4 4 ■' -'f 44^... ,:,V n ■■'■■•■'.: K) ■' 4 ,.../^ 4' '.4:'4.y... 4s4->"4-'4 s 4 J i 44~-44^^/v - y ■ ,s44 ■'■'• ' -■ .. — ' 4 ■ '''■ '■'• ,., 4-- ’'•■'W-,‘ ■ ", . ',4' ■'/44')--.;; ■■ 4^ -'S- ,,, S' s ' '’''■'j ” '/'■ ’'-J ^4 ' ^ ‘ ^ ^ * ■' , ,, .,■, , ' , ■ ' .)■ , ■- ';' ':v yv'-x,: , ,’44 . .;',v ’ ^ ' " ' s' Vi4 , ."'4-/'''4- ' ■ ' ,' ■■ : ■ , ’ I // ■■ ■ ■--, ■ . I ‘ .1 ^ ' , * ' ^ . ■ ' " 4444-:: s ■■ \s}4-- 44,;,'"- „'^' ; •I' .-■ r ■ ■4' .44 4 4- 4:^4,.,. 4^,:-^ "4^ I 'S'! ,4' :444%5s/4 4 :,.s;t-. ... ' ,4 4/ ^ ' .- !44 ■■ ' ’ , , ' -■' , , s' 4 ■ ' , t '''' ' • .j"; "’• 4; ^ , •' ^ ' , S' ' ,4' , 4 .,!" i. S. f ' '' 'V ,- >;'^'''Vyr' ! V; ' 7'^> ■ , • ^ ' > . -y. '4 •■ , 4/„;y_> 7 ^ ; ART. XIV. THE LAND SNAILS COLLECTED DURING THE 1936 VOYAGE OF THE “VAGABONDIA” WITH THE DESCRIPTIONS OF NEW SPECIES OF PLECTOSTYLUS By Stanley Truman Brooks During the winter and early spring of 1936, the Yacht “Vaga- bondia” belonging to Mr. W. L. Mellon of Pittsburgh cruised the waters of the Caribbean, then went through the Panama Canal to the southern Pacific and south to Tierra del Fuego and the Straits of Magellan. Through Mr. Mellon’s thoughtfulness and his realiza- tion of the great scientific value of such a cruise, one of the staff of the Carnegie Museum, Mr. Reinhold L. Fricke, accompanied him. Collections were made at various points during the journey. The land snails were sparse and hard to obtain but the findings in this small collection indicate the necessity of more extensive and specialized efforts within the areas encompassed by this voyage. However, Mr. Fricke should be highly complimented for his efforts as this collection of gastropods is only a minute fraction of the many speci- mens obtained by him, including hundreds of birds, mammals, rep- tiles, and several hundred crustaceans. The latter are now in the hands of the proper authorities and will be reported upon in due time. To Dr. H. A. Pilsbry, of the Academy of Natural Sciences of Philadelphia, I wish to express my appreciation for his aid in the proper allocation of the following forms. His great and profound wisdom is constantly a source of wonder and inspiration to the other students in this field of endeavor. Lastly, as an acknowledgment of the interest of Mrs. W. L. Mellon in the mollusca, and of her kind efforts in behalf of the laboratory of Recent Invertebrates of the Carnegie Museum, I propose that one of the following new species shall bear the name Plectostylus maricE in her honor. In the following list of specimens collected, the species are arranged by localities. 123 Issued July 15, 1936. MAR 25 1940 124 Annals of the Carnegie Museum vol. xxv Locality: Beata Id., San Domingo Brachypodella laterradii (Grateloup) Chondropoma sp. indet. Close to C. weinlandii Pfr. In the absence of more specimens it is thought best not to describe this specimen as a new species. Description: Shell waxy-bluish gray, clouded with white; apex eroded, whorls five with two broken brownish-yellow bands, three on the body whorl; sutures deep, crenulated by the vertical striae; base of each whorl over- lapped by the following one, distinctly sepa- rated from the body whorl at the aperture forming a long (i to 3 mm.) triangular groove; striations well marked, vertical, an indication of spiral striae may be seen on base with high magnification but this is not very clear; umbi- licus moderate, visible from the base; aperture oval, oblique, pointed at its apex, reflected and thickened within; lip bears a well defined sulcus, deepest at the apical and basal extremities, slight along the columellar border. Altitude 16.2, diameter 9.4 mm. Carnegie Museum no. 62.28175. Locality: San Esteban, Chile Plectostylus mariae sp. nov. Named in honor of Mrs. W. L. (Mary) Mellon. Shell imperforate, thin, ovate, yellowish corneous with oblique, longitu- dinal streaks of dark yellow and brown; on the upper whorls irregular, vertical arcs of dark brown appear, the last occurring half way down the body whorl (as in reflexus Pfeiffer) ; surface smooth, shining, under the lens it appears granulated by very indistinct spiral lines; spire short, pointed; whorls five, quite convex; aper- ture long-ovate 19.6 mm. in altitude and 8.3 mm. in diameter, bluish white within, peristome thin sharp, with brownish bor- der; columella granular, rounded and thin. Altitude 29.6, diameter 19.7 mm. Carnegie Museum, holotype no. 62.28184. Collected on trees; a much more convex shell than P. reflexus but undoubtedly of this general group. Chile 1936 Brooks: Land Snails of "Vagabondia” 125 Plectostylus vagabondiae sp. nov. Another shell, presumably found at the same locality, differs from P. maricB in so many particulars that I shall tentatively name it Plectostylus vagahondicB. Shell thin but heavier than preceding; found living but with the shining epidermis only along the columellar border of the aperture, surface chalky bluish white marked with brown bars and patches and with few vertical bands of brown; surface chalky, marked with vertical growth wrinkles and indistinct scattered spiral lines; sutures deep, ir- regular, bordered by the constrictions and lines of the following whorl; spire long, conic; whorls six, convex {not as convex as P. marice but more so than its nearest relative, P. reflexus) ; columella thickened, rounded, granular and almost pustulate under the lens, appressed, forming an angle with the peristome which is slightly reflexed at the columel- lar margin; aperture bluish-white within, peristome thin with dark brown border. Size of aperture is i6.l in altitude and 8.9 mm. in diameter. Altitude of shell 34.2, diameter 14,8 mm. Carnegie Museum, holotype no, 62.28185. 126 Annals of the Carnegie Museum VOL. XXV Locality: Juan Fernandez Island, Chile Fernandezia sp. Juvenile specimens only. Fernandezia bulimoides Pfeiffer These are also very young and the identification is doubtful. Helix aspersa Muller* Oxychilus alliarium Muller* Oxychilus cellarium Muller* Tornatellina plicosa Odhner Locality: Salado Bay, Chile Lissoacme albicans Broderip Locality: Cocos Island Guppya pacifica (Pfeiffer) Ochrodermella cumingiana (Pfeiffer) Nesopupa cocosensis (Dali) Guppya hopkinsi Dali *First reported from Juan Fernandez in 1931, by Odhner, N. Hj., “Mol- lusca from Juan Fernandez” (addenda), Nat. Hist. Juan Fernandez, and Easter Island, Uppsala, Vol. 3, part 4, pp. 481-482. I dt THE AMPHIBIANS OF THE PULITZER ANGOLA EXPEDITION > BY Karl Patterson Schmidt . ' 4-'-- -'v, ' '.v; : " : A.:'. J-': ■ ,K --'-.A" .p, . .V 4J:- A r: ') : ;y : . > ■ > ' ' A„ J T'T^ ;..A - •-' ■ ->,A\ A '■• ■■''' ",, i ’■ - "r. 'v ' ■ ■" . (■' \A,- ^ /'/ ' -.-'a i . ^ -■■' . ' I i ■'■,''■■ > '■ ■ ■' , A •'it''" -- ',- .ji ' 'tv Reprinted from Annals of the Carnegie Museum Vol. XXV, p. 127-133, 1936 '4'. . ' - V Issued July 24, 1936 .1 ‘-,. J, t' . f ( /. -•7l'-;' ' 'r!^- ■ - ■' A'" • '■ ■ ' ' ■■■■ -3 V- -■ ^ ^-’:vUA;’t--A'^ '-'W . x'''; "'’ ’ Av-; AtvA'^ ./''"rVAvA’^l 'AVA/f : ['-yh V ' \' :-^/- V- ' /4:/'A • ''AA. ^■■A-' ('Ai^ ''•. A <'1' ; ■ V'.,, ' i-ji. , I-' "C' ' S' -'' ' /■ ' ;i. " -AA;AfiAT- AV'a v ■ S «' \ A.v4- ' .:-T-v-:p"A;i''';'i>;:v^',A; ;>A;;"..PA3 v\.'" t i A ' : t^, /" ;'X- ■,■ ■ 'A -A''^ ' • ■ ' VA/ rvA Ta: -f- "'ij -A. A , Ay ” ' '■ "\ j I' / \; -■ . 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'.v^y A:yAv/:y k A i* - v"'' Kx, A 'A 'a 'A| -■-.' '^ . A--A77 .-,1' , J , A' , - . ' I , ,'s ^ V A^.y,'* /I ..,. ■ -■ vyy AA vvv^. A : ■_ ' ‘A ,.. ,v I V A - A- , . -y'"']. ',y I- - ''A';? , ■ !•' '"' ■•' '.'■•V ,-A '. A vl -A V -M'y ^ > '■V '' ^'V ’f : A'm ,L:;nA;'7. f A A A'- 1 A AAm- /li.S Ml.')- r,“ , 77 / ' ■ //■ " i^'.v; r*; : ., ,7 A'- aY- ' ■ ' mAv'’ :. ../-vA,, Al. <-P . yv[ -/ K AA^'' ^y ' A A A.' A.' YoiA^rAAr AA''AAi;.A2 ART. XV. THE AMPHIBIANS OF THE PULITZER ANGOLA EXPEDITION By Karl Patterson Schmidt, Assistant Curator of Reptiles, Field Museum of Natural History Introduction The collection of frogs and toads made in the course of the Pulitzer Angola Expedition in 1930 and 1931, by Mr. and Mrs. Ralph Pulitzer and Mr. and Mrs. Rudyerd Boulton, amounts to 442 specimens, representing seventeen species. The reptiles of this expedition have been reported upon in a previous paper by myself (Schmidt, 1933), which includes a map of Angola with the collecting stations of the expedition indicated. While no new forms are represented in the collections listed below, the number of references to synonymy, of revivals of names pre- viously placed in synonymy, and of extension of range of forms well- known in East Africa or in the Katanga, is sufficient to indicate the important place of the Angolan fauna in the elucidation of the taxo- nomic and distributional relations of the African forms. The large collections made by Messrs. Rudyerd Boulton and Herbert Lang for the American Museum of Natural History, on the Vernay Angolan Expedition, in 1925, are still unreported. The taxonomic maze in the African frog fauna presented by ffie genus Hyperolius with its hundreds of species, can be clarified only by the combination of work in field and laboratory exemplified in the collections made by Mr. H. B. Cott (1932) in Mozambique and reported upon by himself and Mr. H. W. Parker (1931). It may be added that experience with the similarly difficult genus of American frogs Eleutherodactylus points to the fact that many of the species will prove immediately distinguishable by attention to their voices in the breeding season. 127 Issued July 24, 1936. MAR 25 IMO 128 Annals of the Carnegie Museum VOL. XXV SALIENTIA PIPIDiE 1 . Xenopus petersii Bocage Xenopus petersii Bocage, 1895, Herp. Angola, p. 187. Xenopus poweri Hewitt, 1927, Rec. Albany Mus., 3, p. 413, pi. 24, fig. 3. Twenty-five specimens, Carnegie Museum Nos. 6601-6612, 6892- 6893, Gauca, Jan. 8-10, 1931; and 6699-6701, 6724-6728, 6852, 7010- 7011, Chitau, Jan. 11-15, 1931. These specimens are all very heavily marbled with black beneath, and are readily distinguishable by this character from Xenopus Icevis Icevis. The largest specimen. No. 6601, measures 78 mm. from snout to vent, and the smallest 23 mm. Juvenile specimens all exhibit a striking dorsal pattern of ocellar spots, which tend to become obscure in the adults. I have no hesitation in referring Hewitt’s Xenopus poweri from Northern Rhodesia to the Angolan petersii in view of the normal east-west range of Angolan savanna forms south of the rain forest. (Schmidt, 1919, p. 447, map 6; 1933, p. 10). BUFONID^ 2. Bufo regularis regularis (Reuss) Bufo regularis Reuss, 1834, Mus. Senck., i, p. 60. Bufo regularis regularis Loveridge, 1932, Occ. Papers Boston Soc. Nat. Hist., 8, p. 53- Seventy-three specimens, Carnegie Museum Nos. 6613-6645, 6671-6673, 6681, 6685, 6689, 6897-6904, Gauca, Jan. 2-10, 1931; 6658-6666, 6694, 6809-6820, Chitau, Jan. 12-16, 1931; 7027, Angola, 1931- I follow Loveridge, (1933, p. 354) in referring the Bufo regularis of Angola to the typical subspecies, on the ground of geographical probability. RANID2E 3. Rana fuscigula angolensis Bocage Rana angolensis Bocage, 1866, Jorn. Sci. Lisboa, i, p. 54. Rana fuscigula angolensis Loveridge, 1933, Bull. Mus. Comp. Zool., 74, p. 362. 1936 Schmidt: Amphibians of Pulitzer Angola Expedition 129 Seventy-two specimens and seven lots of tadpoles and transforming juvenile specimens, Carnegie Museum Nos. 6670, 6676, 6686-87, 6894-95, 7020, Gauca, Jan. 5-10, 1931; 6690-93, 6705-7, 6719, 6739-53, 6762, 6764, 6766-82, 6796-6808, 6821, 6826, 6834, 6856-61, 6869, 6874, 6891, Chitau, Jan. 12-15, I93i; 7024, 7028, Angola, 1931. 4. Rana mascareniensis subpunctata Bocage Rana subpunctata Bocage, 1866, Jorn. Sci. Lisboa, 1, p. 54. Rana anchietoc Bocage, 1867, Proc. Zool. Soc. London, 1867, p. 843, fig. i. Rana porosissima Steindachner, 1867, Reise, Novara, Amphib., p. 18, pi. i, fig. 9. Rana mascareniensis uzungwensis Loveridge, 1932, Bull. Mus. Comp. Zool., 72, p. 384. Five specimens, Carnegie Museum No. 6647, Gauca, Jan. 8, 1931; 6714, 6763, 6823, 6825, Chitau, Jan. 12-16, 1931. I have applied the oldest Angolan name to these frogs, whose final definition will require the study of abundant material and com- parison with the types. The specimens at hand agree excellently with the description and figure of Rana anchietcB. Angola is included by Loveridge in the range of uzungwensis, and it is clear that if this is correct, one of the names based on Angolan material must be em- ployed; a paratype of uzungwensis from Tanganyika Territory in Field Museum agrees excellently With the Angolan frogs. In the small series at hand two specimens have the spots of the posterior surfaces of the thighs confluent into well-defined stripes. 5. Rana oxyrhyncha Smith Rana oxyrhynchus Smith, 1849, Ulus. Zool. S. Africa, 3, pi. 77, fig. 2. Thirteen specimens, Carnegie Museum Nos. 6704, 6757-61, 6765, 6822, 6871-73, 7019, Chitau, Jan. 12-15, 1931; 6896, Gauca, Jan. 10, 1931. 6. Rana bunoderma Boulenger Rana bunoderma Boulenger, 1907, Ann. Mag. Nat. Hist., (7) 19, p. 214. Twenty-eight specimens. Carnegie Museum Nos. 6708-9, 6711-13, 6715, 6737-38, 6824, 6827-33, 6866-67, 6870, 6875-77, 6881-82, 6886- 89, Chitau, Jan. 12-16, 1931. These specimens agree with the original description of Rana buno- 130 Annals of the Carnegie Museum VOL. XXV derma except for the presence of an outer metatarsal tubercle, and the somewhat ill-defined tympanum. The specimens key out to buno- derma in Witte’s key to the subgenus Ptychadena (Witte, 1921, p. 7). Males have an extremely well-defined subgular vocal sac, which expands exteriorly through a slit which extends to the base of the arm. The species is readily distinguished from R. mascareniensis by the short webbing of the toes and the rounded or elongate dorsal tubercles which take the place of the regular folds of mascareniensis. 7. Rana albolabris Hallowell Rana albolabris Hallowell, 1856, Proc. Acad. Nat. Sci. Phila., 1856, p. 153. Five specimens, Carnegie Museum Nos. 6754-56, 6788, 6793, Chitau, Jan. 12-16, 1931. 8. Rana tuberculosa Boulenger Rana tuberculosa Boulenger, 1882, Cat. Batr. Sal. Brit. Mus., p. 30. Four specimens, Carnegie Museum Nos. 6862-65, Chitau, Jan. 15, 1931- Rana cryptotis Boulenger (1907, p. 109) agrees in detail with tuber- culosa except in its smaller size, hidden tympanum, and subequal first and second fingers. It seems possible that it may prove to be the young of tuberculosa. 9. Rana sp. Five specimens, Carnegie Museum Nos. 6669, 6679, 6682, Gauca, Jan. 2-6, 1931; and 7016-17, Chitau, Jan. ii, 1931. These specimens clearly represent the young of a species of Rana not otherwise represented in the collections. It is to be expected that they will be identifiable when more Angolan material becomes available. 10. Phrynobatrachus natalensis (Smith) Stenorhynchus natalensis Smith, 1849, Illustr. Zool. S. Africa. 3, App., p. 23. Phrynobatrachus natalensis Gunther, 1864, Proc. Zool. Soc. London, 1864, p. 480. One hundred and sixty-five specimens, Carnegie Museum Nos. 6648-57, 6674, 6677-78, 6683-84, 6905-7009, Gauca, Jan. 4-10, 1931; 6710, 6716-18, 6720, 6731-36, 6783-86, 6789-91, 6835-51, 6868, 6878- 80, 6883-85, Chitau, Jan. 12-15, 1930 7023, 7025, Angola, 1931. 1936 Schmidt: Amphibians of Pulitzer Angola Expedition 131 11. Arthroleptis parvulus Boulenger Arthroleptis parvulus Boulenger, 1905, Ann. Mag. Nat. Hist., (7) 16, p. 109, pi. 4, fig- 3- Eighteen specimens, Carnegie Museum Nos. 6721, 6890 (4), 7012- 15, 7018, Chitau, Jan. 11-15, 1931; 6675 (5), 7021, Gauca, Jan. 4-5, 1931; and 7022, 7026, Angola, 1931. A considerable variation in the dorsal rugosity in this series appears to be due to differences in preservation. I am indebted to Mr. Arthur Loveridge for the identification of this species. POLYPEDATID2E 12. Leptopelis anchietae (Bocage) Hylambates anchietce Bocage, 1873, Jorn. Sci. Lisboa, 4, p. 226. Leptopelis anchietce Noble, 1924, Bull. Amer. Mus. Nat. Hist., 49, p. 234. Four specimens, Carnegie Museum Nos. 6696-97, 6729, and 6795, Chitau, Jan. 12-16, 1931. 13. Leptopelis angolensis (Bocage) Hylambates angolensis Bocage, 1893, Jorn. Sci. Lisboa, (2) 3, p. 119; 1895, Herp, Angola, p. 179, pi. 17, fig. I. Four specimens, Carnegie Museum Nos. 6695, 6722-23 and 6794, Chitau, Jan. 12-16, 1931. These specimens agree exactly with Bocage’s descriptions and figures of angolensis. The species is referred to Hylambates hocagii by Boulenger (1906, p. 166), and this species, retained provisionally in the genus Hylambates by Noble, (1924, p. 247) has subsequently been referred to Leptopelis by Loveridge (1925, p. 787). Until further studies can be made, it is preferable to retain angolensis as distinct from bocagii, since the original description of bocagii differs con- spicuously from the specimens at hand. 14. Hyperolius marmoratus Rapp Hyperolius marmoratus Rapp, 1842, Arch Naturg., 8, pt. i, p. 289, pi. 6. Four specimens, Carnegie Museum Nos. 6668, 6680, 6688, Gauca, Jan. 6-8, 1931; 6854, Chitau, Jan. 15, 1931. Three distinct types of coloration are included in these few speci- mens, one of which corresponds well with H. decoratus Ahl (from 132 Annals of the Carnegie Museum VOL. XXV Cameroon) and a second with H. graueri Ahl, (from Lake Tanganyika). In the present state of taxonomic confusion in the genus Hyperolius it is impossible to come to any satisfactory conclusions regarding the forms deserving of recognition in any given area from laboratory material; but it may be pointed out that these Varieties’ were named long since by Bocage on the basis of Angolan specimens, (Bocage, 1895, p. 164). 15. Hyperolius nasutus Gunther Hyperolius nasutus Gunther, 1864, Proc. Zool. Soc. London, 1864, p. 482, p. 33, fig. 2. A single specimen, Carnegie Museum No. 6667, Chitau, Jan. 14, 1931- This specimen, a male with well-developed vocal sac, measures 19 mm. It exhibits the two dorsolateral light lines mentioned by Bocage (1895, p. 169), as occasionally present in this species. 16. Hyperolius seabrai (Ferreira) Rappia seabrai Ferreira, 1906, Jorn. Sci. Lisboa, (2) 7, p. 163, fig. Three specimens, Carnegie Museum Nos. 6702-3, 6855, Chitau, Jan. 12-15, I93i» are referred here. They are close to Hyperolius cinnamome-ventris Bocage, but lack the lateroventral dark line which appears to be very characteristic in specimens of undoubted cin- namome-ventris from Cameroon which are available for comparison. 17. Kassina angeli Witte. Cassina angeli Witte, 1933, Rev. Zool. Bot. Afr., 23, p. 172, 1934, Ann. Mus. Congo Beige, Zool., (i) 3, p. 183, pi. 8, fig. 3, pi. 10, fig. 8. Four specimens, Carnegie Museum Nos. 6646, Gauca, Jan. 8, 1931; and 6730, 6792, 6853, Chitau, Jan. 12-16, 1931. These specimens agree in their shorter hind limbs with the speci- mens from the Katanga described by Witte, and females have the curious denticulated anal flaps, which are less well developed in East African K. senegalensis. Kassina modesta Ahl, described from Natal, is also shorter-legged than the typical senegalensis. Without much more South African material, it is impossible to attempt a subspecific rearrangement of the forms of Kassina. 1936 Schmidt: Amphibians of Pulitzer Angola Expedition 133 LITERATURE CITED Bocage, J. V. Barboza du. 1895. Herpetologie d’Angola et du Congo. Lisbonne, xx -j- 203, 19 pi. Boulenger, George Albert. 1906. Report on the batrachians collected by the late L. Fea in West Africa. Ann. Mus. Civ. Stor. Nat. Genova (3) 2, 157-172. 1907. Description of a new frog discovered by Dr. W. J. Ansorge in Mossamedes, Angola. Ann. Mag. Nat. Hist., (7) 20, 109. CoTT, Hugh B. 1932. On the ecology of tree-frogs in the lower Zambesi Valley. Proc. Zool. Soc. London, 1932, 471-541, 2 pi. Loveridge, Arthur. 1925. Notes on East African batrachians collected 1920-1923. Idem. 1925, 763-791, 2 pi. 1933. Reports on the scientific results of an expedition to the southwestern Highlands of Tanganyika Territory. VII. Herpetology. Bull. Mus. Comp. Zool., 76, 197-416, 3 pi. Noble, Gladwyn Kingsley. 1924. Contributions to the herpetology of the Belgian Congo. III. Amphibia. Bull. Am. Mus. Nat. Hist., 49, 147-347, pi. 23-42. Parker, H. W. 1931. A collection of frogs from Portuguese East Africa. Proc. Zool. Soc. London, 1930 (publ. 1931), 897-905, i pi. Schmidt, Karl P. 1919. Contributions to the herpetology of the Belgian Congo. 1. Turtles, crocodiles, lizards, and chameleons. Bull. Am. Mus. Nat. Hist., 39, 385-624, pi. 7-32. 1933. The reptiles of the Pulitzer Angola Expedition. Ann. Carnegie Mus., 22, 1-15, 2 pi. Witte, Gaston-Fr. de. 1921. Description de batraciens nouveaux du Congo Beige. Rev. Zool. Afr., 9, 1-22, 5 pi. ARDYNOMYS AND DESMATOLAGUS IN THE NORTH AMERICAN OLIGOCENE BY J. J. BURKE Reprinted from the Annals of the Carnegie Museum Vol. XXV, p. 135-154, 1936 o. Issued December 12, 1936 I ART. XVI. ARDYNOMYS AND DES MATO LAG US IN THE NORTH AMERICAN OLIGOCENE By J. J. Burke (Text figures 1-7) Study of the fossil rodents and lagomorphs in the collections of the Carnegie Museum reveals that the genera Ardynomys and Desma- tolagiis, which have previously been known only from the Oligocene of Asia, are also represented in deposits of that age in western North America. Through the courtesy of Dr. Walter Granger I have been able to compare the specimens described in this paper with fossil material from Mongolia preserved in the collections of the American Museum of Natural History. The illustrations which accompany this article are taken from drawings by Mr. Sydney Prentice. Order SIMPLICIDENTATA Lilljeborg Eamily ISCHYROMYIDT: Alston Genus Ardynomys Matthew and Granger Ardynomys occidentalis sp. nov. Holotype: Right ramus of mandible with incisor, P4, Mi_2 and base of M3, Carnegie Museum No. 1056. Referred specimens: Left ramus of mandible with broken incisor. Fig. I. Ardynomys occidentalis Burke. Right lateral view of skull, C. M. No. 1055 and holotype right ramus of mandible, C. M. No. 1056. Squamosal region of skull restored from left side. X 2. 135 ^AR 2 s Issued Dec. 12, 1936. 136 Annals of the Carnegie Museum vol. xxv worn Mi_2, P4 and M3 broken off, C. M. No. 1105; partial skull lack- ing on both sides and with LM^ missing, C. M. No. 1055; right maxilla with root of Ph worn P^ and C. M. No. 1083. Horizon: McCarty’s Mountain Oligocene. Localities: C. M. Nos. 1056, 1105, “S. of McCarty’s Mt.”, C. M. No. 1083 “McCarty’s Mt.’’; C. M. No. 1055 “S. W. McCarty’s Mts.”^ Madison County, western Montana. Diagnosis: Lower jaw essentially as in Ardynomys olseni Matthew and Granger, but hypoconids of molars better developed, trigonid of P4 less transverse, its cusps not as widely separated and the external valley of P4 not as crowded. Species of smaller size than Ardynomys chilli Matthew and Granger (P4^ — M3 = 10 mm.). Perhaps the most striking features of the skull (C. M. No. 1055) which I am referring to this species are the robust construction throughout, the flattened brain-case, the sharply constricted and narrow interorbital region, and the antorbital breadth, coupled with an angular and heavy type of rostrum. Unfortunately, a great deal of the dorsal portion of the skull has been broken away, but enough remains to show that the brain-case was somewhat wider than the rostrum. The left squamosal is com- plete to the extent of preserving some of the characters of the glenoid fossa, 2 which is in general like that of Arctomys, broad and shallow. A small postglenoid foramen is shown, postero-median to which occurs a large subsquamosal foramen. Despite the loss of the bones investing the brain-case dorsally, the brain cast which remains indicates that this region of the skull must have been flattened to a degree comparable with the same region in the skull of Aplodontia. This cast shows broad cerebral lobes and a fairly distinct superior longitudinal fissure, but the hemispheres have not the heart-shaped anterior tapering which characterizes Cynomys iThe locations included in quotation marks are taken directly from the field labels of these specimens. I assume that C. M. Nos. 1056, 1105 and 1083 were collected from the locality on the southeastern slope of McCarty’s Mountain, about sixteen miles north and a little east of Dillon, Montana. But C. M. No. 1055 (the skull) coming from “S. W. McCarty’s Mts.” suggests the second locality which Earl Douglass mentions in Mem. Carnegie Museum Vol. II, No. 5, p. 21 1, 1905 — the locality two miles west of the main collecting area. The supposition is also strengthened by the fact that of the two specimens found there, one was a Colodon, the other “an apparently new rodent.” 2Part of this squamosal root of the zygomatic arch was accidentally shattered after Mr. Prentice had made the drawings which accompany this article. 1936 Burke: Ardynomys and Desmatolagus 137 and Ischyromys and, but to a lesser degree, Aplodontia\ the outline is decidedly more rounded and becomes almost truncate directly behind the olfactory bulbs. Most of the dorsal surfaces of the olfac- tory bulbs appear to be exposed; they are relatively large, apparently foreshortened longitudinally, expanded laterally and inflated. It is true that the absence of the bones of the dorsal surface of the skull probably exaggerates the interorbital constriction to some extent, but in any case it is quite pronounced. The lateral walls of the brain-case round rather sharply forward to make the constriction, anterior to which the skull again fans outward fairly rapidly to the zygomatic region. There is no means of determining, from this damaged skull whether postorbital processes were present or not; in all likelihood they were absent. The skull under description (C. M. No. 1055) has undergone some damage in the lacrimal region; the extra-orbital part of the lacrimal bone has been lost on both sides, and there has been crushing of the skull walls within the orbits. However, on the right side, the naso- lacrimal foramen is shown. The lacrimal does not enclose the canal entirely (the condition noted by Hill^ in the porcupine) ; in general its construction seems to be as in Arctomys. Of considerable interest, and particularly to be taken into account when the relationships of Ardynomys and the later Oligocene genera Tsaganomys and Cyclomylus are considered, is the possible incipient development, in Ardynomys, of an antorbital crest such as Matthew and Granger^ noted in Tsaganomys. In anterior view the skull of Ardynomys shows the zygomatic region as widely expanded; even in the absence of the malar, which is not preserved in our specimens, this outjutting of the arch is pronounced. However, it is most evident below, where it would probably be accentuated if the malar were preserved. Above, the resemblance to Tsaganomys, as figured by Matthew and Granger, is less evident. The skull, C. M. No. 1055, upon which most of the present description is based, does not, as noted before, preserve the orbital rim entire. The maxillary, C. M. No. 1083, preserves a little more of the rim. Judging from these two speci- mens, the skull of this species had not yet developed an orbital ridge John Eric, “The Cranial Foramina in Rodents.” Jour, of Mamm. Vol. 16, No. 2, p. 123, 1935. ^Matthew, W. D., and Granger, Walter, “New Bathyergidae from the Oligocene of Mongolia.” Amer. Mus. Novit. No. loi, p. 3, 1923. 138 Annals of the Carnegie Museum VOL. XXV comparable with that of Tsaganomys; there appears to have been no marked or abrupt building out of the orbits above the zygomatic root, but there seems nothing to preclude the development of the ridge in the later history of a skull of this type. The orbital and zygomatic regions of Ardynomys occidentalis m. do not show a great deal in the way of specialization beyond the same regions in, say, Aplodontia; there is a strong lateral expansion of the anterior zygomatic root, but the latter is heavy and sturdy, not thin and sharp as in Tsaganomys, In another feature the zygomatic structure deviates from that found in either Aplodontia or Tsaganomys] the upward tilt of the zygoma is more marked than in either of these two genera. The infraorbital foramen is some- what ovate, as in Tsaganomys. I cannot find any evidence that the infraorbital foramen (which is not enlarged) transmitted a slip of the masseter. The rostrum, seen from above, appears unusually heavy; it is quite broad posteriorly and tapers fairly uniformly anteriorly. The premaxillaries reach to the interorbital region; the premaxillary- frontal suture is shown on the right side of C. M. No. 1055, a little posterior to the naso-lacrimal foramen. The nasals are not preserved, but the right nasal must have been broken away practically along the suture between it and the pre- maxillary. Assuming that the nasals were flattened, or but slightly arched, the rostrum must have presented a crudely hexagonal trans- verse section (similar to that of Tsaganomys)] there is a prominent incisive swelling produced by a sharp outward arching of the pre- maxillaries; below the incisive swelling the sides of the rostrum have a long slope downward and inward to the palatal surface. Of the antero-posterior extent of the palate, the premaxillaries constitute about a third. An interpremaxillary foramen is not definitely indicated; if present it must have been quite small. The palate narrows posterior to the incisors but expands again about the middle of the rostrum. The incisive foramina are of fair size and con- fined to the premaxillaries. The tooth rows diverge anteriorly; the palate is broadest in the region of the premolars. The posterior palatal notch extends anteriorly to the posterior region of M^; the palatines are short. A median ridge extends the length of the palate anteriorly as far as the anterior palatine foramina; it is flanked on either side by rather vague ridges bordering the alveoli of the cheek teeth. In the 1936 Burke: Ardynomys and Desmatolagus 139 Fig. 2. Ardynomys occidentalis Burke. Ventral view of skull, C. M. No. 1055. Left side of rostrum restored from opposite side. X 2. two channels separating these ridges the slit-like posterior palatine foramina are located opposite the first molars. Crushing and distortion makes the determination of some of the cranial foramina rather tentative. From its position in what appears to be the pterygoid fossa I take the foramen which shows in ventral view to be the sphenopterygoid. A foramen showing external to the fossa, and dorsal to the last, which is directed posteriorly in the alisphe- noid, would appear to be the masticatory. In advance of the latter appear two openings in the region of the sphenoid fissure; the posterior of these must be the anterior exit of the sphenopterygoid canal, or what is more likely, the sphenopterygoid -j- the alisphenoid canals. The opening in advance of this seems to be the sphenoid fissure proper. Further forward, and well separated from the last, is found the prominent optic foramen. The incisors of this form are especially large when compared with the cheek teeth. The superior incisors are flattened along the anterior surface (showing only a slight rounding) and are a little more extended transversely than antero-posteriorly. On the sides of the rostrum the courses of the incisors are indicated by well-defined incisive swellings. 140 Annals of the Carnegie Museum VOL. XXV In the skull, C. M. No. 1055, the cheek teeth anterior to the molars are not shown. However, in a maxilla, C. M. No. 1083, is preserved, although somewhat broken, and the root of is also retained. The latter tooth was quite small — merely a slender spike which rested Fig. 3. Ardynomys occidentalis Burke. Occlusal view showing root of P^, and worn Mi‘2 right, C. M. No. 1083. X 4. along the antero-internal side of Pk P'^, on the other hand, was the largest tooth in the maxilla. As exhibited in this specimen it is badly worn, showing a broad shelf of wear on the internal side. Externally, however, most of the original crown structure can be made out. Traces of the anterior and posterior valleys remain; the posterior valley (now represented by a shallow and small fossete) was ap- parently the larger of the two. Worn cusps represent the paracone and the metacone, from which converging lophs extend toward the worn internal shelf. Separating these lophs appears a broad V- shaped central valley, the outlet of which is blocked by two low intermediate tubercles conjoined at their bases. In the skull, C. M. No. 1055, the alveolus for P^ indicates a three- rooted tooth commensurate in size with P^ of C. M. No. 1083. In advance of the internal root socket of this tooth appears the alveolus for a small tooth occupying the same position as P^ in C. M. No. 1083. The cheek teeth of Ardynomys occidentalis m. combine unilateral hypsodonty (by extension rootward of the crown enamel of the protomere) with oblique implantation (the plane of the superior tooth row slants outward, that of the inferior tooth row slants inward, in both cases strongly oblique to the vertical plane of the skull). The combined effect of these dental constructions is the conservation of the enamel of the tooth with wear. With age, the teeth of Ardy- nomys occidentalis m. show marked broadening of the protomere shelf, the crown surface of which is worn away long before that of the metamere. The molars of the skull, C. M. No. 1055, show little wear. Seen at the crown surface they are subrectangular in outline. They taper somewhat rootward (especially M^) when viewed internally. At the 1936 Burke: Ardynomys and Desmatolagus 141 present stage of wear have the same crown dimensions, but M^, being more rounded below, might be expected to vary from at a later stage of wear. is reduced in size over The molar pattern on the external side follows the same general plan as that described for P^, consisting of anterior and posterior Fig. 4. Ardynomys occidenlalis Burke. Occlusal view of C. M. No. 1055. X 4. cingular crests, separated by anterior and posterior valleys from the protoloph and the metaloph. The latter lophs converge and extend to the internal shelf in the form of a sharp V, enclosing between them the central valley. The shelf is not strongly bowed in outline at the crown surface, though, a feature which contributes to the sub-rec- tangular outline of these teeth. In all of the molars the metaloph is somewhat oblique to the protoloph. The external outlets of the anterior and posterior valleys are nearly completely dammed off (except in M^, where the posterior valley exit cuts rather deeply), and are short notches considerably above the level of the central valley exit. The exits of the posterior valleys are in reality postero- external, internal to the metacones. The posterior valley extends further internally in these molars than does the anterior valley. The paracone and the metacone are nearly the same size in but the paracone is in excess of the metacone in There is a small intermediate tubercle in the outlet of the anterior valley of RM^ which is not present in that place in LM^. There is an intermediate tubercle in the outlet of the central valley of which forms a dam; the same condition holds in M^. In the intermediate tubercle is not present, but a small tubercle buds off the posterior flank of the metacone, somewhat crowding the exit of the posterior valley. A definite metaconule is not shown in or in although some swelling of the metalophs probably indicates it. In it is indi- cated as a vanishing tubercle on the unworn metaloph. Incidentally, the metaloph in IVP sinks below the level of the triturating surface before connecting with the internal shelf — -a condition which may have held, to some extent, in the metalophs of before they were worn down. On the internal side, the molars show slight anterior 142 Annals of the Carnegie Museum VOL. XXV and posterior enamel grooves or notches, between which the proto- cone swells internally. The lower jaw, except for its smaller size, is remarkably close to that of Ardynomys olseni M. & G. In both cases the jaws are heavy and are deep anteriorly; the masseteric fossa is not in advance of Mi. Ventrally the symphysis is extended downward to produce a sharp flange-like projection. There is a large mental foramen, below and just anterior to P4. The American form agrees with Ardynomys olseni M. & G. and Ardynomys chihi M. & G., in showing the ramus much pinched from the external side in the region of the diastema. The diastema has about the same relative extent as in Ardynomys olseni M. & G. The lower incisor is very much like that of Ardynomys olseni M. & Go It shows the flattened anterior face characteristic of the genus, and the incisor of the holotype, C. M. No. 1056, shows the shallow grooving or concavity on the anterior face, noted by Matthew & Granger.® This grooving is not apparent in the incisors of the referred specimen, C. M. No. 1105. As in Ardynomys olseni M. & G. the mandibular cheek teeth are not only oblique to the antero-posterior plane of the ramus, the Fig. 5. Ardynomys occidentalis Burke, holotype. Occlusal view of P4. M1-2 right, C. M. No. 1056. X 4. tooth rows converging posteriorly, in keeping with the maxillary tooth rows, but also they are implanted obliquely to the vertical plane of the ramus, and overhang on the internal side. P4 of Ardynomys occidentalis m., as shown in the holotype, bears on the trigonid two cusps, which are separated by a notch-like median valley anteriorly. The internal of these two cusps is the larger and doubtless was, before wear, the higher of the two, as in Ardynomys olseni M. & G. Its posterior flank is considerably drawn out to meet with the base of the postero-internal cusp, raising the internal outlet ®Matthew, W. D. and Granger, Walter, “New Creodonts and Rodents from the Ardyn Obo Formation of Mongolia.” Amer. Mus. Novit. No. 193, p. 5, 1925- 1936 Burke: Ardynomys and Desmatolagus 143 of the central basin high above the floor of the basin. The antero- and postero-external cusps are connected by an ectolophid. The hypoconid is an over-sized, jutting cusp, becoming a shelf with wear. From it extends a hypoconulid crest which sweeps around to join with the entoconid. The central basin is broad and rounded, and there is no definite evidence of any crests from the trigonid cusps within it. Also, at first glance, there appears to be no hypo- lophid crest present. However, on close inspection it can be detected forming the posterior boundary of the central basin. It is low, curl- ing parallel with the hypoconulid crest, and is evidently being elimi- nated, although it still shows a connection with the entoconid. The posterior valley is little more than an entrenched shelf behind it. The illustration of the type of Ardynomys olseni M. & G.® does not depict a hypoiophid in P4. I have not checked the accuracy of the drawing, but P4 of A. M. No. 20369, a specimen referred to Ardynomys olseni M. & G. shows a hypoiophid very much like that in P4 of the specimen under description — with the exception that it is perhaps better de- veloped. I have at hand also one of the paratypes of Ardynomys chihi M. & G., A. M. No. 20372, which shows a stubby but well- defined hypoiophid in P4. The illustration of the holotype of this species does not indicate a hypoiophid in P4 either.'^ The cheek teeth increase in size from P4 to Ms; M3 was considerably smaller than Ms (M3 is represented only by the roots in both the holotype, C. M. Nd. 1056, and in the referred specimen, C. M. No. 1105). The molars of C. M. No. 1105 are so worn as to obliterate all details of the crown pattern at the occlusal surfaces. The holotype, however, still preserves the central valleys on Mi-s; the valleys have a V-shape with wear, and at their outlets are flanked by the entoconids and metaconids; the entoconids, on P4, Mi and Ms, do not extend as far internally as do the metaconids. On Mi of C. M. No. 1056 I can detect the last traces of the posterior valley. Judging from what is preserved of the pattern, though, and from the con- figuration of the teeth in general, the unworn crown must have re- sembled rather closely that of Ardynomys olseni M. & G. In general construction the molars as now preserved, except for more transverse trigonids, resemble P4. The oversized hypoconids are broad shelves, which bore the brunt of wear. The hypoconid juts strongly externally ^Ibid, text fig. 8 (p. 6). "^Ibid, text fig. 9 (p. 7). 144 Annals of the Carnegie Museum VOL. XXV in Ml (more so than in P4) and still more prominently in M2. Not only is the hypoconid a greatly enlarged element of the crown in these teeth, but also the enamel extends disproportionately down along its enlarged root; one is tempted to call this “unicuspal,” rather than unilateral, hypsodonty. The above characters might well serve as an enumeration of those which also show in the worn lower cheek teeth of Ardynomys olseni M. & G. or of A. chihi M. & G. To a considerable degree they also apply to those of P seudocylindrodon neglectus m. and to Sespemys thurstoni Wilson. In a previous paper,® I have pointed out some of the differences which exist between Ardynomys and Pseudocylin- drodon : “In some respects, P seudocylindrodon is intermediate between Cylindrodon and Ardynomys. The cheek teeth are nearer those of Ardynomys in crown height and their tendency toward hypertrophy of the hypoconid regions; the tooth pattern in general shows simi- larities. The incisor of P seudocylindrodon is not flattened, however, as it is in Ardynomys] the central basins of the cheek teeth are deeper in P seudocylindrodon, have rounded floors, and are closed internally by the upgrown intermediate cuspule. The jaw of Ardynomys is deeper anteriorly than that of P seudocylindrodon, the posterior basins of the cheek teeth of Ardynomys have internal exits, and the pro- tolophids are less transverse than in P seudocylindrodon.'’'’ It might be added to the above, however, that Ardynomys chihi M. & G. shows the central valley of M3 closed off on the internal side. Matthew and Granger state that it is “closed by a marginal inner crest.”® On M3 of one of the paratypes, A. M. No. 20371, the intermediate tubercle is still indicated as an element in this crest — so that the closure of the central valley in this tooth has come about in essentially the same manner as in the inferior molars of Pseu- docylindrodon neglectus m. The intermediate tubercle is still indi- cated on Mi_2 of A. M. No. 20371, also, where it is fused below with the posterior flank of the metaconid. Incidentally, this same paratype shows an accessory cuspule in the posterior valley of M3, and a spur crossing the central valley from the hypolophid to the metaconid in ®Burke, J. J., “Pseudocylindrodon, a New Rodent Genus from the Pipestone Springs Oligocene of Montana.” Ann. Cam. Mus. Vol. 25, Art. i, p. 3, 1935. ®Matthew, W. D. and Granger, Walter, “New Creodonts and Rodents from the Ardyn Obo Formation of Mongolia.” Amer. Mus. Novit. No. 193, p. 7, 1925. 1936 Burke: Ardynomys and Desmatolagus 145 the same tooth. This spur had its origin in two tubercles, a small one on the anterior side of the hypolophid, a larger one at the base of the metaconid; the two have fused, but are still indicated. In M2 the tubercles are present but not joined. They are also apparent in Ml, but are rather poorly shown. Another paratype, A. M. No. 20372, preserves P4, and as I have mentioned previously in this article, a short hypolophid is present. The entoconid in P4 of this specimen is quite well-defined and attenuated externally to meet the hypolophid. Apparently Ardynomys chihi M. & G. is a well established species, quite distinct from Ardynomys olseni M. & G., and probably, as re- gards the pattern at least, even further removed from the latter species than is Ardynomys occidentalis m. At the time of the description of Pseudocylindrodon neglectus m. I had not seen Wilson’s excellent description and illustrations of Sespemys thurstonid^ Sespemys, in several respects, shares characters with both Pseudocylindrodon and Ardynomys. Sespemys tends to- ward hypertrophy of the hypoconid as do Pseudocylindrodon and Ardynomys \ in depth of lower jaw it seems nearer Pseudocylindrodon than Ardynomys, it preserves the internal intermediate tubercle of the molars, found also in Pseudocylindrodon and in Ardynomys chihi M. & G., although the tubercle in Sespemys tends to be attenuated into the central valley as a sort of spur — a condition which I have not noted in Pseudocylindrodon and Ardynomys. The tubercle in Sespemys does not appear to fuse with the metaconid and the ento- conid to block the central valley as it does in Mi_3 of Pseudocylindrodon and in M3 of Ardynomys chihi M. & G. Sespemys preserves the protolophid in P4, the protolophid does not appear to occur in P4 of Ardynomys, there are only traces of it in P4 of Pseudocylindrodon, if it is indicated at all. Sespemys shows short molar protolophids which compare better with those of Ardynomys than with the molar protolophids found in Pseudocylindrodon. The hypolophid seems well developed in P4 of Sespemys-, it is present, but shows poor develop- ment, in P4 of Ardynomys-, it is absent in Pseudocylindrodon. The posterior valley of the molars is open in Sespemys, open but crowded in Ardynomys, definitely closed in Pseudocylindrodon. In depth of valleys, the molars of Sespemys appear to be nearer to those of Ardy- ^°Wilson, Robert W. “Two Rodents and a Lagomorph from the Sespe of the Las Rosas Hills, California.” Cam. Inst. Wash., Publ. 453, No. 2, pp. 13-16, text fig. I, pi. I, figs. 2, 2a, 2b, 1934. 146 Annals of the Carnegie Museum VOL. XXV nomys, the molar valleys of neither of those two forms seem as deep as those of Pseudocylindrodon. The anterior face of the incisor is flattened in Sespemys and in Ardynomys, rounded in Pseudocylindrodon. All three genera agree in showing an enlarged central valley and re- duced posterior valley, and in having hypolophid crests that are transversely shortened. From the above intermingling of characters I conclude that Sespe- mys, Ardynomys and Pseudocylindrodon are rather closely related genera. Wilson has observed, in Sespemys, and Matthew and Granger have pointed out,^^ Ardynomys various Ischyromyid features. Matthew and Granger, stated^^ of Ardynomys, that its teeth “retain the partly subcircular outlines of Tillomys and have not taken on the rectangular proportions of Ischyromys, but the relationships of the genus are probably with this family, as Troxell has also urged for Tillomys. When distinguishing Pseudocylindrodon from Cylindrodon, I found the two genera had much in common with Ardynomys, and remarked that ^^Cylindrodon, Pseudocylindrodon and Ardynomys are probably Oligocene representatives of the stock of rodents typified in the Bridger Eocene by the forms which Troxell has included under Tillomys.^’’ To the “Oligocene representatives” I believe that we should now add Sespemys. In deriving these genera from the Tillomys stock, and in suggesting their interrelationships, I am basing my conclusions on certain funda- mental cusp-crest details of the tooth pattern which characterize Tillomys and Tillomys-W^e rodents of the Eocene — characters which in these Oligocene forms have undergone alterations, modifications of the Tillomys dental mechanism that fitted the group for invasion of certain environmental locales which during the Eocene appear to have been generally the province of various rodents of the Paramys group. Possibly, during the Eocene, the Tillomys stock was pre- dominately arboreal, its diet principally one of seeds and fruits; the same may have been true for the majority of the Paramyidce. But i^Wilson, Robert W., “Two Rodents and a Lagomorph from the Sespe of the Las Rosas Hills, California.” Cam. Inst. Wash., Publ. 453, No. 2, p. 15, 1934. i^Matthew, W. D. and Granger, Walter, “New Creodonts and Rodents from the Ardyn Obo Formation of Mongolia.” Amer. Mus. Novit., No. 193, p. 5, 1925. ^Hbid. i^Burke, J, J., “Pseudocylindrodon, a New Rodent Genus from the Pipestone Springs Oligocene of Montana.” Ann. Cam. Mus., Vol. 25, Art. i, p. 3, 1935. 1936 Burke: Ardynomys and Desmatolagus 147 by Bridger time certain of the members of the Paramys stock de- veloped specializations of the dental mechanism and skeleton which point to a terrestrial life. In Matthew’s Mans group^^ characterized by “broad robust molars” and wide incisors we find a departure from the more typical Paramyid dentition, and an increase in body size, which might indicate trends toward a coarser vegetable diet and better adaptability for a semi-arboreal, perhaps terrestrial, existence. Later, in the Uinta, we find Ischyrotomus (which Matthew suggested may have been derived from the Mans group^*^) attaining maximum size for the Paramyidce; its skeleton indicates not only a terrestrial but also even a fossorial habitat. Ischyrotomus, incidentally, fur- nishes an instructive comparison with Ardynomys. In both genera the skulls are stoutly constructed and robust, and Matthew’s de- scription of the muzzle of Ischyrotomus peter soni as “peculiarly angular” might also apply to the skull of Ardynomys. The lower jaws in both genera are heavy. Ardynomys olseni M. & G. and occidentalis m. develop a flange-like downward extension in the symphisial region of the mandible not altogether unlike that in Ischyrotomus. The incisors are flattened in both genera. The broaden- ing of the molar basins is of course evident in Ardynomys as well as in Ischyrotomus, hypertrophy of the hypoconid region characterizes the lower molars of Ischyrotomus, and I might add that there is some unilateral heightening of the cheek tooth crowns in Ischyrotomus also. Yet the possession of these characters by the two genera in common seems nothing more than another interesting case of paral- lelism; they are to be found in various combinations in a number of terrestrial or fossorial rodents, both living and extinct. Ardynomys is removed from any close relationship with Ischyrotomus through the cusp-crest pattern of its cheek teeth, already complicated beyond that of Ischyrotomus before being secondarily modified along the same lines as in the Paramyid. With the passing of the specialized Paramyidce at the close of the Eocene it is reasonable to suppose that faunal places were open for the invasion of the Ischyromyidce which, even in the Bridger, gave evidence i^Matthew, W. D., “On the Osteology and Relationships of Paramys, and the Affinities of the Ischyromyidae.” Bull. Amer. Mus. Nat. Hist. Vol. 28, Art. 6, p. 50, 1910. ^^Ibid, phylogenetic chart, p. 51. ^Ubid, pp. 56-57. 148 Annals of the Carnegie Museum VOL. XXV of considerable plasticity. Ardynomys and its relatives probably replaced the hians group of the Paramyidce in the North American Oligocene, at least, and Ardynomys seems particularly well fitted to have superseded Ischyrotomus, with which it possessed a number of habitus characters in common. MEASUREMENTS Holotype, Carnegie Museum No. 1056 mm. Diastema 5.0 Incisor antero-posterior 3.2 Incisor transverse 2.6 P 4 antero-posterior 2.3 P 4 transverse 2.2 M 1 antero-posterior 2.2 Ml transverse 2.8 M 2 antero-posterior 2.6 M 2 transverse 2.7 Tooth row at alveoli lo.o Molars at alveoli 7.7 Carnegie Museum No. 1105 Diastema - 5.2 Incisor antero-posterior 3.7 Incisor transverse 2.8 Ml antero-posterior 2.5 Ml transverse 3.1 Tooth row at alveoli 10. o Molars at alveoli 7.5 Carnegie Museum No. 1055 Height of skull measured at M^ 13-5 + Greatest width of muzzle 13 0 Width of interorbital constriction 6.9 Diastema 8.4 Width of palate at P® 4.9 With of palate at M^ 4.2 Length of palate from incisors to post-palatal notch 10.3 Incisor antero-posterior 2.4 Incisor tiansverse 2.5 M^ antero-posterior 2.1 Ml transverse 2.3 M2 antero-posterior 2.1 M2 transverse 2.3 M^ antero-posterior 1.6 1936 Burke: Ardynomys and Desmatolagus 149 mm. transverse i . 6 Tooth row at alveoli 9.1 Molars at alveoli 6.4 Carnegie Museum No. 1083 antero-posterior 2.5 transverse 3.8 Ml antero-posterior 2.0 Ml transverse 3.0 M2 antero-posterior 2.1 M2 transverse 2.8 Order DUPLICIDENTATA Illiger Family LEPORID.F Gray Genus Desmatolagus Matthew and Granger Desmatolagus dicei* sp. nov. Holotype: Left maxillary with P^"^, Mi"^; right ramus of mandible with P4, Mi_3; left ramus of mandible with Mi_2; C. M. No. 814. Horizon: Pipestone Springs Oligocene. Locality: Pipestone Springs, Jefferson County, Montana. Diagnosis: Species larger than Desmatolagus robustus M. & G.; premolars not as well developed, molars less reduced; external outline Fig. 6. Desmatolagus dicei Burke, holotype. Ventral aspect of left maxillary with p3-4^ Mi'2, occlusal view of P4, M1-3 right and lateral view of right ramus of mandible, C. M. No. 814. X 2. *This species is named in honor of Dr. Lee R. Dice, in appreciation of his works on fossil Duplicidentates. VOL. XXV 150 Annals of the Carnegie Museum of maxillary tooth row more evenly rounded; M2 the largest inferior cheek tooth. Desmatolagus gazini* sp. nov. Holotype: Left maxillary with M^^^. q ]\/[^ Horizon: Oreodon Beds Oligocene. Locality: Badland Creek, Sioux County, Nebraska. Diagnosis: Species intermediate in size between Desmatolagus gobiensis M. & G. and Desmatolagus robustus M. & G.; external outline of maxillary tooth row less rounded than in Desmatolagus dicei m. and premolar-molar proportions nearly as in Desmatolagus robustus M. & G. and Desmatolagus gobiensis M. & G., but not as compressed along its anterior side and M'^ with unusually short posterior wall crest, external roots of superior cheek teeth stronger. Fig, 7. Desmatolagus gazini Burke, holotype. Ventral aspect of left maxillary with P3-4, Mi-2, C. M. No. 37. X 2. It would appear that species of Desmatolagus in the North American Oligocene may be distinguished from contemporary species of Mega- lagus by their lower-crowned and more transverse cheek teeth, re- duced Mf and somewhat simpler P^. In the lower Oligocene the excess in size of M2 over Mi and P4 may also prove characteristic of Desmatolagus, but at present I am not inclined to place much trust in this peculiarity. It may be anticipated that when the third lower premolars of Desmatolagus dicei m. and D. gazini m. are found they will also show more reduction and secondary simplification than in Megalagus, while P^ may be expected to show more reduction also. While Desmatolagus in the lower Oligocene in general resembles Megalagus, the two genera do not keep apace in their later develop- ment. Middle and Upper Oligocene species of Megalagus strongly *The specific name is in recognition of the fossil lagomorph studies of Dr. C. Lewis Gazin. 1936 Burke: Ardynomys and Desmatolagus 151 parallel Palceolagus in the “modernization” of their tooth structure. Desmatolagus, however, continues the reduction of M| and if not the reduction, certainly the simplification of P3; appears to be re- duced and there are changes in the pattern of P^ (this tooth also under- goes compression along its anterior face). There is some equalizing of the proportions of P^”^ and perhaps better expressed as due to the increase in size of the premolars concerned. There seems to be, as regards the premolars at least, an increase in hypsodonty (Teil- hard de Chardin^* even describes P| of his Desmatolagus pusillus as of continuous growth). But it is noteworthy that Desmatolagus also preserves, in the Upper Oligocene, an assemblage of characters which are found in the Eocene Mytonolagus and also in specimens of Mega- lagus from the lower Oligocene — curved superior cheek tooth shafts, relatively poorly developed unilateral hypsodonty, transverse cheek teeth and persistence of the crown pattern with wear. Certain of the above characters have doubtless been interpreted by some authors as indicative of Ochbtonid affinities of Desmatolagus. To my way of thinking they are merely retained primitive characters, indicative of the unprogressiveness of the genus as a whole. That lower jaws of the type which VireP® describes and figures as of Amphilagus antiquus closely resemble those of Desmatolagus goes without saying, and judging from what I can learn of these forms they must have a close relationship with Desmatolagus. But from the same deposits Viret has also described and figured^*^ certain maxillaries upon which he bases his genus Piezodus, and these maxil- laries also resemble those of Desmatolagus. On the other hand, the maxillaries which Viret refers to Amphilagus antiquus, and the lower jaws which he includes under Piezodus brannsatensis Viret, do not appear at all to be of the Desmatolagus type. Unfortunately, I do not possess any of these European forms for comparison with Desmatolagus, and I can only judge them from the scant literature at my disposal. I have the feeling that there has been some confusion of genera in Viret’s treatment of his material. The lower jaws which i^Teilhard de Chardin, P., “Description des Mammiferes Tertiaries de Chine et de Mongolie.” Ann. Pal. tome XV, p. 23, 1926. i®Viret, J., “Les Faunes de Mammiferes de I’Oligocene Superieur de la Limane Bourbonnaise.” Ann. Univ. Lyons, N.S. fasc. 47;tpp. 85-88, et seq. text fig. ii, PI. 29, figs 12, 13a, 13b, 14, 1929. ^^Ibid, text fig. 14, b, b’, c, c’, d, e, PI. 29, figs. 15a, 15b, pp. 94-96. 152 Annals of the Carnegie Museum VOL. XXV he refers to Amphilagus antiquus, and the maxillaries which he places in Piezodus brannsatensis Viret might possibly belong in the same genus or even species. The resemblance of this material to Desma- tolagus is so well marked that I suspect that here we have European representatives of the Desmatolagus stock. But it does not follow, of necessity, that the mere presence of Desmatolagus-\\]^e. forms in the European Oligocene presumes a “Lagomyid” and therefore Ochotonid disposal of the Desmatolagus line. From present evidence, Desmato- lagus (and Viret’s material mentioned immediately above) appear to me to have far too many characteristics of the Leporidm to be ancestral to any Ochotonidce or to warrant their inclusion in the latter family. Desmatolagus is already specialized along the lines of the Leporidce in the pattern of its cheek teeth. Despite the marked persistence of the internal valley in P4 and in the inferior molars (a condition which, by the way, can be found in certain species of Palceolagus, even in the Upper Oligocene) these inferior cheek teeth are thoroughly leporine as regards their construction; they show the characteristic crowding of the paramere cusps and a main valley extending transversely across the tooth from the protomere which separates the trigonid from the talonid. In the OchotonidcB the paramere cusps are widely separated, and while both the internal and the external valleys appear to have conjoined to form the separation between the trigonid and the talonid, it is of note that the external valley is directed postero-internally, while the internal valley is directed antero-internally, becoming con- fluent with the external valley by cutting across the latter in advance of its postero-internal channel. The superior molars of Desmatolagus show a narrow internal notch or valley, such as characterizes the Leporidce generally, and the course of the valley is postero-external, or at the most transverse, not in advance of the “crescent”; the external portion of the valley becomes isolated as an enamel island as in Mytonolagus, Megalagus, and PalcBolagus. In the Ochotonidce the internal cusps of the molars are well separated by a wide internal valley which is antero-externally directed, and which in the earlier forms is seen to be directed anterior to the “crescent.”^! 2iBut this “crescent,” or main lunate valley, in the Ochotonidce may be an en- tirely different crown element from that called the “crescent” in the Leporidce. J, Ehik, in Ann. Mus. Nat. Hungarici, Vol. 23, pp. 178-186, figs. 1-5, 1926, de- {Continued on next page) 1936 Burke: Ardynomys and Desmatolagus 153 The type of pattern torsion exhibited in the OcJiotonidcB is not without parallels among the simplicidentate rodents; something very much like it is found among the Castoridce, where it is associated with considerable preservation of the original elements of a complex talon- talonid pattern. I am inclined to think that the Ochotofiidce, too, retained some of the talon-talonid elements of the crown pattern longer than did the LeporidcB. Atrophy and fusion of the various crown elements of these regions appear to have been early develop- ments in the LeporidcB, and certainly must have had their initiation well back in the history of the group, as a preliminary to the attain- ment of the basic leporine pattern which is already well established by Upper Eocene time. (MEASUREMENTS ON FOLLOWING PAGE.) scribes and figures as Titanomys fontannesi two maxillaries which preserve excel- lent details of the crown pattern. In and of his material Ehik found what appears to be the last trace of an anterior valley in advance of the main lunate valley. The internal valley in his specimens is directed antero-externally, as in Ochotonidce generally, and is anterior to the main lunate valley, but is pos- terior to the presumed vestige of an anterior valley. I do not agree with Ehik’s interpretation of molarization of the premolars in the Duplicidentata, and I am not in accord with him in his cusp terminology in general as he has applied it in this case, as I have stated in a previous article (Ann. Cam. Mus., Vol. 23, Art. 9, p. 408, 1934). But if the structure anterior to the lunate valley in and of Ehik’s specimens is a vestigial anterior valley, then it must correspond to the anterior valley in the Leporidoe, and the wall which separates it posteriorly from the main lunate valley may very well represent the paracone, with which Ehik identifies it. Such a disposition of the paracone would seem to accord very well with the evident pattern torsion in the Ochotonidce. If this construction, or one similar to it, can be shown to have been a stage in the evolution of the pattern of P^ and in the Ochotonidce generally I need not point out to students of the Duplicidentata that it not only explains the chief differences in superior cheek tooth pattern in the Ochotonidce and the Lepor idee, hut also it emphasizes the early separa- tion of the two Duplicidentate families. 154 Annals of the Carnegie Museum VOL. XXV MEASUREMENTS * Desmalolagus Desmatolagus dicei gazini C. M. No. 814 C. M. No. 37 antero-posterior 2.5 transverse 5.2 P4 antero-posterior 2.9 P'* transverse 6.3 Ml antero-posterior 2.5 M 1 transverse 6.5 M2 antero-posterior 2.6 M2 transverse 5.6 M2 transverse 5.6 P4 antero-posterior 3.9 P^ transverse 3.3 M2 antero-posterior 2.9 M„ transverse 3.1 M2 antero-posterior 3.3 M2 transverse 3.5 M2 antero-posterior 1.5 M2 transverse 1.6 Length of inferior molar series 10.6 2.15 3- 6 2 . 2 4- 3 2.15 4-3 1.8 3-4 3-4 'All measurements taken at triturating surfaces of teeth. OCCURRENCE OF THE FAMILY CARYCHIID^ IN WEST VIRGINIA , By Stanley Truman Brooks and Gordon m. Kutchka % Reprinted from the Annals of the Carnegie Museum Vol. XXV, p.lSS-161, 1937 ^ - Issued February 3, 1937 ART. XVII. OCCURRENCE OF THE FAMILY CARYCHIIDiT IN WEST VIRGINIA* By Stanley Truman Brooks and Gordon M. Kutchka (Text Figures 1-14) Introduction: For several years the Laboratory of Recent In- vertebrates of the Carnegie Museum has been increasingly active in the study of the molluscan fauna of West Virginia. Usually a study of the forms living within the boundaries of a political unit is value- less in regard to the distributional features involved. West Virginia, however, is unique. It is one of the last molluscan frontiers of the eastern states, and has only been slightly investigated by conchologists. But more important is the fact that West Virginia lies within two of the areas of migration for the northern dispersal of the so-called southern fauna. A full discussion of the distribution of the land Gastropoda will be taken up in the final papers of this series and the accumulated material will then be examined in the light of the ecology of the region studied. We hope that these efforts will in no way deter any student who wishes to take up a study of the molluscs of West Virginia. On the contrary, it should stimulate him to make a more precise survey of the field. Our methods of mass collecting are painstaking and exhaustive, but they undoubtedly leave much to be discovered in the future. Acknowledgments : In the list issued in 1935 (Brooks), mention was made of the Carnegie Museum Collection and those responsible for its accumulation. Since that time other students have been active and much more material has flowed into our hands. The greatest advance was made during the summers of 1935 and 1936 when the junior author made two trips of six weeks each with the University of West Virginia Biological Expedition. Through his tireless efforts of collecting and sifting forest loam, and through the regulation *This is the first in a series of studies being prepared by the Section of Recent Invertebrates. Others will follow covering the other families of the Land Gastro- poda. ■sued Feb. 3, 1937 1937 Brooks & Kutchka: Carychiid^ in West Virginia 156 methods of collecting, Mr. Kutchka has added over 16,000 specimens, mainly minute, to our collection. Professor G. R. Hunt of Fairmont Teachers College, Fairmont, West Virginia, also visited a few locali- ties during 1935 and his material has added several important stations Fig. I. Greater Diameter. Fig. 2. Lesser Diameter. Fig. 3. Height or altitude. Fig. 4. Greater Diameter. Fig. 5. Counting the number of whorls. Fig. 6. Aperture - L. = length, W. == width. and records. Mr. Neil Richmond collected at several new stations and has turned his material over to this laboratory for study. Others responsible for additions to our collection, although not to the ma- terial utilized in this individual paper, will be mentioned from time to time as the need arises. The above collectors will be designated as (GMK), (GRH), and (NDR), respectively, in the following pages. 157 Annals of the Carnegie Museum vol. xxv Genus Carychium Muller Some of the smallest of the land snails belong to this group. The shell is spiral, with an elongate or oval aperture; lip reflected, thick- ened, and appears toothed. The columellar folds begin at the lip and revolve within the whorls and in some species are visible ex- ternally. The shells somewhat resemble those of the PupillidcB but in the living animal the eyes occur below and behind the tentacles near their base. All of the species are terrestrial in habit and may be found living among damp leaves and at the roots of grasses. In some localities they are found near springs and nearly submerged in the water. Carychium exile H. C. Lea Shell ovate-conical, elevated, with five to five and one-half convex whorls, striated longitudinally or ribbed; apex obtuse; suture deeply impressed; aperture small, very oblique, one-third of the length of the shell; outer lip more or less thickened but may be slightly flattened; the columellar lamellae beginning at the aperture appear as two teeth, the upper one the largest, the lower reduced to a mere denticle, lamellae well developed within the body whorl; size of shell, altitude 1.70 mm., diameter 0.60 mm. Type Locality: “Wissahickon Creek, Philadelphia, Pennsylvania” (Lea, H. C. 1842). Range: Ontario, south to Alabama and west to Minnesota. West Virginia Records Braxton County, Gassaway (GMK) Greenbrier County, North Caldwell (GRH) Greenbrier County, Alderson* (Muddy Creek Valley) (NDR) Hampshire County, Romney (GMK) Harrison County, near Bristol (NDR) Kanawha County, Tornado (GMK) *Alderson occurs on County line. 1937 Brooks & Kutchka: Carychiid^k in West Virginia 158 Marion County, Kingmont (GRH) Marshall County, Bannon (NDR) Monroe County, Alderson* (GRH) Monroe County, Peters Mountain (GRH) Nicholas County, Summersville (GMK) Nicholas County, Lockwood (Panther Mtn.) (GMK) Pendleton County, Franklin (GMK) Pendleton County, Ruddle (GMK) Pendleton County, Judy Gap (GMK) Pocahontas County, Marlinton (GMK) Pocahontas County, Mill Point (GRH) (GMK) Randolph County, Huttonsville (GMK) Summers County, Talcott (GMK) Remarks: In order to clarify the descriptions of the columellar folds a part of Miss Winslow’s (1922) remarks may be appended here: “The upper columellar fold is very large in proportion to the dia- meter of the last whorl, almost touching the wall of the whorl at the upper angle of the aperture. Typically it is bent sharply downward at its widest part, the edge turning toward the columella. The lower fold is leaflike, curling at its edge, wider than in exigimm, and con- spicuously projecting from the columella beneath the upper fold. Both lamellae are more persistent in the penultimate whorl than in the case of exiguum, becoming wider in that whorl before disappear- ing in the upper whorls.” The typical northern Carychium exile (Brooks 1936) is not common in West Virginia, its place being taken, in great part, by a narrower form averaging 1.9 mm., in altitude, and 0.60 mm. in diameter. At first, this was thought to be the large form described by Dr. G. H. Clapp, Carychium exile canadense (1906), but dissection of the body whorl showed that the lamellae were like those of exile. How^ever, it was concluded that this form is a geographical race. The aperture. *Alderson occurs on County line. 159 Annals of the Carnegie Museum VOL. XXV texture of the shell and the proportions, all tend to separate it from the broader exile of Pennsylvania. The lamellae vary from the “typi- cal” exile form to one approaching that of canadense. The difference in length causes the “dip” of the upper lamella to vary its position relative to the aperture. It would almost seem that an addition of several tenths of a millimeter to some of these shells would produce the characteristic lamellae of canadense. The specimens from Prince- ton, Alabama, mentioned by Dr. Clapp (1906), are not this form but in their proportions and lamellae seem to be true exile. Carychium exiguum (Say) Shell tapering, oblong, with minute grooved lines; apex obtuse, whorls five; suture deeply impressed; aperture with two teeth, the superior one large, the lower reduced; lip flattened and reflected; umbilicus distinct. Usually a shining, quite transparent shell. Size of the average, altitude 1.75 mm., diameter 0.75 mm. Type Locality: “Vicinity of Philadelphia” (Say, Thomas 1822). Range: Ontario south to Alabama and west to Texas and Iowa. Remarks: This species may be distinguished from exile by the more inflated or “bellied-out” appearance of the body whorl and by the upper lamella which is much “smaller in proportion to the diameter of the last body whorl” than in exile, “and never deflected down- ward,” Winslow (1922). The shell when removed from the living animal is usually shining 1937 Brooks & Kutchka: Carychiid^ in West Virginia 160 and transparent, but when found weathered is opaque. There is a chemical change in the structure of the shell after the animal is gone and the shell exposed to the elements. Those in the collection ob- tained living have retained their transparency. The opaque shells have become thinner and more brittle and are easier to open for the observation of the lamellae. West Virginia Records Hampshire County, Romney (GMK) Surprising as it may be, this species seems to be uncommon in WTst Virginia. In this study, over forty large sacks of loam were collected, sifted, and the siftings examined under a lens. It does not seem possible that it has been overlooked by us. It will probably be found in other localities with further collecting as it appears to be fairly common in the surrounding areas. Carychium nannodes Clapp “In shape this species resembles C. exile, being long and slender, but differs in being absolutely smooth, without any traces of growth lines, even when magnified 6o diameters; under high magnification the surface shows a faint granulation; color waxy-white, transparent, the columellar fold showing distinctly through the shell; whorls about four and one-half, regular, tapering from the body whorl to the apex; sutures deep, whorls slightly shouldered; lip wide and well reflected especially at the columella where it forms a distinct umbilical chink, outer curve of the lip decidedly flattened, hardly thickened within; viewed from the back the lip is more squared below than in exile and exiguum; upper columellar fold of good size, lower one almost obsolete.” Clapp (1905). Size 1.4 mm., in altitude, and 0.5 mm., in diameter. Winslow (1922) describes the lamellae as follows: “Upper fold small in proportion to the size of the last whorl, evenly sinuate. Lower fold scarcely more than a cord, becoming somewhat flattened and slightly projecting during the course of the first turn.” 161 Annals of the Carnegie Museum VOL. XXV Type Locality: Monte Sano, near Huntsville, Alabama (Clapp 1905)- Range: Alabama to West Virginia. West Virginia Records Greenbrier County, Renick (GRH) Marion County, Kingmont (GRH) Monroe County, Alderson (GMK) Pendleton County, Franklin (GMK) Pocahontas County, Mill Point (GRH) Summers County, Talcott (GMK) Summers County, Wolf Creek (GMK) Remarks: This form is easily distinguished from the other species by its size. It is smaller than any other American member of the genus. The ribless, granulated appearance of the shell also dis- tinguishes it from the other more northerly forms. The finding of this species in West Virginia greatly extends its range, the exact limits of which are unknown. BIBLIOGRAPHY Brooks, S. T. 1935. Annals of the Carnegie Museum, Vol. XXIV, p. 61-68. 1936. The Nautilus, Vol. 49 (4), p. 115. Clapp, G. H. 1905. The Nautilus, Vol. XIX, p. 91, pi. Ill, figs. 7, 8, 9. (C. nannodes) . 1906. The Nautilus, Vol. XIX, p. 138-140, pi. VHI, figs. i-io. (C. e. canadejise) . Lea, H. C. 1842. American Journal of Science, (i), Vol. XLII, p. 109, pi. I, fig. 5. (C. exile). Say, Thomas. 1822. Journal of the Academy of Natural Sciences, Vol. II, p. 375. (C. exiguum). Winslow, Minna L. 1922. Occasional Papers of the University of Michigan, No. 128, p. 1-16, pi. 1-5. HYBRIDIZATION AND THE PHYTOGENY OF THE GENUS PLATYSAMIA ■ i ■ k By W. R. Sweadner Vi ^ r ■ 'i I C 4 k/ r . i - ' sv:\ k- ! _r - ‘ k Reprinted from the Annals of the Carnegie Museum Vol. XXV, p. 163-242, 1937 Issued May 15, 1937 ^ ART XVIII. HYBRIDIZATION AND THE PHYLOGENY OF THE GENUS PLATYSAMIA By W. R. Sweadner (Text Figures 1-5; Charts I-III) (Plates XV- XX) TABLE OF CONTENTS I, Introduction 167 Selection of the Problem 167 Resume of Previous Work on Lepidoptera Hybrids. 167 Work of Standfuss 167 Work with the Sphingid hybrids 168 Work of Harrison and his colleagues 168 Classification of hybrids. 169 Work of Pictet 170 The bearing of all these on the present problem. . . 170 Preceding work with the genus Platysamia 170 II. Material 171 The Genus Platysamia 171 Descriptions and Distribution of the Forms 173 P. cecropia (Linnaeus) 173 P. gloveri (Strecker) 176 P. Columbia (Smith) 179 P. nokomis (Brodie) 181 P. winonah (Brodie) 183 P. euryalis (Boisduval) 184 P. kasloensis (Cockerell) 185 P. cedrosensis {CoehereW) 187 Aberrations 187 Summary 188 HI. Methods 190 Methods of Collecting and Cross Breeding 190 Methods of Rearing 192 Methods of Pattern and Color Analysis 193 Methods of Genitalic Analysis 194 Summary 195 163 Issued May 15, 1937. 164 Annals of the Carnegie Museum vol. xxv IV. Observations. 195 The Breeding Experiments 195 Time of flight and normal matings 195 Record of experimental cross matings 196 Cross matings in confinement 196 Cross matings in the field 198 Matings of hybrids and secondary hybrids 201 Summary 201 Viability of the Hybrids 201 Per cent of Hatch 201 Factors in larval mortality 202 Summary 204 Morphology and Biology of the Hybrids 204 Larval inheritance 204 Adult color and pattern 205 Size and vigor 205 Sex ratio and fertility 205 Summary 207 Zones of Contact and Intergradation 207 The northwestern contact 207 The southwestern contact 209 The Canadian contacts 210 Matching of intergrades with hybrids 212 Summary * 213 Genitalic Analysis 214 V. Discussion 216 VI. Conclusions 224 VH. Appendix,. 225 VHI. Bibliography 228 FOREWORD It would require too much space to acknowledge individually all of the many persons who have assisted in the work recorded in this paper. The writer wishes to thank those hospitable people who helped him during his survey trips, the many entomologists throughout the United States and Canada who have supplied information and ma- terial, and the scientific workers in the University of Pittsburgh and the Carnegie Museum who have generously given their time and advice. The writer particularly wishes to acknowledge his indebtedness to Dr. Andrey Avinoff who has directed this work and whose inspira- tion, suggestions, and criticisms have been the principal forces in shaping this research; also to express his gratitude for time, instruc- tions, and criticisms graciously given and for material loaned by the late Mr. Foster H. Benjamin and his associates in the United States National Museum; and to declare his appreciation to Dr. Robert T. Hance and his colleagues in the Department of Zoology of the University of Pittsburgh for counsel and guidance during the progress of the investigation. 165 I. INTRODUCTION A. The Selection of the Problem The work presented in this paper is a by-product of an investiga- tion into the underlying principles involved in the wing patterns of the Lepidoptera. Hybridization was proposed as a tool with which to pry into the secrets of this tremendously varied expression of nature’s art. The apparent duplication of homologous markings in a hybrid between Mimas tilice and Smerinthus ocellata suggested this line of attack. Accordingly a considerable number of species belong- ing to the families Saturniidae and Sphingidae were used in attempts to produce hybrids. The ease with which cross matings were ob- tained in the genus Platysamia (Grote) = Sarnia (authors) of the former family led to speculation as to what would happen if the ranges of two species should overlap. A careful search of the literature and examination of specimens in the local collections yielded clues which might be construed as indicating that hybridization was taking place in nature, but no concrete evidence was uncovered. A five year study of this genus both in the laboratory and in the field has dis- closed facts which may be of considerable importance in moulding our concepts of the much debated “Species Problem.” B. Resume of the Previous Work on Lepidoptera Hybrids A very considerable amount of work has been done on hybridiza- tion in the Lepidoptera covering a period of more than fifty years. It can be grouped conveniently into three classes: first, the pioneer work of Standfuss and his colleagues on hybridization in the genus Saturnia; second, the very extensive work with the Sphingidae of Europe and to a lesser extent of America; and third, the work of Harrison and his colleagues with the Bistoninae and of Pictet with the races of butterflies and moths in the Swiss Alps. THE WORK OF STANDFUSS Dr. Standfuss of Vienna crossed the various species of the Pale- arctic genus Saturnia and obtained a low percentage of matings (they 167 168 Annals of the Carnegie Museum VOL. XXV do not cross in nature) and even smaller percentages of fertile eggs and of imagoes. A large number of his hybrid broods were composed of only one sex. His general conclusions may be summarized as follows, (i) The offspring tend to resemble the phylogeneticly older parent in morphological and physiological characters. (2) There appears to be a certain predominance of characters inherited from the male. (3) The females are sterile and the males more or less fertile. (4) The individuals of the second generation are very variable, tend to approach the phylogeneticly older parent, and include a con- siderable percentage of gynandromorphs. THE WORK WITH THE SPHINGID HYBRIDS. The principal workers with the Sphingid hybrids were Denso, Standfuss, Lenz, Kysela, Dannenburg, and Rebel. The crosses were made between the various species of the genera Celerio, Smerinthus, Mimas and Chcerocampa. Practically all possible combinations within the various genera were obtained and many intergeneric crosses made. Among these were a number of interfaunal crosses such as those in- volving Smerinthus ocellata or Amorpha populi of Europe as one parent and Smerinthus jamaicensis, S. cerysii, Paonias myops or P. astylus of North America as the other. Secondary and tertiary hybrids in all combinations were produced in many cases, particularly using the species Celerio galli and Celerio euphorhice. The principal contributions to the knowledge of hybrids made by this group of investigators were the production of intergeneric hybrids and the filling in of the gaps in the knowledge of the types of hybrids. THE WORK OF HARRISON AND HIS COLLEAGUES Harrison has attacked the problem of hybridization principally from the angle of phylogeny. By means of his experiments on the Bistoninae he was able to demonstrate that the fertility of the hybrid is a function of the specific divergence between the parents. It was also demonstrated that the phenomena of dominance in the hybrid depends on this same factor, being not evident when there is consider- able divergence. Harrison correlated the data from geographic dis- tribution with that obtained from hybridization to formulate the re- lationships and probable origin of the species in the genera Lycia, Pcecelopsis and Nyssia. Perhaps the most important contribution of Harrison and his co-worker, Doncaster, was the finding of an irregular 1937 Sweadner: Hybrids and Phylogeny of Platysamia 169 chromosome behavior in the hybrids. .This provides a possibile ex- planation for the infertility of some, for the gynandromorphism and for the queer sex ratios often obtained. THE CLASSIFICATION OF HYBRIDS As a result of these various experiments with hybrids of the Lepi- doptera, a classification of hybrids was proposed by Raepke. This classification, reported by Morgan in the American Naturalist, Volume XLIII, is reproduced here with an additional class added to each end. 0. Those hybrids which, although they undergo considerable embryonic differentiation, fail to hatch, or hatching, fail to complete the larval development, because of disharmony of parts. 1. “First, those hybrids that are so abnormal (atypic or sexless) that their sex cannot be determined. 2. “Second, those hybrids in which only one sex developes, gener- ally the male; females also rarely appear, but are so imperfect that reproduction is impossible. The males also are sterile. 3. “Third, those hybrids in which both sexes appear in normal proportions; the females sterile, the males crossed back to the parent species fertile in various degrees. The offspring of such a union are, however, very abnormal and monstrous both in their primary and secondary sexual organs. In certain series gynandromorphs appear in surprisingly large numbers. 4. “Fourth, those hybrids in which the females although appearing normal lay either no eggs or abnormal eggs. The males are like those of the last category or like those in the next. 5. “Fifth, those hybrids in which the females produce only embryos, or if the caterpillar stage is reached at all, the young are weak. When it has been possible to rear moths by back crossing these females to the parent species (or from male hybrids of the same cross) only males develope but in such scanty numbers that they have not been tested further.” To this we might add another class, 6. Those hybrids in which the fertility of both sexes approaches that of the parent species in that secondary, tertiary, or other hybrids can be obtained. Example: Celerio galli x Celerio eiiphorhicB. It may be further stated that these classes, while they show the manner in which the hybrids grade from one extreme to the other in 170 Annals of the Carnegie Museum VOL. XXV viability and reproductivity, are artificial groupings of a more or less uniform gradation. THE WORK OF PICTET The work of Pictet in his analysis of the Lepidopterous fauna of the Swiss National Park, while not directly involving species- hybridization is important in that he was able to show in a number of cases that intermediate forms found on the barrier between two subspecies could be duplicated by hybrid forms produced in the laboratory. He appears to be the first to demonstrate by synthesis the possible hybrid nature of an intergrade. THE BEARING OF ALL THESE ON THE PRESENT PROBLEM From the above incomplete and much abbreviated account of the previous work, on hybridization, it is hoped that the reader may have gleaned an understanding of the phenomena as they are manifested in the Lepidoptera. The experiments described in the following pages partake of the nature of all of those listed above, in that the relationships between the various species and forms in the genus are determined from the data obtained through hybridization as well as from distribution and comparative morphology, as in Harrison’s work; while the hybrid nature of certain intergrading forms is demon- strated through synthesis in much the same way that Pictet analyzed his intermediate forms in the Swiss Alps. C. Preceding Work with the Genus Platysamia The various species and forms of the genus Platysamia have been crossmated and the hybrids reared to the imago many times as re- corded in the literature. Every sizable collection in North America contains one or more of these hybrids. As yet no secondary hybrids have been reported. For a number of years after the publication of the description of Platysamia Columbia (Smith) = Sarnia Columbia (Smith), a controversy raged as to whether or not it was of hybrid origin. Dr. H. A. Hagen of Cambridge, Massachusetts, suggested that it might be a hybrid, possibly between P. cecropia (L) and Callosamia promethea (Drury). ^ This was soon disproven. Most of the published records of hybridization in this genus are mere isolated notes to the effect that a hybrid had been more or less ac- ^Hagen, H. A. “On Attacus (Sarnia) Columbia and its Parasites.” Bulletin of the Buffalo Society of Natural Science, Jan. 1875. 1937 Sweadner: Hybrids and Phylogeny of Platysamia 171 cidentally obtained. To my knowledge, the only careful study was made by Miss E. L. Morton of Newburg, New York.^ She had P. cecropia and P. gloveri emerge and mate in the same cage. The hybrids from this and from other crosses were reared to the imago stage and all were reported to be completely sterile. The best sum- mary of the early studies in this genus was made by Tutt in the third volume of his work “A Natural History of British Lepidoptera” in which he assigned names to Miss Morton’s hybrids and to those produced by Heyer.^ The names which are not recognized under the International code were P. griffithsi (Tutt) = P. cecropia cf x P. gloveri $ ; P. watsoni (Tutt) ==P. cecropia (T x P. euryalis 9 ; P. ameri- cana (Tutt)=P. Columbia cf x P. cecropia 9 ; and P. heyeri (Tutt) = P. euryalis cf x P. cecropia 9 . II. MATERIAL A. The Genus Platysamia The forms of insects herein discussed are referable to the scaly winged order Lepidoptera. Since they have bipectinate antennae they belong to the suborder Heterocera, the moths. The hind wings have the most posterior vein (3rd Anal or Ic) absent and the most anterior one (Sc-Ri or 8) diverging from the cell at the base. The fore wings have vein M2 or 5 arising from above the middle of the discocellular vein. In addition, they contain “no sensory organs of radiating bristles behind the antennae, no tympanal cavity at the base of the abdomen, no proboscis, no tooth on tarsal claw, and the frenulum is completely absent,”"^ placing them in the family Saturniidae. In the most recent check list of the North American Saturniidae^ they are placed in the genus Platysamia.^ The species of the genus Platysamia are characterized by having ^Morton, E. L. “Hybrid Saturniid Moths and their Larvae.” Proceedings of The Entomological Society of London, 1895, PP- 34-35- ®Tutt, J. W. A Natural History of British Lepidoptera, London, Vol. 3, pp. 292-293. ^Jordan, K. “Note on the families of moths in which M2 (vein 5) of the fore wing arises from near the center or above the center of the cell.” Novitates Zoologica, Vol. 30, 1923, pp. 163-166. ^Barnes, W. and Benjamin, F. H. “Check List of the Lepidoptera of Boreal America, Superfamilies Sphingoidea, Saturnoidea and Bombycoidea of the Families Syntomidae and Arctiidae.” Pan. Pacific Entomologist, Vol. 4, pp. 87-89, 1927. ®See Appendix 1. 172 Annals of the Carnegie Museum VOL. XXV bipectinate antennae, the pectinations of the male being much longer than those of the female; heavy furry bodies; and large broad wings. The markings common to all of the species consist of a basal and an extradiscal white cross band and a curved discal spot on both wings and an apical ocellus on the forewing. The eggs are ovate, white, covered with a gummy brown cementing material and are laid in groups of from three to six. The larvae are cylindrical and equipped with eight rows of warty tubercles armed with spines. The first seg- ment has six, segments two to five have eight, segments six to ten have six, segment eleven has only five because of the fusion of the tubercles representing the two dorsal rows, and segment twelve has only four. In the last instar the dorsal tubercles of segments two, three, four and eleven are large and swollen. The cocoon is densely woven, composed of an inner and an outer shell, is open at one end, and is rigidly attached to some support, usually a branch of the food plant. The first to be described, of the species concerned in this report, was PhalcBna cecropia (Linnaeus) 1758. Hiibner placed it in his generic subdivision Sarnia"^ along with cynthia (Drury) and promethea (Drury) in 1816. In 1855 Walker redescribed the genus Sarnia^ and dropped cynthia’ (Drury) from the list of its species. In 1865 Grote fixed cynthia (Drury) as the type of Sarnia, confirmed promethea (Drury) as the type of Packard’s genus Callosamia^ and named cecropia (L.) as the type of his new genus Platysamia.^^ The species cecropia (L.) and its relatives were referred to a variety of genera in the succeeding years including Attacus, Bombyx, Hyalophora, Sarnia and Platysamia. On the whole, Platysamia predominated until about 1900. This name was then replaced by Sarnia. The most recent check list, that of Barnes and Benjamin, reestablishes the name Platysamia. The forms included in the genus are: Platysamia 1. cecropia Linn. 2. Columbia Smith. nokomis Brodie. winonah Brodie. 3. gloveri Strecker. 4. euryalis Bo\sd\iw3\ = ceanothi Behr = calif or 7tica Grote = rubra N. & D. kasloensis Cockerell. cedrosensis Cockerell. ^ ^^See Appendix I. 1937 Sweadner: Hybrids and Phylogeny of Platysamia 173 B. Descriptions and Distributions of the Forms In the following descriptions no differentiation will be made be- tween species, forms, etc., because the writer is not in accordance with the accepted status of the organisms. Their relationships and tax- onomy will be discussed later. Platysamia cecropia (Linnaeus) Original description in Systema naturae. Tome I, Carl Linnaei, 1758. Cecropia 3. P. Bombyx elinguis, alis patulis subfalcatis griseis: fascia fulva, superiorihus ocello subfenestrato ferrugineo. M.L.U. Catesb. car. 2. p. 86. t. 86. Habitat in America septentrionali. Supplementary Description: Head, thorax and abdomen brick red, tegula white, abdomen with white and black intersegmental bands, spiracles red, ringed with white. Ground-color of wings black suffused with white or yellow scales giving it a grizzled appearance. Fore wings with basal area red; basal line white, edged externally with black. Medial area usually suffused with red, densely along the basal line and fading out distally. The extradiscal line is white, edged outwardly with red. It may be straight, angled, curved or scalloped, but always tapers towards the costal margin of the wing, sometimes not reaching the costa. The extradiscal line is bordered distally by a band of the ground color which reaches to the ocellus and bears irregular black blotches between the veins at its outer edge. There is a fine brownish gray subterminal line much indented between the veins. This line is separated from the band of ground color by a yellowish gray area and from the termi- nal line by a pale cream area. The terminal line is narrow, uniform in width and pale brown in color. The ocellus, located in cell R5 (or II2) is oval, black with a blue crescent whose horns point towards the outer margin. A narrow red bar extends distally from the an- terior edge of the ocellus parallel to the veins. A zigzag white line extends from the ocellus to the costa and is bordered proximally by a patch of lavender scales and distally by a red spot near the costa. The discal spot which is shaped like an inverted comma with the narrow end directed outwards parallel to the costa, varies greatly in size. It is brick red, bordered with a narrow black line and usually has a white center. 174 Annals of the Carnegie Museum VOL. XXV The hind wing is marked much like the front wing with the fol- lowing exceptions. The area between the costa and subcosta is yellowish white. The red basal area is obsolete, the white basal band extending to the wing base. The extradiscal line is wider, particularly the red component, and does not taper. There is an additional pale scalloped line which separates off small spots from the outer edge of the band of ground color that is distal to the extradiscal line. There is no ocellus. The discal spot is comma shaped with the point ex- tending outwardly between veins m2 and m3 but never reaching the extradiscal line. It is brick red with a fine black border and a white center. The undersides of both wings have the basal white band and the red suffusion missing. Aside from this and the addition of a general suffusion of white or yellow scales, the markings of the upper side are reproduced below. Variation: The ground color may vary from deep black to pale speckled gray. The amount of red suffusion in the medial area may vary from none, as in some eastern specimens, to almost entire red, as in specimens from North Dakota. The white part of the extradiscal line may be lost on the front wing and reduced to the width of a hair on the hind wing. The red band may be reduced to a few scales or, rarely, entirely lost. The discal spots vary greatly in size but always retain their characteristic comma shape. Larva: The larvae are black marked with yellow in the first instar, yellow marked with black and with black tubercles in the’ second, yellow or green in the third, and pea-green in the remaining instars. In the third, fourth and fifth instars the side tubercles are blue or blue tipped with white. In the third and fourth stages the dorsal tubercles on segments two and three are bright cherry red while the remainder are straw yellow. In the last instar (fifth) all of the dorsal tubercles may be pale yellow, but usually those on segments two and three are bright red while the others are pale yellow. Very rarely, the dorsal tubercles may be a burnt orange on segments two and three, smoky yellow on segments four and eleven and pale yellow on the others. The shade of each dorsal tubercle varies from time to time in the same instar but the fundamental color remains the same. The amount of heavy black chitin on the tubercle also varies. All of the species and hybrids of this genus that have been reared by the writer have shown this variability in these two respects. The full 1937 Sweadner: Hybrids and Phylogeny of Platysamia 175 grown larva of Platysamia cecropia attains a length of from four to six inches and may be as thick as a man’s thumb. Cocoon: The inner and outer cocoons are loosely connected. The inner cocoon is a long ellipsoid with the head end closed as with a draw string, while the outer may be one of two forms: either closely applied to but easily separated from the inner, or a loose envelope many times the size of the inner cocoon. The outer cocoon is usually pointed at both ends and attached throughout its length. Both cocoons are a light tawny color. Food Plants: The larvae of P. cecropia may feed on almost any decidu- ous tree or shrub and even occasionally on the broad leafed evergreens, but the preferred hosts are the elderberry {Sambuciis), maple (Acer), willow (Salix) and the wild cherries { Primus). Distribution: Platysamia cecropia is found throughout the eastern states from Maine to Florida and westward into the foothills of the Rocky Mountains. In Canada, it occupies the Maritime Provinces and extends an undetermined distance north into Quebec. Its north- ern limit in Ontario appears to be a short distance south of Cochrane in the east and the Lake of the Woods district in the west. In Mani- toba the species is not found a dozen miles east of Winnepeg, but the vanguard has pushed northward along the Red River to Lake Winne- peg, perhaps farther, and west of Winnepeg extends only to the Assiniboine River. While its western limit in general is the Rocky Mountains, if the records of capture are accurate, P. cecropia has replaced P. gloveri in eastern Montana within the past fifty years, has become plentiful in southern Alberta within the past few years and has invaded the Black Hills within the past decade. Cockerell reported that P. cecropia had invaded the territory of P. gloveri at Boulder, Colorado in 1929. The species is found in eastern and southern Texas and there is a specimen in the United States National Museum from Jalapa, Mexico. There appears to be a wave of P. cecropia in numbers sufficient to completely defoliate trees pushing across the northwestern part of its range. Whether this represents a new invasion made possible by the tree planting activities of the settlers, or to a temporary lack of parasite and disease control, there is no way of determining. Diagnostic characters: The grizzled ground color and tapering extradiscal band on the fore wing and to a certain extent, the red part of the extradiscal band will separate this species from all the others 176 Annals of the Carnegie Museum VOL. XXV in the genus. The larva may be distinguished in the third and fourth and usually the fifth instars by the fact that the dorsal tubercles on segments two and three are red while the remainder are yellow. The cocoon may be recognized by its tawny color, its long line of attach- ment and the fact that it is pointed at both ends. Platysamia gloveri (Strecker) Original Description: P. gloveri was described by Herman Strecker in his “Lepidoptera Rophaloceres and Heteroceres” in 1872. “Platysamia gloveri: Male. Expanse, four and one-half inches. “Antennae pectinated. Head and thorax dark brownish red; collar white; abdomen alternately banded with dark brown and white. “Upper surface, primaries, inner half dark reddish brown, with a dirty white band near the base and a small white oval spot bordered on the outer and inner sides with black; the dark red colored space is bounded outwardly by a white transverse band extending from the costa to the inner margin, widening at the latter termination; the space immediately beyond the white band occupying fully two- thirds of the exterior half of the surface, is dark gray,, composed of black and white scales, the exterior portion being the darkest; the remaining space between this and the exterior margin is dirty gray, transversed by a serpentine black line; a short space in from the apex on the costa is a small black spot, from which a zigzag white line extends a short distance to another larger black spot of an oval shape, which has within it a pale blue crescent; the space from this latter black spot to the costa, and from the zigzag line almost to the white transverse band is violet. “Secondaries have a white basal patch, also a white transverse band as in Cecropia; the color between this band and the base is dark brownish red; in the center is a moderately large lunate discal spot, white edged with black; of the remaining space, between the transverse band and the exterior margin, the inner two-thirds is com- posed of black and white scales, as in the primaries, and the outer third is light, dirty gray, which latter is divided by a black line, on the inside of which line, running parallel with it, is a row of irregularly shaped black spots. “Under surface same as above, with the exception of the ground color of the secondaries between the transverse bands and the base which is composed of black and white scales instead of being dark red as on the upper surface. Female. Expanse, five and one-quarter inches. “Antennae not as broadly pectinated as in the male. Head and thorax brownish or brick red, much the same as in ordinary forms of Cecropia; abdomen alternately banded with the same color and white. 1937 Sweadner: Hybrids and Phylogeny of Platysamia 177 “On the primaries the transverse white band is somewhat irregular and rounded outwardly; the basal band also rounded; the discal spot larger than in male, and slightly lunate; the ground color between the transverse band and the base is brownish red, of a slightly dif- ferent tint than the thorax; the portion beyond the transverse band, light dirty gray as far as the serpentine line, the inner third of it powdered with a band of reddish scales, broader near the exterior margin, and narrowing towards the costa; margin outside of the serpentine line dirty white, shaded slightly exteriorly; the black apical spots and violet patch same as in male. “Secondaries. Color same as in primaries; ornamentation same as in male, except the discal spot which is double the size. “Undersurface marked same as the upper; coloration same as in underside of male. “Habitat: Arizona. i^Dr. Strecker stated further that he had received “examples of it, which were said to come from lower California, but as L. weidermeyerii, Parn. smintheus, and other northern montane species were sent in company with it, I expressed my doubts regarding that locality, which have since been confirmed by my receiving a female from Arizona.” Present knowledge of the distribution of the Platysamia indicates that the specimen figured could not possibly have come from Lower California. With the assistance of Mr. Benjamin, the writer had the pleasure of tracing the history of Strecker’s specimens at the United States National Museum. The specimens in question (six) were received by Dr. Glover from a Dr. Edw. Smart along with several Nemophila, a Syneda and the specimens listed by Strecker. This happened before the establishment of the National Museum and so the letter accompanying the donation has not been found. Dr. Glover recorded the speci- mens as having come from “South Calif.” and made copper plate etchings of all of the species. The plates were subsequently destroyed and never published, but manuscript copies are deposited in the National Museum library. The specimens accompanying the gloveri, particularly the Basolarchia weider- meyerii, as delineated in the plates could only have come from the Rocky Moun- tain region or from a colony of Rocky Mountain fauna in California. Such a colony is not known to exist. On searching through the files of the National Museum, a donation of a portion of a fossil swan by a Dr. Edw. Smart of Hastings, Nebraska was found recorded. In his letter. Dr. Smart stated that he had dug up the specimen from an Indian mound in 1875. This would place him east of the Rockies about the time that the gloveri were taken. In the handwriting of the letter, there appears to be little difference between the o’s and the a’s. It is quite possible that the donations of Lepidoptera and of the fossil swan came from the same man, and that Dr. Glover mistook the longhand “South Col.” to be “South Cal.”. This would make the type locality of the male figured and described by Strecker to be South Colorado and this fits perfectly with sub- sequent material from that district. 178 Annals of the Carnegie Museum VOL. XXV Variation: In the central part of its range, in Colorado and Utah, the ground color of P. gloveri is wine red. The discal spots are uni- formly reniform, without any appreciable point and vary greatly in size. Also there are no red scales bordering the extradiscal line distally although there is an occasional purplish tinge to the gray subterminal area. Neumoegon described a form, P. gloveri reducta, from Gibson Gulch, Colorado at an altitude of ii,ooo feet that is characterized by having only about half of the expanse of the normal form and by having the gray subterminal band greatly widened at the expense of the central red ground color, the gray band occupying the entire terminal half of the wing. In Montana, Wyoming, South Dakota (Black Hills) and in Alberta the moths approach the form reducta in size, but have the normal gloveri markings, although one specimen from Alberta has the markings as in reducta. The size is apparently a function of the severity of the climate. At the southern- most known limit of the range, in Pima County, Arizona, the char- acters by which the female type specimen differs from the male type specimen are accentuated. Thus, the discal spots have a long point extending distally from the anterior end and sometimes reaching al- most to the extradiscal line. The extradiscal line is edged outwardly with pink scales. The gray subterminal band is more or less suffused with rusty brown. Larva: The larvae resemble those of P. cecropia with the following exceptions. The green color usually has a bluish tinge. In the third and fourth instars the dorsal tubercles are either all red or all yellow, but all the same shade. In the third instar, there is a con- siderable development of the black chitin which partially obscures the red in some specimens or may even completely cover it. In the last instar the dorsal tubercles are all straw yellow, but in a few ex- ceptions, those on segments two, three and four are slightly deeper in shade. The only Montana specimen that I have seen had the dorsal tubercles of segments two and three a dull burnt orange, those on segment four a dirty yellow while the remainder were pale yellow, but this specimen was taken when full grown in Glacier Park on the edge of the zone of intergradation with P. euryalis and is probably not pure P. gloveri. Cocoon: The cocoons are small in proportion to the moth that emerges from them, and have the diameter larger in comparison with the length than in P. cecropia. They are pointed only at the open end 1937 Sweadner: Hybrids and Phylogeny of Platysamia 179 and are attached for less than half their length. The inner and outer shells are so closely woven together that they can scarcely be separated. The silk is white when woven and the outside of the cocoon is covered with ridges. When the spinning is finished, the silk is impregnated with a fluid that forms a blackish gum that does not reach the ridges which remain silvery against a dark background. Food Plants: In Utah P. gloveri feeds on the sand-bar willow (Salix) and on the wild currant (Ribes). In Montana it feeds on the wild currant and on alder {Alniis). Perhaps there are other hosts. Distribution: Platysamia gloveri is found throughout the Rocky Mountain region in the United States and its range probably extends southward into Mexico. It extends westward to the deserts of the Great Basin and in Montana to a line a few miles west of the Conti- nental Divide. A specimen was taken in the Black Hills of South Dakota in 1933 and the species was once found as far east as Miles City in Montana but is no longer to be found east of the mountains. In Alberta it tends to intergrade with the form P. nokomis from which it can scarcely be distinguished. Platysamia Columbia (Smith) Original Description: Proceedings of the Boston Society of Natural History, Vol. 9, December 2, 1863, by S. I. Smith, read by Mr. A. E. Verrill. Printed in 1865. “Description of a species of Sarnia supposed to be new, from Nor- way, Maine. Sarnia Columbia Male — Antennae black and broadly pectinated. Palpi dark maroon brown. Thorax with a white band before; upper side dark maroon, with a short grey band behind; beneath black, the legs also black, slightly tinged with brownish towards the extremities. Abdomen annulated with alternate black and dirty white. “Above. Primaries with a greyish white band near the base, ex- tending from the inner nearly to the costal border, and enclosing a reddish brown patch at the base. The middle area of the wing is dark brown, tinged with reddish towards the center, and contains a triangular white discal spot; bordered on the side towards the base with black, and on the other sides with greyish brown. There is a narrow white transverse band, wider towards the inner border, be- tween the middle and the outer area. A sinuous black line on a grey colored ground, crosses the posterior border. Near the apex there 180 Annals of the Carnegie Museum vol. xxv is a round black spot, containing a bluish white crescent, with its horns toward the outer border; between this and another small ob- long black spot at the apex there is a zigzag white line in the form of a W, with the upper side toward the outer border. A space along the costal border, extending from this zigzag line almost to the mid- dle area is bluish white, growing darker and more indistinct as it approaches the transverse band. A short band between the middle area and the greyish outer border, extending from the inner border a third of the way across the wing is dark greyish brown becoming lighter as it leaves the inner border. “Secondaries with a small dirty white spot on the shoulder and the anterior border just edged with the same. A white transverse band similar to the one on the primaries. The space between this band and the base of the wing is dark brown, with the discal spot large and white. The outer border is margined with clay color, bound- ed on the inside with an arcuate black line. Just inside of this line there is a band of black spots on a greyish ground; the space between this band of spots and the transverse band is occupied by a wide greyish brown band. “Beneath, the markings of the upper side are repeated; but all the reddish tints are wanting, so as to leave the ground color of the wings black, intermingled with whitish scales. The discal spots are bordered with black. Female. The antennae are less broadly pectinated than in the male and all the colors less intense. Discal spots on the primaries almost obsolete; being only short lines bordered with black, and parallel to the transverse band. Discal spots of the secondaries much smaller than in the male and more rounded. “Expanse of wings: cf 4"; 9 4.9" i & 19.” Variation: The discal spots of the hind wings may be oval, or reniform with the upper end drawn out into a slight point. Out of 57 specimens in the writer’s collection, eight in the Redpath Museum at Montreal, four in the Museum of Comparative Zoology at Cam- bridge, three in the Carnegie Museum at Pittsburgh, and 23 in the United States National Museum, 95 specimens in all, 42 have red along the outer margin of the extradiscal band varying from a thin line of scales visible only on the underside to a well defined band wider than the red band of some P. cecropia. Larva: The larvae of P. Columbia are characterized by the great development of the heavily chitinized patches on the tubercles with the result that these tubercles are more slender than those of the other forms and the color is usually hidden. The dorsal tubercles of the third and fourth instars, when they show any color, resemble those of P. cecropia except that those of the fourth segment are a reddish 1937 Sweadner: Hybrids and Phylogeny of Platysamia 181 orange instead of yellow. Also in the last instar, this set of tubercles is the same shade of red as those on the second and third segments while the remainder are yellow with a pink tinge. The larvae of P. Columbia are smaller in all stages than those of P. cecropia or P. gloveri. Cocoon: The cocoon resembles that of P. gloveri in structure and color but is slightly darker, smaller and more slender. The puckered opening, however, is about the same size as that of P. gloveri, making it much more conspicuous on the small cocoon. Distribution: According to records from before 1900, P. Columbia extended from western Ontario to Maine and down into southern Michigan. At the present time, it is limited to the swampy region east of Winnepeg, along the northern shore of Lake Superior and east to the region of North Bay, Ontario. A small colony of the species still remains in Michigan and it has been recently reported again from Maine. No specimens are recorded as taken in southern or eastern Ontario or in Quebec for thirty years. The principal cause for the restriction of range in Canada appears to have been the almost com- plete destruction of the larch {Larex americana), the food plant of P. Columbia, by the. larch sawfly although disease may have been an equally important factor. Platysamia nokomis (Brodie). Original Description: The form nokomis was first described by Mr. William Brodie in a paper read before the Natural History Society of Toronto. The paper was not published, and both it and the types of P. nokomis and P. winonah were lost. Later Mr. Brodie redescribed the forms in the Biological Review of Ontario, 1894. The type locality is listed as Carberry, Alanitoba. The description follows: “The following points of difference may be noted between the Columbia nokomis form and the Columbia form, as represented by Ontario specimens, and as compared with Smith’s description of Columbia, parts of which are given in brackets. The standard of colors is Ridgeway’s Nomenclature of Colors. “Antennae, central shaft, bright reddish brown; pectinations darker [black]; palpi, light liver brown [dark maroon brown]; dorsum of thorax, bright reddish liver brown with a posterior pure white band [dark maroon with a short grey band]; underside of thorax red- dish liver brown [black] ; legs, reddish brown, pile darker [black, slightly tinged with brownish]; abdomen with alternate annulations, bright liver brown and pure white [black and dirty white]. 182 Annals of the Carnegie Museum vol. xxv “Primaries above with a rather sharply elbowed pure white line [greyish white] ; the middle area of the wing is bright reddish liver brown [dark brown], and contains a central ovate white spot [triangu- lar]; this bright area is separated from the costa by a moderately wide longitudinal greyish stripe. Secondaries with a large white spot at the shoulder [small dirty white]; the central area, bright reddish liver brown [dark brown], having a central white spot, which varies from kidney form to curved pear form and varying much in size, but always larger that the corresponding spot on the primaries; but no sexual difference could be observed either in the size or in the form of these central spots. “The primaries beneath have the space from the shoulder to the median white crossband of a maroon brown [black], and generally the underside of the wings of Columbia nokomis is brighter colored than that of Columbia.^’’ Supplementary Description: The collar is white and the discal spots (Brodie’s central white spots) are bordered with tan and outwardly ringed more or less completely with black. Variation: On the whole there is a tendency for the reddish por- tions to be brighter than the “reddish liver brown” of Brodie’s de- scription, usually as bright as the claret of P. gloveri or even brighter. In three out of six specimens at the U. S. National Aluseum, in 15 out of 21 specimens in the writer’s collection and in about half of a considerable number of specimens noted in the collection of Manitoba entomologists there is a red band along the distal border of the extradiscal white crossband ranging from a few scattered scales to a solid band equal in width to the white band. Distinguishing Characters: Platysamia nokomis can be separated from its eastern neighbor, Columbia, much more easily than from its western neighbor, gloveri. From the former it may be distinguished by its lack of melanic suffusion, its wider extradiscal band and its wider ovate discal spots. While an experienced lepidopterist can separate most of the nokomis from gloveri, there is no single constant character or combination of characters that can be listed as diagnostic of the “species.” This opinion is based on examination of more than a hun- dred of each “species” collected over almost their entire ranges. Larva: The larvae of Platysamia nokomis differ structurally from those of P. Columbia in that they lack the heavy black chitinization of the tubercles in the third and fourth stages, resembling those of P. gloveri or P. cecropia in this respect. The first instar is black show- ing yellow spots when full fed. The second is a brilliant orange- 1937 Sweadner: Hybrids and Phylogeny of Platysamia 183 yellow fading to a pale canary; the tubercles are black. The third instar is a pale pea green with canary yellow at the base of all the dorsal tubercles except the first. The fourth instar is bluish green, the dorsal tubercles from segments two to eleven all bright yellow, the side tubercles blue. The fifth stage is a greenish slate color as in P. gloveri, the dorsal tubercles all being ochre yellow, those on segments two, three and four having a patch of heavy black chitin on the side towards the head. Cocoon: The cocoon resembles that of P. gloveri except that it is smaller and more silvery. It is slightly larger than that of P. Columbia and is less slender. Food Plants: The principal food plant of P. nokomis is the wolf willow (Elceagnus). It also feeds to a certain extent on the willow {Salix). Distribution: P. nokomis is found in Manitoba west of Winnepeg and north of the Assiniboine River. It is also found in Saskatchewan and Alberta, although many, if not all, of the specimens from the latter province may be P. gloveri. Platysamia winonah (Brodie) Original description: Platysamia winonah was described by Mr. William Brodie along with P. nokomis. In his redescription of the latter he makes the following comment: “Early in 1883 I received a package of cocoons of P. Columbia and T. polyphemus collected by W. G. A. Brodie near Pelley, N.W.T. Only one imago emerged from this lot, from a P. Columbia cocoon, and it dilTered so very much from the Manitoba form that I considered it a well marked variety, being much less in size and of much brighter colors, and the boundaries of the colors much more distinct . . . . for the Pelley form [I proposed the name] P. Columbia winonah. . . “As I have not seen but one specimen of the Columbia winonah type, little need be said about it. My specimen may have been re- presentative of an extreme northern group, or it may have been only a strongly marked specimen of Columbia nokomis.^' This is all that is known of P. winonah, and until more specimens are collected at or near the type locality, the writer is unable to com- ment further regarding the exact status of the organism represented by the name. 184 Annals of the Carnegie Museum VOL. XXV Platysamia euryalis (Boisduval) This species of Platysamia from California was first described and named by Boisduval in 1855. A month later Behr described it in California, but applied no name and a descriptive adjective used has crept into use as a name. A little later in the same year the name ceanothi was indirectly attributed to Behr when specimens donated by him were recorded under that name. Grote, refusing to recognize Boisduval’s description, again described it in 1865, applying the name calif ornica. Origmal Description: Boisduval in “Bulletin Entomologique,” i" trimestre, 1855, Societe Entomologique de Erance. “Tels sont entre autres la Saturnia euryalis, voisins de la cecrop(i)a, de I’Amerique du nord, mais beaucoup plus petite et fort distincte au premier coup d’oeil, par la longue lunules des ailes inferieures qui traverse la bande mediane; . . . .” Grote’s description is somewhat more complete and will serve much better than a supplementary description by the writer. ‘‘‘‘Platysamia calif ornica’^ “Male and Female — smaller than P. cecropia. Primaries reddish- brown shaded subterminally with a brighter plum-colored tinge, with no gray scales whatever. [Italics mine] A broad basal white arcuate band, running from the interior margin, at extreme base, to just below the costa at basal fourth, narrowly lined outwardly with dark scales. Beyond the disc a moderate, whitish, sub-luniform spot, shaded with buff. A nearly straight sub-terminal whitish band narrower than the basal band, lined outwardly with dark scales. Subterminal space with a brighter, somewhat peach-colored shade, which becomes less distinct outwardly. As in P. cecropia, the apical interspace has a W-shaped pale mark preceded by lilac scales. Be- low, in the post apical interspace, is a black ocellus, margined with a blue annulus, obsolete outwardly. Terminal space dull pale wood- color, much the same shade as in P. cecropia, but narrower. The terminal line is narrow and but slightly waved. “Secondaries resemble primaries in coloration. A few whitish scales at extreme base. A larger, similarly colored sub-luniform spot to that of the primaries, which is produced so as nearly to attain to the outer transverse white band, which latter matches that on the anterior wings. Under surface darker, but resembling upper surface i^Augiistus R. Grote. “Notes on the Bombycene Moths of Cuba.’’ Pro- ceedings of the Entomological Society of Philadelphia, Vol. 5, p. 229, 1865. (The reference is to the footnote.) 1937 Sweadner: Hybrids and Phylogeny of Platysamia 185 in ornamentation; secondaries with a white patch at base on costa.” “Head, capital appendages, thorax and abdomen reddish-brown very nearly concolorous with the wing. 'Collar’ entirely white. Abdomen with long white hair fringing the segments posteriorly, very distinct in female; in the male, bands are more confused. The sexes resemble each other. o' 9 “Expanse 3.85 in. 4.25 in. Length of body i . 10 in. 1.85 in. Habitat: California (San Francisco)” Variation: The length of the discoidal lunules varies greatly; the point often penetrates and extends beyond the extradiscal line. The extradiscal line on specimens from high altitudes is usually bordered on both sides with dark scales while that of lowland specimens is not. Many of the Oregon specimens are darker than those of California. Larva: The writer has never reared pure P. euryalis and the published notes describing the larval stages do not check with data obtained from hybrids. The first stage is black, becoming spotted with bees- wax yellow when full fed. The second instar is straw yellow with black tubercles. In the third instar, the body is green, the dorsal tubercles are concolorous, and yellow, salmon or red, while the side tubercles are turquoise blue. In the fourth and fifth instars the body is pea green and the dorsal tubercles, as far as known, are concolorous and straw yellow. The side tubercles are blue or blue tipped with white. Cocoon: The cocoon of P. euryalis is a tawny brown color as in P. cecropia and lacks the dark gummy impregnation found in P. gloveri. It is flask shaped, the main body being ovoid, almost round, with a slender neck at the open end. It is seldom fastened longitudinally to a support as are the cocoons of the other members of the Platysamia. Distribution: This species is found throughout California, western Nevada, Oregon and Washington to Vancouver Island, British Columbia. In eastern Washington, in Idaho and in British Columbia it intergrades with P. gloveri, the mid-intergrade having been assigned the varietal name, kasloensis, by Cockerell. Platysamia kasloensis (Cockerell) Original Description: The following description appears as a footnote in Part III, page 226, of Packard’s “Monograph of the Bombycene Moths of North America,” the third part of which was edited by Dr. T. D. A. Cockerell after Dr. Packard’s death. “S. rubra kasloensis, collected by Dr. H. G. Dyar at Kaslo. This 186 Annals of the Carnegie Museum vol. xxv is the form described by Mr. Cockle. ... It is a submelanic race, darker above than the typical; much blacker and less red below.” Mr. Cockle’s description follows. (It was never intended as a scientific description and is consequently rambling. Only the de- scriptive portions are quoted.) “There is undoubtedly a local race of the species [rubra = euryalis] to be found in the interior of British Columbia, the general color of which is purplish-brown above on all wings, whilst the underside is always greyish. . . . The markings of both forms [rubra and the British Columbia form] are very variable, the only difference being in the submarginal line which, instead of being deeply dentate as in the California species, has a tendency to lose this dentation, the line being a series of loops bent outwardly with, in some cases, a slight dentation between the veins.” Mr. Cockle continues with the following comment which will be referred to later: “I have frequently been met with remarks from collectors to whom I have sent this form of rubra that it was nearer to gloveri than to rubra, [Italics mine] but the two species are sufficiently distinct in the median lines to prevent all possibility of mistake, but at the same time the northern form of rubra is so different in appearance to warrant a separation and possibly, when the investigation is carried to completion, even a new name.”^^ It may be said in passing that judging from about two hundred specimens of the forms involved, either specimens in the writer’s collection or photographs of specimens in other collections, the shapes of the submarginal and median lines upon which Mr. Cockle effects his separation are not valid characters but subject to great variation in all of the species. Supplementary Description: The general tone of the ground color may vary from red as bright as that of P. euryalis to a deep reddish brown. The discoidal lunules may vary from comma-shaped spots reaching to the extradiscal line on the hind wing to spots that are almost obsolete. All of the specimens have a mixture of red and grey scales in the band of ground color just outside the extradiscal line. This last character may be considered as diagnostic of the form. Larva and Cocoon: The writer has never seen either larva or cocoon, all of his specimens having been taken on the wing, but the hybrid larvae of the cross P. kasloensis cf x P. gloveri 9 in their last instar have all of the dorsal tubercles a bright coral red whereas the homo- i^Cockle, J. W. Entomological News, Vol. 19, July 1908, p. 340. 1937 Sweadner: Hybrids and Phylogeny of Platysamia 187 logous tubercles of P. gloveri are always yellow. Further, the hybrid larvae from the cross P. kasloensis d' x P. cecropia 9 in their last three instars have the dorsal tubercles on segments two, three and four bright red, and the remaining dorsal tubercles a bright pinkish- orange, about the shade known as tango. This seems to indicate that the dorsal tubercles of P. kasloensis are either red or reddish- orange in all of the last three instars. Distrihutiofi: Platysamia kasloensis is found throughout northern Idaho, southeastern British Columbia and western Montana. It intergrades with P. euryalis on the west and with P. gloveri on the east. Later in this paper it will be shown that P. kasloensis itself is the intergrade between these last two species. Platysamia cedrosensis (Cockerell) The variety cedrosensis was described in the same footnote as was kasloensis. “5. rubra cedrosensis , marked “Cedars I,” (evidently Cedros I) Mexico. Male. Margins of upper side of wings broadly and suffusedly blackened, the submarginal markings almost entirely lost; ocellus of primaries smallish; discal mark on hind wings longer and more slender than in kasloensis; beneath the wings are very black, but the region basad of the bands is suffused with brownish vinaceous.” Cockerell’s types were not marked because they were named after he had left the Museum and there is no specimen in the U. S. National Museum, where the type is supposed to be, that bears a label "Cedars I.” The only specimen, unlabeled, that comes close to fitting the description is one of the kasloensis type, whose wing tips appear to have been blackened with soot as is characteristic of a moth that has been flying around an oil lantern. In fact the microscope revealed that the darkening of the outer third of the wings of this particular specimen was due to soot. Whether the type has been destroyed or this is it, we cannot say. Dr. Cockerell believes that it is not. LIntil other specimens are obtained from the type locality or near it, the form cedrosensis, like winonah, must remain merely a “list” name. Aberrations The usual color aberrations caused by somatic mutation or by the failure of some developmental process occur in this genus. In P. cecropia the red may be replaced by yellow, or the brown and black 188 Annals of the Carnegie Musfum VOL. XXV be replaced by yellow and orange. A fairly common aberration in P. gloveri and P. nokomis is a general suffusion of white hairlike scales over the surface of one or more wings, which gives them a mouldy appearance. The discal spots may become obsolete in all of the species. There also occur ‘breaks’ in which a part of the pattern becomes distorted without a corresponding distortion of the wing membrane. These ‘breaks’ appear more often in the hybrids than in the pure species. The effect of cold applied to the pupa at the time of scale forma- tion results in a thinly scaled, semitransparent condition; in an in- crease of red pigment and its migration towards the base of the wing; and in an increase of black pigment and its migration towards the margin. This produces a reduction or elimination of the white areas and a general haziness of pattern. The mechanism of this shifting of pigment, if indeed it takes place, is unknown, but the phenomena have been observed in other forms, particularly the Nymphalidae. Summary If we still refrain from distinguishing between species and form, there are eight names applied to the various forms of the genus Platysamia; P.cecropia, P. Columbia, P. nokomis, P. gloveri, P. kasloensis and P. euryalis representing groups of organisms found distributed throughout temperate North America in the manner indicated on the accompanying map. The other two names, P. winonah and P. cedrosensis are represented by single specimens, both of which have been lost. Until more specimens are taken, we may disregard these two names. The forms may be distinguished as follows: P. cecropia by its grizzled ground color, by its tapering extradiscal line on the fore wing and by the more or less wide red outside border to the extradiscal line on the hind wing; P. Columbia by its submelanic ground color, by its white extradiscal line reaching to the costa on the fore wing and by the absence or near absence of red bordering this line on the hind wing. While an experienced lepidopterist can distinguish be- tween P. nokomis and P. gloveri, there is no character that can be cited as separating them, although the former is much less variable than the latter. Both forms have a red ground color inside the extradiscal line and a pure grey ground color outside it. Both have 1937 Sweadner: Hybrids and Phylogeny of Platysamia 189 Fig. I. Reproduced from “University of Pittsburgh Bulletin” (Graduate School Abstracts, Vol. X), Vol. 31, 1934, p. 347. approximately reniform discal spots. P. euryalis is distinguished by its brick red ground color, by its long comma shaped discal spots and by the absence of grey from the ground color outside of the extradiscal line. P. kasloensis may be distinguished from both P. euryalis and P. gloveri by the mixture of red and grey scales in the ground color outside of the extradiscal line. 190 Annals of the Carnegie Museum VOL. XXV The larvae are most easily separated in the fourth instar, the dis- tinguishing character being the color of the dorsal tubercles, although structurally, P. Columbia has a greater hard chitinization of these tubercles than the others and P. cecropia has more spines. On P. cecropia the dorsal tubercles of segments two and three are red, the rest being straw yellow; on P. Columbia, they are red on segments two, three and four while the remainder are yellow. On P. nokomis all are bright yellow; on P. gloveri all are yellow or all red; and on P. euryalis, all are yellow, or possibly salmon colored. The cocoons are of three types. Those of P. cecropia are more or less smooth, tawny and pointed at both ends, the inner and outer shells being easily separated. Those of P. gloveri, P. nokomis and P. Columbia are dark, covered with silvery ridges, pointed only at the open end and impregnated with a dark gummy substance. The cocoon of P. euryalis resembles that of P. cecropia in color and tex- ture but is ellipsoidal, almost spherical and has a slender neck at the open end. III. METHODS Methods of Collecting and Cross Breeding In the' early part of the work, cocoons were collected in the Fall or purchased from collectors in other localities. They were kept over winter in a cold place in order that they would not emerge before the wild moths around Pittsburgh. Males of one species were confined with females of another in cheesecloth cages of about one cubic yard capacity. This method, which was used by Miss Morton, yielded a number of crosses but many refused to mate. A second method which was more successful was the method known as ‘tying out.’ In it the female to be mated is tied to a protecting bush by a piece of soft string around the body between the primaries and secondaries. The only variation in the use of this time honored method was that the female was always of a species not occurring in the district. Females ‘tied out’ in this manner were seldom unmated. It became evident early in the experiments that cocoons of the species other than cecropia could not be obtained in sufficient quantities, if at all. Also, a means for determining the nature of the insects in the zones of contact between the species was imperative. The knowl- edge that the females of any one species readily attract and mate with 1937 Sweadner: Hybrids and Phylogeny of Platysamia 191 the males of any other species in the genus, as was learned the first season, and the use of a truck for transportation of the insects solved these problems. During 1932 and 1933, collecting, breeding, and the analysis of the zones of contact, were combined into one operation. A road was selected through the country to be explored, and virgin females (usually P. cecropia) were tied in the shelter of bushes suf- ficiently far back from the road so as to not be disturbed by traffic. These ‘plants’ were spaced at intervals of from five to ten miles for a distance of about a hundred miles, and camp was made in a par- ticularly promising locality. The remaining females were placed in a wire cage on the engine hood. If there were Platysamia males in the neighborhood, the operator was awakened about an hour before dawn by their flapping against the top and sides of the truck as they tried to locate the females. He then caught them in his hands as they fluttered against the cage, dropped those he wished to mate into the cage and those he wished for specimens into a cyanide bottle. The rejected ones were placed in a bucket covered over with netting, to be released at dawn. The number of males dropped into the cage must always be one less than the number of females there because the attracting odor is not emitted after mating. Although the males normally remain in coitus for from fifteen to twenty hours, it has been found that two hours is sufficient for complete ensemination. At the end of about two hours the cage was placed inside the truck and the route of the previous day retraced. The jarring of the wheels on the road soon dislodged the males and as soon as the union was broken, the male was killed and given a number. The female was placed in a small cardboard box or paper sack with the same number. Notes must be taken concerning any peculiarity of color or pattern before the female is placed in the box because she remains in there five days and batters her wings to pieces. At the end of the egg lay- ing period the female was killed and preserved and the eggs which she had glued to the inside of the box were picked off and kept in vials until they hatched. Along the ‘trapline’ all the females were found mated if one was lucky. We thus got a sample male from each in- terval of the district covered, and a fairly complete record from the district of the camp. In this manner also, up to twenty-five cross matings have been secured in a single night. Factors which prevented mating or the securing of the males were (i) a cold night, (2) high winds which forced early separation, (3) birds, mice or ants may have 192 Annals of the Carnegie Museum VOL. XXV killed the female, and (4) the tied female may have been discovered by some other human. Methods of Rearing The eggs hatch in from ten to twenty days, turning dark a day or two before the larvae crawl from the shell. Experience has demonstrated that wild cherry {Primus serotina) is best suited as a food plant for the hybrids regardless of upon what plant their parents feed. If this is not available, elderberry {Sambucus) , willow {Salix) or alder (Alnus) may be used in the order named. As soon as the larvae emerged from the shell, they were placed in a large vial with several leaves, and the vial was plugged with cloth and put in a cool place. Fresh leaves were substituted at six hour intervals, care being exercised that the tiny larvae were not pulled from the old leaves but cut out on a small sec- tion of the leaf to which they clung. After twenty-four to thirty-six hours they were ready to be placed in the breeding cages. The hybrids were too delicate to be reared on cut leaves in boxes and cans as the parent species may be. They had to be fed on the living food plant under ideal conditions. A cage was devised that gave the larvae protection against all of the parasites and predators except boys and the ants. The latter were excluded by banding the trunk of the food plant with tanglefoot. The cage was constructed as follows: thirty inch, sixteen mesh window screen was cut into thirty-six inch lengths. Pieces of strong cloth twelve to fifteen inches wide were sewn to the long sides. The short sides were then tacked to a stick of wood so as to form a cylinder, the ends of which were cloth. The edges of the cloth were then sewn together to complete the cylinder which was open at both ends. In use, a branch was selected and violently shaken and carefully inspected lest a spider or reduvid bug remain and be confined with the larvae. The cage was then slipped over the branch and the cloth tied firmly around the stem. The larvae were then introduced through the free end and the end tied shut. This arrangement permited the larvae to feed on the living host under natural conditions with alP of the attendant advantages but at the same time afforded complete protection against predators and insect parasites. The cage was examined from day to day and when nearly all of the leaves had been eaten, it was removed to another branch. The caterpillars, which by that time were in the second instar and had reached a length of about three-quarters of an inch, were trans- 1937 Sweadner: Hybrids and Phylogeny of Platysamia 193 ferred by cutting out the pieces of the leaf to which they clung or by rolling them off the twigs. Great care was exercised in order that the delicate prolegs might not be torn. If the caterpillar was moulting, the entire twig was cut off and carefully placed on the fresh foliage. As the caterpillars grew, fewer and fewer were kept in a single cage, until during the last instar, when they attained a length of from three to live inches, no more than ten were kept in each cage. Methods of Pattern and Color Analysis Intergrades were found in the zones of contact, particularly in the contact between P. euryalis and P. gloveri. When the specimens were pinned to a large board according to their geographic distribution, the way in which they intergraded immediately struck the eye. Such a demonstration could not be put on paper, so a statistical method was devised for illustrating this blending. The specimens were classified according to the degree to which they resembled one or the other of the supposed parent species. For instance, the two most important differences between P. euryalis and P. gloveri are the shapes of the discal spots on the hind wing and the color of the area just out- side of the extradiscal line. The discal spots of P. gloveri are reniform (except in its most southern range) while those of P. euryalis are quite elongated. The area in question is red with no gray in P. euryalis and the reverse in P. gloveri. In the analysis of the discal spots, since the size varies greatly in both species, no single measure- ment such as length or area was found to be of any value for com- parison and a ratio of length to width, which would have eliminated the factor of variable size was found to depend on from what points the measurements were made. A satisfactory measurement, the ratio of average width to longest length, was obtained in the following manner. The spots were carefully traced using a camera lucida and the areas thus obtained were measured with a planimeter. This area divided by the square of its longest length gave the desired index of the shape of the spots. The ratios thus obtained were plotted against the geographic distribution as is shown in Chart II, page 208. The other character (color) did not yield to so objective an analysis. The gradations between the gray and red were divided into eighteen approximately equal classes and the specimens plotted by classes against geographic distribution as recorded in Chart I, page 207. Because of the extreme variability of P. cecropia, and the small 194 Annals of the Carnegie Museum VOL. XXV amount of intergradation detectable, no such comparison could be made in the contacts between P. cecropia on one hand and P. Columbia or P, nokomis on the other in the eastern zone of contact. Method of Genitalic Analysis Mr. Foster H. Benjamin of the United States Bureau of Entomology and Plant Quarantine pointed out that there occur instances where two species intergrade in color and pattern but are quite distinct in their genitalia. He suggested that before any proof could be estab- lished that intergradation was due to hybridization, this possibility must be eliminated. He very generously devoted considerable time to instructing the writer in the art of preparing slides of the genitalia of Lepidoptera, the Saturniidae in particular. The male genitalia were prepared as follows: The insect was re- laxed and a scalpel inserted into the underside of the abdomen just ahead of the genitalia. The latter was then pried off leaving the rest of the body undisturbed. This ‘tip’ was then either soaked over night or boiled for ten to fifteen minutes in a ten per cent solution of potas- sium hydroxide. This treatment destroyed the soft tissues and to a certain extent bleached the chitin. The genitalia were then placed in water and all vestiges of soft tissues teased out under a binocular microscope. When the dissection was finished, all scales and scale- like hairs, all fat and other tissue had been removed leaving only the chitin. The dissection was then dehydrated by dropping into 95 per cent alcohol where it remained half an hour. The valves were then spread open as far as possible without distorting the parts and oil of cloves dropped on. The oil of cloves serves the double purpose of finishing the dehydration and hardening the tissue. After soaking in this reagent for half an hour, the tissue was placed in xylol for a similar period and then mounted in a deep balsam cell on a slide and covered with a cover slip. Because of the comparatively large size of the Platysamia genitalia the cell had to be made from one and a half to two millimeters deep. So deep a cell would require years to dry, so as soon as the slide became set, it was cleaned and the cell sealed with celloidin. The female genitalia were prepared in much the same way but have not been used in this report because no clear cut differences were found between the most divergent species. 1937 Sweadner: Hybrids and Phylogeny of Platysamia 195 Summary Since the color patterns, morphology and distribution of the various forms of the Platysamia suggested hybridization, the following methods were devised or borrowed from other workers for the purpose of analyzing the genus. Cross matings were secured either by con- fining the different forms together in wire screen or cheesecloth cages or by tying out a female to a bush in the territory of another form. The hybrids were reared in specially designed wire and cloth cages on the living food plant. Samples of the indigenous males of the zones of intergradation or zones of contact between two forms were obtained by tying out congeneric virgin females at intervals in the territory. Intergrades were analyzed by determining to what degree the characters that differentiate the supposed parent forms were present and plotting this against the geographic distribution. Identifi- cations of the species were checked by examination of the male genitalia. IV. OBSERVATIONS A. Breeding Experiments TIME OF FLIGHT AND NORMAL MATINGS The moths of the family Saturniidae are elinguate in the adult and consequently very short lived. A mated female will live about five days, and seven or eight if unmated. The male lives a little longer. The span of life is lengthened by cold and shortened by heat as would be expected since temperature controls the rate of metabolism and consequently the rate of depletion of stored food. A life of four- teen days is recorded for Actias selene.^^ Because of the short period of flight, the eggs and sperm are fully formed while the moth is in the pupa and are ready for fertilization immediately after eclosure. Normally the female mates the first night after emergence but can be mated on any night up to four or five days of age. Mating of the species of Platysamia normally takes place just before or just after dawn. Rau^^ reports that the time of flight for P. cecropia in the latitude of St. Louis, Missouri, is sharply i^Klaue, Dr. Wolfgang, “Die Winterzucht von Actias selene (Lep.),” Ento- mologische Zeitschrift. Stuttgart. Vol. 44, pp. 40-42. i^Rau, Phil, and Rau, Nellie L., “The Sex Attraction and Rhythmic Periodicity in Giant Saturniid Moths.” Transactions of the Academy of Science of St. Louis, Vol. XXVI, No. 2, 1929. 196 Annals of the Carnegie Museum VOL. XXV confined between the hours of 3:30 and 4:30 A.M., except in a few cases in which moonlight simulates the light conditions of early dawn. Platysamia ettryalis, P. kasloensis and P. gloveri in Idaho and western Montana also fly just before sunrise, the flight starting at or just be- fore the first lightening of dawn and continuing until just before the sun itself appears. Platysamia Columbia in Ontario and Manitoba and P. nokomis in Manitoba fly under the same conditions. Platysamia cecropia, on the whole, flies at dawn, but I have secured matings in the early evening and have taken males at light (rarely), also in the evening. The flight in these exceptional cases may have been in- fluenced by moonlight; of this I have no record. On the fifth of June, 1933, in a tamarack swamp in Claire County, Michigan, a few P. cecropia females attracted a number of P. cecropia males about two hours after dark. A dense fog had settled over the swamp. The attracting medium is a chemical emanation, as proven beyond doubt by Rau and others. It is unlikely that it is the characteristic odor perceptible to man since the sexual emanation is stopped as soon as coition begins, while the odor remains. The male flies against the wind and usually goes past the female on the first try. Having thus lost the scent he drops back until he is down wind from the female and tries again. There is apparently no selection on the part of the female. She takes the first male that reaches her. Coition usually lasts until the following evening when the female becomes active and dislodges the male which then flies away. This long union is due to the daylight period of inactivity, an hour or so giving ample time for complete insemination. The mated female flies throughout the night laying her eggs in groups of from three to six, usually on the top surface of a leaf or on twigs. Record of Experimental Cross Matings Cross Matings in Conflnement: In the attempts to secure crosses between various Saturniidae and between various American and Palearctic Sphingidae in 1930, three crosses were obtained between Platysamia cecropia males and P. gloveri females confined together in cheesecloth cages indoors. There was a slight current of air through the cage. I'^Collins, C. W. and Potts, S. F. “Attractants for the Flying Gypsy Moths as an Aid in Locating New Infestations.” Technical Bulletin No. 335, U. S. Department of Agriculture, 1932. 1937 Sweadner: Hybrids and Phylogeny of Platysamia 197 In the Spring of 1931 with a larger and more varied supply of cocoons a number of matings were secured using cheesecloth cages placed out of doors. Two cages were placed together so that the wind would blow from the female of one species past the female of another species to the male of the first species. Before the weather became warm enough to place the moths outside, a few were kept in cages in a room having a single north window. On the seventh of May in this cage a P. cecropia male mated with a P. gloveri female, whose wings had not yet completed expanding, at 5:30 p.m., long before the light had de- creased to the intensity normally required for activity in this genus. In the outside cages the following cross matings were obtained: P. gloveri cf x P. cecropia 9 (2) ; P. cecropia cT x P. gloveri 9 (4) ; P. nokomis (p x P. cecropia 9 ; P. cecropia (p x P. nokomis 9 ; P. euryalis cp x P. gloveri 9 : P- euryalis cP x Actias selene 9 ; P. cecropia (p X Sarnia walkeri advena 9 ; and Sarnia walkeri advena cP x Callosamia promethea 9 . All of the above matings proved fertile except the intergeneric crosses, but because of inexperience in rearing and to other extrinsic causes none were reared to maturity except the product of some of the gloveri-cecropia crosses. In connection with the survey trip through the zone of contact between Platysamia euryalis and P. gloveri in Idaho and Montana during 1932, many cross matings were secured between various species while they were confined out of doors in a large cheesecloth cage five by eight by three feet, or in wire cages measuring eighteen inches on a side. Most of the matings involving hybrids reared the preceding year were procured in this manner. The matings included: P. cecropia cp x P. gloveri 9 (7) ; P. gloveri cp x P. cecropia 9 (8) ; P. cecropia-gloveri hybrid