[■■'■■ ■'■•:;--:- ". Skbhmn if tt tii ;',..... HARVARD UNIVERSITY Library of the Museum of Comparative Zoology BREVIORA %° *° MUSEUM OF COMPARATIVE ZOOLOGY Harvard University Numbers 296-325 1968- 1969 CAMBRIDGE, MASS. U.S.A. 1969 CONTENTS BREVIORA Museum of Comparative Zoology Numbers 296-325 1968 No. 296. The genus Dysderina (Araneae, Oonopidae) in Central America and the West Indies. By Arthur M. Checker- ing. 37 pp. October 11. No. 297. Population structure of the Asthenes flammulata super- species (Aves: Furnariidae). By Francois Vuilleumier. 21 pp. October 1 1. No. 298. Monograph of the Cuban genus Viana (Mollusca: Archaeogastropoda: Helicinidae). By William J. Clench and Morris K. Jacobson. 25 pp. October 11. No. 299. What is Lumbricus eiseni Levinsen, 1884 (Lumbricidae, Oligochaeta)? By G. E. Gates. 9 pp. October 11. No. 300. Contributions to a revision of the Lumbricidae. III. Eisenia hortensis (Michaelsen) (1890). By G. E. Gates. 12 pp. October 1 1. No. 301. New echymid rodents from the Oligocene of Patagonia, and a synopsis of the family. By Bryan Patterson and Rosendo Pascual. 14 pp. October 11. No. 302. Geomyseria glabra, a new genus and species of scincid lizard from Bougainville, Solomon Islands, with com- ments on the relationships of some lygosomine genera. By Allen E. Greer and Fred Parker. 1 7 pp. October 1 1 . No. 303. Review of the genera of the tribe Loberini (Coleoptera: Languriidae). By T. Sen Gupta. 27 pp. December 31. No. 304. Redescriptions of Anachis avara (Say) and Anachis translirata (Ravenel) with notes on some related species (Prosobranchia, Columbellidae). By Amelie H. Scheltema. 19 pp. December 31. No. 305. Lytechinus williamsi, a new sea urchin from Panama. By Richard H. Chesher. 13 pp. December 31. No. 306. .Anew species of Eleutherodactylus (Amphibia, Salientia) from the (inavana region. Edo. Bolivar. Venezuela. B\ Juan A. Rivero. 11 pp. December 31. No. 307. A new species of Hyla (Amphibia, Salientia) from the Venezuelan Guayana. By Juan A. Rivero. 5 pp. December 3 1. No. 30S. The molluscan fauna of an unusual Bermudian pond: a natural experiment in form and composition. By Stephen Ja\ Gould. 13 pp. December 31. No. 309. Variability ol rodent incisor enamel as viewed in thin section, and the micro-structure o\ the enamel in fos- sil and recent rodent groups. By John H. Wahlert. 1 S pp. December 31 . 1 969 No. 310. A shovel-tusked Gomphothere from the Miocene of Kenva. Bv Vincent J. Maglio. 10 pp. March 31. No. 311. The genera Apenesia and Dissomphalus in Argentina and Chile (Hymenoptera, Bethylidae). By Howard E. Evans. 23 pp. March 31. No. 312. Ecological observations on Anolis occultus Williams and Rivero (Sauria, iguanidae). By T. Preston Webster. 5 Pp. March 31. No. 313. Louis Agassiz's numbers for Steamer Blake Stations of 1X77-78, 1878 79. By Myvanwy M. Dick. 11 pp. March 31. No. 314. The cranial anatomy of the Permian amphibian Pantylus. By Alfred Sherwood Romer. 37 pp. March 31. No. 315. Stellicola dentifer n. sp. (Copepoda, Cyclopoida) as- sociated with a starfish in Jamaica. By Arthur G. Humes. 11 pp. March 31. No. 316. Cytotaxonomic studies on some unusual iguanid lizards assigned to the genera Chamaeleolis, Polychrus, Poly- chroides, and Phenacosaurus, with behavioral notes. By G. C. Gorman. R. B. Huey, and E. E. Williams. 17 pp. April 30. No. 317. Relationships of two Cretaceous lizards (Sauria. Teiidae) . By Richard Estes. 8 pp. April 30. No. 318. Leucolepidopa sunda gen. nov., sp. nov. (Decapoda: Albuneidae), a new Indo-Pacific sand crab. By Ian E. Efford. 9 pp. April 30. No. 319. Competitive exclusion among anoles (Sauria: Iguanidae) on small islands in the West Indies. By A. Stanley Rand. 16 pp. April 30. No. 320. Taimanawa, a new genus of brissid echinoids from the Tertiary and Recent Indo- West-Pacific with a review of the related genera Brissopatagus and Gillechinus. By Robert A. Henderson and H. Barraclough Fell. 29 pp. June 10. No. 321. Observations on the agonistic and breeding behavior of Leptodactylus pentadactylus and other amphibian spe- cies in Venezuela. By Juan A. Rivero and Andres Elroy Esteves. 14 pp. June 10. No. 322. The fossil record of amphiumid salamanders. By Richard Estes. 11 pp. June 10. No. 323. On the earthworms of Ascension and Juan Fernandez Islands. By G. E. Gates. 4 pp. June 10. No. 324. Polymorphism and evolution of the Hispaniolan snake genus Uromacer (Colubridae). By Henry S. Horn. 23 pp. September 15. No. 325. The genus Phenacosaurus (Sauria, Iguanidae). By James D. Lazell, Jr. 24 pp. September 15. INDEX OF AUTHORS BREVIORA Museum of Comparative Zoology Numbers 296-325 1968 1969 No. Chesher. Richard H 305 Chicki ring, Arthur M 296 Clinch. William J 298 Die k. Myvanwy 313 Eei-ord. Ian I 318 Esns. Richard 317, 322 I steves, Andres Elroy 32 1 1 \ ans. How ard E 311 Fell, H. Barrac lough 320 Gates, G. E 299, 300, 323 Gorman, G. C 316 Gould, Stephen Jay 308 Greer. Allen E 302 Henderson, Robert A 320 Horn, Henry S 324 No. Huey, R. B 316 Humes, Arthur G 315 Jacobson, Morris K 298 Lazell, James D., Jr 325 Maglio, Vincent J 310 Parker, Fred 302 Pascual, Rosendo 301 Rand, A. Stanley 319 Rivero, Juan A 306, 307, 321 Romer, Alfred Sherwood 314 scheltema, amelie h 304 Sen Gupta, T 303 VUILLEUMIER, FRANCOIS 297 Wahlert, John H 309 Webster, T. Preston 312 Williams, E. E 316 BREVIORA Museum of Comparative Zoology Cambridge, Mass. 11 October, 1968 Number 296 THE GENUS DYSDERINA (ARANEAE, OONOPIDAE) IN CENTRAL AMERICA AND THE WEST INDIES Arthur M. Chickering Abstract. A total of twenty-four species of the genus Dysderina are recognized in this paper. Seventeen species are described as new. Nine of these are from Panama, two are from Costa Rica, two are from Trini- dad, W. I., one is from Jamaica, W. I., two are from St. Vincent, B. W. I., and the remaining one is from Dominica, B. W. I. Dysderina principalis Simon from St. Vincent, B. W. I. is not D. principalis (Keyserling) from Colombia and, therefore, is described as a new species, D. soltina sp. nov. Dysderina antillana Bryant, described from St. Croix, U. S. Virgin Islands in 1942 and reported from Hispaniola in 1948, has been shown to be Ischnothyreus peltifer (Simon) and is treated in another publication. The Oonopidae include a group of very small spiders usually oc- cupying concealed habitats such as leaf litter, debris, especially grass and weed debris, and other similar habitats. They are par- ticularly numerous in tropical and subtropical regions but are now known from many other parts of the world. The distinctive features of the family are treated in such publications as the following: Simon, 1892-1895; Petrunkevitch, 1939; Comstock, 1940; Kas- ton, 1948, and others, and will not be treated in this paper. Since early in my work of collecting and studying spiders in Panama, I have continued to be interested in these minute members of the order Araneae. For the past several years I have made a special effort to collect members of the family Oonopidae in Central America and the West Indies. As a result of this effort I have accumulated a rather large number of species of several genera belonging to this family, and the time has arrived for me to put the results of these years of study and collecting into a permanent record. It had been my intention to publish the results of these studies in a single monograph. Recently, however, the decision was made to publish a series of shorter papers each dealing with a single genus or a group of genera as conditions seem to warrant. The genus Dysderina is one of a group of genera to be treated early in the series. 2 rreviora No. 296 My appreciation and gratitude are again expressed for the con- tinued aid and encouragement in the pursuit of my studies ex- tended by the staff of the Museum of Comparative Zoology. My thanks are also extended to Dr. W. J. Gertsch, American Museum of Natural History, Dr. G. Owen Evans and Mr. D. J. Clark, De- partment of Zoology, British Museum (Natural History) for the loan of important species of the genus Dysderina. Grants GB-1801 and GB-5013 from the National Science Foundation have made it possible for me to continue my work in the Museum of Com- parative Zoology and to spend a total of nearly twelve months during the last lour years collecting in Panama, Costa Rica, and the West Indies. Except as otherwise stated in later parts of this paper, all types together with my entire collection oi the genus Dysderina will be deposited in the Museum of Comparative Zoology. Genus Dysderina Simon, 1891 The type species is Dysderina principalis (Kevserling) by mono- tvpv. The genus was established on the basis of a male from 1 ilombia identified by Keyserling as Oonops principalis. Simon (1891 ) correctly recognized that this species could not be regarded as belonging to the genus Oonops and. therefore, placed it in a new genus Dysderina. Simon also described two additional species from St. Vincent Island. B. W. I. Dysderina plena O. P. -Cam- bridge was described from Mexico in 1894. Dysderina antillana Br\ant was described from St. Croix, U. S. Virgin Islands in 1942, and also reported from Hispaniola in 1948, but is now known to be Ischnothyreus peltifer (Simon) as will be shown in another paper. In 1951 I reported four species of Dysderina from Panama. Since that time I have collected these small spiders at every opportunity and as a result I now have a rather large collection consisting of numerous species from parts of Central America and the West Indies but none has appeared east of Dominica, W. I. The most important features of this genus observed during my study of the group may be stated as follows: Total length varies from about 1.6 mm to 2.75 mm, with females usually somewhat larger than males. There is a remarkable similarity of general appearance among the species recognized in this paper. Males can be readily separated into species on the basis of the distinctive features of the palpal tarsi. Females, on the other hand, are very difficult to separate into species with any degree of certainty. In my treatment of this sex I have placed great emphasis on the epigynal areas 1968 DYSDERINA IN C. AMERICA AND W. INDIES 3 which occasionally are quite distinctive but more frequently are at best only obscurely so. Nearly all outer parts are strongly chitin- ized. The carapace is moderately high; regularly arched from pos- terior eyes to beginning of posterior declivity; about three-fourths as wide as long nearly opposite second coxae; surface irregularly granulate; usually with no indication of a median groove or pit. Eyes: six in two rows; AME lacking; posterior row gently recurved as viewed from above; quite compactly arranged; little difference in size of eyes but shape often varies (long diameter always used for measurements); clypeus typically heavily bordered. Chelicerae moderately developed; usually vertical, parallel; with fang typically slender and evenly curved; fang groove with a single tooth, at least in certain species. Maxillae with a peculiar terminal notch seen only in males and thus far only in a few species (Fig. 2). Lip: strongly chitinized; considerably wider than long; transversely con- cave in middle. Sternum: strongly chitinized; usually deeply grooved with conspicuous lobules opposite the coxae; continued laterally between coxae and united with a sclerite surrounding the cephalothorax to make a strong enclosure; sternal suture strongly procurved; fourth coxae usually widely separated. Legs: usually 4123 in order of length but with minor variations occasionally; first and second legs with conspicuous ventral spines which vary somewhat among different species; third and fourth legs without true spines. Male palp: all segments except tarsus simple and un- modified; apparently without true spines; tarsus somewhat inflated and with more or less distinctive terminal structures. Abdomen: ovoid; with pedicel strongly chitinized and deeply corrugated; dor- sal scutum varies somewhat in size but typically covers most of dorsum; epigastric scutum continued around pedicel and far dor- sally (Fig. 8); ventral scutum covers much of remainder of venter but varies in extent; a sclerite typically partially surrounds the spinnerets on ventral side; black bristles may mark the position of the reduced colulus; openings of book-lungs, tracheal spiracles, and genital organs more or less distinct. Genital area somewhat dis- tinctive but usually obscure. Where the species under consideration agrees fully with the stated features of the genus no mention will usually be made of the specific characteristics in the description of the species. As a result of my study of the genus Dysderina presented in this paper I am obliged to recognize a total of twenty-four species from the region under study (exclusive of D. principalis (Keyserling) from Colombia). The number is undoubtedly inflated because of the great difficulty in accurately matching up males and females. 4 breviora No. 296 Thirteen of these species are from Panama; two are from Costa Rica; one is from Mexico; two are from Trinidad. W. I.; one is from Jamaica. \V. I.; four are from St. Vincent, B. W. I.; and one is from Dominica. B. W. I. Fifteen of these species, ten of which arc regarded as new. are represented by males; nine are represented only b\ females of which seven are regarded as new; six of the dif- ferent kinds of males are accompanied by what are believed to be properly assigned females. It should be remembered, however, that matching females with the proper males is a very uncertain task. Key to the males of 1)">si>i kina from Central America and the n est Indies la. Species in which the palpal tarsus terminates in a pair of slender distal apophyses, one sickle shaped and one somewhat feather- shaped (Fig. 43) D. obtina, p. 2 1 lb. Species without such palpal, tarsal, distal Structures as given above 2 2a. Species with B group of four or five distal, terminal apophyses on the palpal tarsus (Fig. ;" i D. globina, p. 14 2 ). Species without such palpal, tarsal, distal structures as given above . .3 Species with three definite, distal, palpal, tarsal apophyses (Figs. 41, 77 ) ( meridina, spinigera ) 4 3b. Species without such distal, palpal, tarsal structures as given above . .5 4i Species with distal, palpal, tarsal apophyses as in Figure 41 D. meridina, p. 20 4b. Species with distal, palpal, tarsal apophyses as in Figure 77 D. spinigera, p. 3 1 Sa Species with a single, short, broad, distal, palpal, tarsal apophysis divided into an opaque half and a transparent half ( 1 ig. 4) D. abdita, p. 5 5b. Species without such a single distal apophysis as given above 6 Species with a long, slender, distal, palpal, tarsal spine and a relatively massive, somewhat twisted, closely associated apophysis (Figs. 20-22) D. craneae, p. 1 0 6b. Species without such a pair of distal, palpal, tarsal structures as given above 7 7a. Species in which the distal, palpal, tarsal apophysis turns at a right angle near its base and then divides into a pair of flattened structures ( Figs. 37-38 ) D. intempina, p. 1 (> 7b. Species without such a distal, palpal, tarsal apophysis as given above 8 8a. Species with a single, distal, palpal, tarsal extension divided into a pair of relatively short, incurved apophyses (Figs. 46-47) . . /). plena, p. 23 8b. Species without such a pair of apophyses as given above 9 1968 DYSDERINA IN C. AMERICA AND W. INDIES 5 9a. Species with two distal, palpal, tarsal structures, a slender curved spine and a relatively large, broad, compound apophysis (Fig. 63) D. seclusa, p. 27 9b. Species without such a pair of distal, palpal, tarsal structures as given above 10 10a. Species with palpal, tarsal, distal apophysis terminating in a broad, angular, somewhat concave structure (Fig. 16) . . . .£>. concinna, p. 9 10b. Species without such a distal, palpal, tarsal structure as given above 11 11a. Species with palpal, tarsal, distal apophysis a single robust structure divided distally into a short, slender spine and a sharply pointed robust spine (Fig. 65) D. simla, p. 29 lib. Species without such a palpal, tarsal, distal apophysis as given above 12 12a. Species with a main palpal, tarsal, distal apophysis sharply bent mid- way, enlarged and flattened distally and with a minute spine (Figs. 58-59) D. recondita, p. 25 12b. Species without such palpal, tarsal, distal structures as given above 13 13a. Species with a robust palpal, tarsal, distal apophysis terminating in a series of short, pointed structures obscured by a cluster of hairs (Figs. 69-70) D. soltina, p. 30 13b. Species without such a palpal, tarsal, distal apophysis as given above 14 14a. Species with a pair of palpal, tarsal, distal apophyses curved toward one another; clypeus about three-fourths as high as diameter of ALE (Chickering, 1951, figs. 2, 3; and Fig. 24 this paper) . .D. dura, p. 12 14b. Species with a long, curved, palpal, tarsal, distal apophysis together with a slender spine and a short, blunt process between the two; clypeus about as high as diameter of ALE (Fig. 81 ) . .D. watiua, p. 33 DYSDERINA ABDITA Sp. IIOV. Figures 1-8 Holotype. The male is from El Volcan, Republic of Panama, August, 1950. The name of the species is a Latin adjective re- ferring to its concealed habitat. Description. Total length 2.28 mm. Carapace 1.04 mm long; 0.85 mm wide opposite second coxae where it is widest; well rounded along ventral border from opposite base of palp to pos- terior margin; 0.52 mm tall; quite evenly arched from PME to beginning of posterior declivity which begins opposite interval be- tween second and third coxae; with surface unevenly granulate with median arched area nearly devoid of granulations; without a median, longitudinal thoracic groove or pit; with sparsely situated short recurved hairs or fine bristles. Eyes: six in two rows as usual; posterior row occupies about five-sixths of width of carapace 6 breviora No. 296 at that level; viewed from above, posterior row gently recurved; viewed from in front, posterior row gently procurved (Fig. 1). Ratio of eyes ALE : PME : PLE = 10 : 9 : 9; boundaries of ALE somewhat irregular but nearly circular, others somewhat oval. ALE separated from one another by nearly their radius, from PLE by one-sixth of their diameter, from PME by a little less than one- third of their diameter. PME contiguous only for a short distance and separated from PLE by about one-ninth of their diameter. Height of clypeus (Fig. 1 ) slightly less than the diameter of ALE. Chelicerae: vertical; essentially parallel; basal segment 0.29 mm long; apparently without a basal boss; fang slender and evenly curved; fang groove apparently with a very small tooth, probably promarginal. Maxillae: moderately long; slender; distinctly con- vergent and almost meeting anterior to lip; palp inserted into basal third; with a well developed distal scopula; distal end divided (Fig. 2). Lip: strongly chitinized; extended posteriorly into sternal area and with an obscure groove separating the horizontal part from the more vertical portion; wider at base than long in ratio of about 9:7; anterior border with a row of stiff bristles. Sternum: scutiform in general; as wide as long; widest between second coxae; strongly chitinized and distinctly but less intricately grooved than in several other species (Fig. 3); posterior border recurved and not continued between fourth coxae which are separated by about 1.5 times their width; coxae three and four short and stout; coxae one and two somewhat more elongated; sternal suture procurved. Legs: 4213 in order of length; tibial index of first leg 10, of fourth leg 9; trichobothria present but exact number and placement undeter- mined. Tarsal claws apparently as recorded for D. seclusa (Chickcring. 1951 ). First femur with two pairs of ventral spines in distal half; first tibia with five pairs of ventral spines, first very long but diminishing distally so that last pair is hardly more than a pair of bristles; first metatarsus with three pairs of ventral spines, irregularly spaced. Second femur apparently with only one con- spicuous spine on prolateral surface at beginning of distal third; second tibia with four pairs of ventral spines; second metatarsus apparently with only two pairs of ventral spines, irregularly spaced. Third and fourth legs lacking true spines. Palp: only tarsus in- flated; important features, more or less distinctive, shown in Figures 4-6. Abdomen: with a well developed and strongly corrugated pedicel; 1.24 mm long exclusive of pedicel; 0.9 mm wide near middle; spinnerets as usual in the genus; genital aperture as in Figure 7; dorsal scutum covers nearly the entire dorsum; epigastric scutum continued around anterior end of abdomen; boundaries be- tween epigastric and ventral scuta not clear; ventral scutum covers 1968 DYSDERINA IN C. AMERICA AND W. INDIES Figs. 1-8. Dysderina abdita sp. nov. Fig. 1. Eyes and clypeus; from in front. Fig. 2. Right maxilla; nearly lateral view (from dissected paratype). Fig. 3. Sternum; from below. Figs. 4-5. Left palpal tarsus; nearly ventral and retrolateral views, respectively. Fig. 6. Tip of left palpal tarsus; nearly dorsal view. Fig. 7. Genital aperture. Fig. 8. Abdomen; left side (from paratype.) nearly the entire venter except for a small area at posterior end and laterally where the two scuta nearly meet (Fig. 8). Color in alcohol: in general, a rich reddish brown, essentially as described for D. dura and others (Chickering, 1951 ). Records. Seven paratype males from El Volcan, Panama, Au- gust, 1950. It seems probable that one of the two kinds of females described from the same locality and collected during the same period should be matched with these males but this is not at present possible to do with any certainty. Dysderina Belinda sp. nov. Figures 9-12 Holotype. The female is from Boquete, Panama, August, 1950. The name of the species is an arbitrary combination of letters. 8 BREVIORA No. 296 Description. Total length 2.5 mm. Carapace: 1.08 mm long; 0.9 mm wide opposite posterior border of second coxae where it is widest (Fig. 9); 0.49 mm tall opposite interval between second and third coxae where it is tallest; distinctly arched from PME to beginning of steep posterior declivity; surface distinctly granulate and with granulations often in rows; with a recurved groove just Figs. 9-12. Dysderina belinda sp. nov. Fig. 9. Carapace; dorsal view. Fig. 10. Posterior end of sternum; from below. Fig. 1 1. Venter, from below. Fig. 12. Epigynal area; from below. Figs. 13-19. Dysderina concinna sp. nov. Fig. 13. Carapace; left side. Fig. 14. Eyes and clypeus; from in front. Fig. 15. Sternum; from below. Figs. 16-18. Distal end of left palpal tarsus; nearly prolateral, nearly ventral and nearly dorsal, respectively. Fig. 19. Genital aperture. 1968 DYSDERINA IN C. AMERICA AND W. INDIES 9 behind posterior eyes; thoracic part regularly rounded along ventral margin and sharply narrowed shortly behind eyes. Eyes: six as usual in a fairly compact group; posterior row occupies about two- thirds of width of carapace at that level; viewed from above, posterior row slightly recurved, measured by posterior borders. Ratio of eyes ALE : PME : PLE = 9.5 : 9 : 8.5. ALE separated from one another by about one-half their long axis; separated from PME by about one-fourth of their long axis and from PLE by a line at one point. PME contiguous for nearly one-fourth of their circumference; separated from PLE by a line at one point. Pos- terior row only slightly wider than anterior row. Height of clypeus equal to fully three-fourths of the long axis of ALE. Chelicerae, maxillae and lip typical of females of the genus as far as observed. Sternum: rather strongly convex; with the usual intricate grooves and lobes typical of the genus; posterior end very bluntly termi- nated and only extended between bases of fourth coxae which are separated by nearly 1.5 times their width (Fig. 10). Legs: tibial index of first and fourth legs 9; spines essentially as described for D. silvatica Chickering (1951) with no very important differences. Abdomen: essentially as described for D. silvatica Chickering (1951); scuta and sclerite partly surrounding spinnerets quite typi- cal of the genus (Fig. 11). Epigynal area: very simple as usual (Fig. 12). Color in alcohol: essentially as described for other species; with no important differences. This species appears to be closely related to D. silvatica Chicker- ing, 1951 (Fig. 6) but differs from that species definitely with respect to the features of the epigynal area. In the former species the epigynal area is a nearly semicircular area with a minute dot near the posterior border. In D. belinda sp. nov. the epigynal area (Fig. 12) appears as a nearly semicircular area with a complete, conspicuous border and a faint central, longitudinal stripe. Records. Three females taken in the same locality with the holotype and during the same short period of nine days in Boquete, Panama, August, 1950, appear to belong here as paratypes. Dysderina concinna sp. nov. Figures 13-19 Holotype. The male holotype is from El Volcan, Panama, Au- gust, 1950. The name of the species is a Latin adjective referring to its neat, pleasing appearance. Description. Total length 2.27 mm. Carapace 1.01 mm long; 0.86 mm wide opposite interval between first and second coxae 10 breviora No. 296 where it is widest; 0.44 mm tall, and therefore, about half as tall as wide (Fig. 13); otherwise essentially as described for D. abdita sp. nov. Eyes: posterior row occupies about seven-ninths of width of carapace at that level; gently recurved as seen from above. ALE separated from one another by about five-eighths of their diameter; separated from PLE by one-eighth of their diameter and from PME by about three-eighths of their diameter. PME contiguous for about one-fifth of their circumference; only slightly separated from PLE. Posterior row wider than anterior row in ratio of 9 : 7. Height of clypeus equal to about nine-eighths of the diameter of ALE. Viewed from in front, posterior row gently procurved (Fig. 14). Chelicerae: basal segment 0.33 mm long; fang slender and evenly curved; otherwise essentially as recorded for D. abdita sp. nov. No teeth observed along fang groove. Maxillae and lip: ap- parently as described for D. abdita sp. nov. but no distal notch observed on maxillae (scarcity of paratypes prevents dissection for closer examination ). Sternum: scutiform in general; slightly longer than wide; intricately and obscurely grooved (Fig. 15); not con- tinued between fourth coxae which are separated by twice their greatest width; right first coxa abnormally small. Legs: tibial index of first leg 9, of fourth leg 10; several trichobothria observed but exact number and placement not determined; two tarsal claws. Spines: first leg with femoral ventral spines 0-lr-lr-2-2, with last two on retromargin reduced to little more than bristles; tibia with ventral spines 2-2-2-2-2-0; metatarsus with five ventral spines, three along promargin and two along retromargin, all ir- regularly placed. Second tibia with five ventral spines along pro- margin and four along retromargin; second metatarsus with two pairs of ventral spines; no true spines observed on legs three and four or on palp. Palp: essential features shown in Figures 16-18. Abdomen: 1.17 mm long; 1.04 mm wide near middle; pedicel, scuta, tracheal spiracles, openings to book lungs and spinnerets all essentially as described for D. abdita sp. nov.; genital aperture as indicated in Figure 19. Color in alcohol: essentially as described for other species in this paper with slight variations. Records. Two male paratypes were taken at El Volcan, Pan- ama, August, 1950. Dysderina craneae sp. nov. Figures 20-23 Holotype. The male is from Simla, Arima Valley, Trinidad. W. I. in the close vicinity of the William Beebe Tropical Research 1968 DYSDERINA IN C. AMERICA AND W. INDIES 11 Station, April 25, 1964. The species is named after Miss Jocelyn Crane, Director of the Station. Description. Total length 2.2 mm. Carapace 1.05 mm long; 0.79 mm wide opposite second coxae where it is widest; 0.42 mm tall; surface conspicuously but very irregularly granulate. Eyes: eye group occupies fully four-fifths of width of carapace at level of PE; viewed from above, posterior row gently recurved; ratio of eyes ALE : PME : PLE =10:9:9; positions and relationships essentially as stated for D. globina sp. nov. Chelicerae, maxillae and lip: all essentially typical of the genus as observed in this study. Sternum: deeply grooved in general pattern usually found in this genus; anterior third with a conspicuous transverse groove but otherwise nearly smooth; posterior two-thirds intricately grooved with primary and secondary grooves; fourth coxae sepa- rated by about six-fifths of their width. Legs: tibial index of first and fourth legs 9; spines almost identical with those recorded for D. globina sp. nov. Palp: all segments typical of males of the genus except the enlarged tarsus with its embolus and associated structures (Figs. 20-22); the degree of twisting of these structures seems to vary somewhat among the observed male paratypes. Ab- domen: typical of males of the genus with respect to scuta, indis- tinctive genital region, spinnerets, book lungs, tracheal openings, etc.; spinnerets partly surrounded by a ventral semicircular sclerite. Color in alcohol: carapace brown with heavily granulate areas darker than middorsal and nongranulate areas; dorsal ab- dominal scutum lighter brown; other parts as usual with variations yellowish and light brown. Female paratype. Total length 2.45 mm. Carapace 1.12 mm long; 0.88 mm wide; 0.45 mm tall; granulate essentially as in male. Eyes: eye group occupies about three-fourths of width of carapace at level of PE; seen from above, posterior row gently recurved; ratio of eyes ALE : PME : PLE = 10.5 : 10 : 9; ALE separated from one another by slightly less than their radius; other relationships typical of females of the genus as seen in this study; clypeus heavily bordered; height equal to about seven-tenths of the diameter of ALE. Mouth parts essentially typical of females of the genus. Sternum: essentially as in male except that anterior region is more irregular because of secondary grooves. Legs: 41=23 in order of length; tibial index of first and fourth legs 10; first leg with femoral ventral spines 0-0-lr-2-2-0, tibial ventral spines 2-2- 2-2-2-0, metatarsal ventral spines 2-2-2-0; second leg essentially as in first; third and fourth legs devoid of true spines. Abdomen: dorsal and ventral scuta nearly cover this part of body; chitinous 12 BREVIORA No. 296 scleritc completely surrounds spinnerets and anal tubercle but is very narrow dorsally. Epigynal area quite distinctive (Fig. 23). Records. Described female paratype taken with holotype from hay and weed debris by sifting. Simla, Arima Valley, Trinidad, W. L, April 25, 1964. Five male and seven female paratypes taken in vicinity of Simla, Arima Valley, or along roadside to Blanchesseusc, Trinidad, W. I., April, 1964. I igs. 20-23. Dysderina craneae sp. nov. Fig. 20. Left male palpal tarsus; nearly prolatcral view. Figs. 21-22. Tip of left male palpal tarsus; dorso- retrolateral view and nearly retrolateral view, respectively. Fig. 23. Epigynal area; from below. Figs. 24-25. Dysderina dura Chickering. Fig. 24. Tip of left palpal tarsus; Dearly prolatcral view. Fig. 25. Genital aperture of male. Figs. 26-27. Dysderina fitrtiva sp. nov. Fig. 26. Eyes from above. Fig. 27. Epigynal area; from below. Dysderina dura Chickering Figures 24-25 Dysderina dura Chickering, 1951: 208. The male holotype is in the Museum of Comparative Zoology; collected on Barro Colorado Id., Panama Canal Zone, July, 1936. 1968 DYSDERINA IN C. AMERICA AND W. INDIES 13 A detailed description of this species was published in 1951 to- gether with two figures illustrating certain features of the palp. Two more figures are added here to still further clarify the de- scription. About three dozen specimens have been added to the collection since the recognition of the holotype. All of these have come from the vicinity of the Panama Canal Zone. The species is by far the most numerous in the collection from Panama. The female is still not certainly known but is suspected of being D. sil- vatica Chickering. Dysderina furtiva sp. nov. Figures 26-27 Holotype. The female holotype is from Jamaica, W. I., St. Catherine Parish, 3 mi. north of Spanishtown, Oct. 16, 1957. The name of the species is a Latin adjective referring to its concealed habitat. Description. Total length 1.98 mm. Carapace 0.84 mm long; 0.69 mm wide opposite second coxae where it is widest; well rounded from just behind PE to posterior border which is only slightly notched; gently rising along median region from PME to beginning of steep posterior declivity; 0.25 mm tall; with no tho- racic groove; with a sparse covering of black hairs most conspicu- ous at beginning of posterior declivity. Eyes: posterior row occupies about eleven-fifteenths of width of carapace at that level; seen from above, posterior row very gently recurved measured by posterior borders but nearly straight (Fig. 26); ratio of eyes ALE : PME : PLE = 7.5 : 7 : 6.5; ALE separated from one another by a little less than one-third of their diameter, from PLE by about one-fifth of their diameter and from PME by about one- third of their diameter; PME contiguous for a short distance, barely separated from PLE; posterior row wider than anterior row in ratio of about 22 : 17; viewed from in front, posterior row defi- nitely procurved; height of clypeus equal to about four-fifths of the diameter of ALE. Chelicerae: basal segment 0.25 mm long; fang groove with a minute tooth on each margin near tip of the slender fang; otherwise essentially typical of females of the genus. Maxil- lae: somewhat more robust than described for D. silvatica Chicker- ing (1951) but essentially as in that species. Lip: wider at base than long in ratio of about 13 : 10; slightly concave along distal border; sternal suture gently procurved. Sternum: scutiform in general; as wide as long; strongly convex; only faintly lobulated opposite coxae; grooves and ridges lacking; this last feature very 14 breviora No. 296 unusual in the genus; posterior end not extended between fourth coxae which are separated by four-thirds of their width. Legs: 4123 in order of length; tibial index of first leg 8, of fourth leg 7; first leg with only one spine on prolateral side about two-thirds from base of femur; first tibia with ventral spines 2-2-2-2-1 p; first meta- tarsus with ventral spines 2-2- lp; second leg with no femoral spines; second tibia with ventral spines 2-2-2-2-0; second metatar- sus with three spines along promargin and two along retromargin with irregularity in placement; legs three and four probably to be considered lacking spines but some of the numerous bristles might be considered weak spines especially on fourth tibia; palp without terminal claw but with main still bristles resembling weak spines Abdomen: pedicel shorter and less corrugated than in typical spe- cies; dorsal scutum covers about nine-tenths of dorsum; ventral scuta extend about seven-ninths of distance from pedicel to spin- nerets; without a chitinous band guarding the spinnerets as in typi- cal species; position of colulus barely indicated. Epigynal area: very simple as usual in the genus but its form (Fig. 27) seems to be more or less distinctive. Color in alcohol: essentially typical of the genus except that the highly chitinized cephalothorax and scuta are somewhat lighter in color. This species also appears to be closely related to D. silvatica Chickering. 1951, but the epigynal area is unlike that seen in any other species. In /). furtiva sp. nov. this region is a simple, broad, slit-like aperture with a narrow border surrounding a plain white area. Records. Paratype females were taken with the holotype and numerous specimens are in the collection from the following locali- ties: St. Andrew Parish. Stony Mill, May 26, 1956 (C. C. Hoff); Coopers Hill, Feb. 10. 1955 (P. F. Bellinger); Jack's Hill Road, Dec. 6, 1957; St. Ann Parish, vicinity of Moneague, Nov. 7, 1957; St. Catherine Parish, Evarton, Nov. 29, 1957. Numerous females from Trinidad, W. I., for some time regarded as representing a new species are now referred to this species. No males have yet been associated with these numerous females. Dysderina globina sp. nov. Figures 28-31 Holotype. The male holotype is from Dominica, B. W. I.. Windward Islands, June 12, 1911. It will be deposited in the American Museum of Natural History, New York, N. Y. The name of the species is an arbitrary combination of letters. 1968 DYSDERINA IN C. AMERICA AND W. INDIES 15 I/// - 34 33 Figs. 28-31. Dysderina globina sp. nov. Fig. 28. Eyes from above. Figs. 29-30. Male palpal tarsus; nearly prolateral and retrolateral views, respec- tively. Fig. 31. Genital region from below. Figs. 32-34. Dysderina Humph- rey i sp. nov. Fig. 32. Eyes from above. Fig. 33. Sternum from below. Fig. 34. Epigynal area from below. Description. Total length 1.83 mm. Carapace 0.91 mm long; 0.73 mm wide opposite interval between first and second coxae where it is widest; 0.4 mm tall; other features essentially typical of males in the region under study. Eyes: eye group (Fig. 28) occu- pies about three-fourths of width of carapace at level of PE; seen from above, posterior row moderately recurved; ratio of eyes ALE : PME : PLE = 9 : 9 : 10 (some irregularities in outline noted); ALE separated from one another by three-tenths of their diameter and barely separated from PLE and PME; PME con- tiguous for more than one-fourth of their circumference and barely separated from PLE; height of clypeus equal to about three-fifths of the diameter of ALE. Chelicerae typical of males in the region under study; maxillae appear to be grooved and possibly divided distally. Lip wide at base and much narrowed distally. Sternum: 16 breviora No. 296 granulate; with a deep transverse groove shortly behind procurved sternal suture; with grooves, intricately developed in mam species, here very much reduced and simplified; fourth coxae separated by nine-sevenths of their width. Legs: tibial index of first and fourth legs 9; 4123 in order of length; first femur with ventral spines 0-0-lr-lp-lp; first tibia with ventral spines 2-2-2-2-2 (last two little more than stiff bristles); first metatarsus with ventral spines 2-2- lp-O; second femur apparently only with ventral spines 0-0-1 p-0; second tibia and second metatarsus essentially as in first with re- spect to ventral spines; third and fourth legs lacking true spines. Palp: all segments as usual in the genus except tarsus with its complicated embolus and associated apophyses (Figs. 29-30). Abdomen: essentially as described for /). meridina sp. now; genital region obscure but apparently distinctive (Fig. 31 ). Records. One male paratype. somewhat damaged, accompanies the holotype. A female, recently moulted, is also with the holotype but is not in good condition for description. Dysderina humphreyi sp. nov. Figures 32-34 Holotype, The female holotype is from Boquete. Panama, Au- gust, 1954. The species is named after Richard L. Humphrey, M.D., who, as an undergraduate student, was my assistant during a period of field work in Panama in the summer of 1954. Description. Total length 2.24 mm. Carapace 0.97 mm long; 0.75 mm wide opposite second coxae where it is widest; 0.41 mm tall; somewhat less regularly arched along median dorsal region than usual in females of the genus; otherwise essentially typical of the genus. Eyes: posterior row occupies about three-fourths of width of carapace at that level; almost straight but very slightly re- curved as viewed from above (Fig. 32); ratio of eyes ALE : PME : PLE = 8.5 : 8.5 : 7.5 (boundaries of eyes, especially of ALE. are quite irregular); ALE separated from one another by about their radius; ALE barely separated from PLE and from PME by about one-fourth of their diameter. Posterior row of eyes very closely crowded together; PME contiguous for one-fourth of their circumference; barely separated from PLE. Posterior row of eyes wider than anterior row in ratio of about 6:5; viewed from in front, posterior row of eyes definitely procurved; height of clypeus equal to about five-eighths of the diameter of ALE. Chelicerae, maxillae, and lip essentially as described for D. silvatica Chickering (1951) as far as observed. Sternum: general features as usual in 1968 DYSDERINA IN C. AMERICA AND W. INDIES 17 the genus; fourth coxae separated from one another by a little less than 1.5 times their width; pattern of grooves essentially as repre- sented in Figure 33. Legs: 4123 in order of length; tibial index of first leg 10, of fourth leg 9. Leg spines: first femur with five ventral spines, two on promargin and three on retromargin, not regularly paired; first tibia with five pairs of ventral spines; first metatarsus with five ventral spines irregularly placed; second femur with four ventral spines, one on promargin and three on retro- margin; second tibia with four pairs of ventral spines; second meta- tarsus with five ventral spines, three on promargin and two on retromargin; legs three and four apparently without true spines. Abdomen: scuta, pedicel, tracheal spiracles, etc., all essentially typical of females of the genus. Epigynal area: very simple as usual; essentially as represented in Figure 34. Color in alcohol: essentially typical of the genus with no unusual features. This species appears to be closely related to both D. plena O. P.-Cambridge and D. silvatica Chickering but the shape of the epigynal area is somewhat different than in the two previously known species and the dots along the posterior border are com- pletely lacking. Records. Eight paratype females collected August 4-11, 1954, in the close vicinity of Boquete, Panama. Dysderina improvisa sp. nov. Figures 35-36 Holotype. The female holotype is from Barro Colorado Island, Panama Canal Zone; collected in a Berlese funnel by Dr. James Zetek, May-October, 1946. The name of the species is a Latin ad- jective meaning unexpected. Description. Total length 2.26 mm, exclusive of the slightly extended spinnerets. Carapace 0.95 mm long; 0.77 mm wide op- posite second coxae where it is widest; 0.4 mm tall; otherwise essentially as in D. silvatica Chickering (1951). Eyes: posterior row occupies about three-fourths of width of carapace at that level (Fig. 35); viewed from above, gently recurved. Ratio of eyes ALE : PME : PLE = 8.5 : 7.5 : 7. ALE separated from one another by a little more than their radius; contiguous to PLE; separated from PME by nearly one-fourth of their diameter; PME contiguous for about one-fourth of their circumference; barely sepa- rated from PLE; posterior row wider than anterior row in ratio of 6:5; viewed from in front, posterior row definitely procurved. Height of clypeus equal to about three-fourths of the diameter of 18 HREVIORA No. 296 ALE. Chelicerae, maxillae and lip essentially as described for D. silvatica C flickering ( 1951 ). Sternum: general features as usual in the genus; pattern of grooves rather complicated; fourth coxae separated from one another by about seven-fourths of their width. Legs: 4123 in order of length; tibial index of first leg 9, of fourth leg 10. Leg spines: first femur with irregularities of ventral spines on right and left; first tibia with five pairs of ventral spines; first metatarsus with three pairs of ventral spines; second femur with ventral spines lr-lr-2-lp; second tibia with ventral spines 2-2-2- 2-1 p; second metatarsus with two pairs of ventral spines; third and fourth legs with few weak spines. Abdomen: essentially typical of females of the genus. Epigynal area: \er\ simple as usual; essen- tially as shown in Figure 36. Color in alcohol: also essentially typical of the genus with minor variations. 37 38 40 I igs. 35-36. Dysderina improvise sp. now Fig. 35. Eyes from above. Fig. 36. Epigynal area from below. Figs. 37-39. Dysderina intempina sp. nov. Figs. 37-38. Distal end of left male palpal tarsus; prolateral ;ind retrolateral views, respectively. Fig. 39. Genital area from below. Figs. 40-41. Dysderina meridina sp. nov.; distal end of left male palpal tarsus; prolateral and retro- lateral views, respectively. D. improvisa sp. nov. is also regarded as closely related to D. silvatica Chickering but its epigynal area is distinctly oval in shape, as compared to the semicircular shape of this region in the 1968 DYSDERINA IN C. AMERICA AND W. INDIES 19 previously known species. The pattern of grooves on the sternum appears to be somewhat distinctive also. Records. Seven female paratypes from Barro Colorado Id., Panama Canal Zone, were collected in a Berlese funnel by Dr. James Zetek, May-Oct., 1946. Six females were taken in the same locality as follows: June-Aug., 1936; April-May, 1942 (Zetek); June-Oct., 1946 (Zetek); Aug., 1954; January, 1958. One female was taken in the Canal Zone Forest Preserve, February, 1958. Dysderina intempina sp. nov. Figures 37-39 Holotype. The male holotype is from Boquete, Republic of Panama, August, 1950. The name of the species is an arbitrary combination of letters. Description. Total length 2.15 mm. Carapace 0.99 mm long; 0.79 mm wide opposite second coxae where it is widest; 0.53 mm tall; otherwise essentially typical of males of the genus. Eyes: pos- terior row occupies about five-sevenths of width of carapace at that level; slightly recurved as seen from above. Ratio of eyes ALE : PME : PLE = 8.5 : 8.5 : 9. ALE separated from one another by their radius; almost in contact with PLE; separated from PME by their radius. PME contiguous only for a short distance; only sepa- rated from PLE by a narrow line. Posterior row of eyes wider than anterior row in ratio of about 5 : 4. Height of clypeus equal to slightly less than the diameter of ALE. Viewed from in front, posterior row of eyes moderately procurved. Chelicerae: basal seg- ment 0.31 mm long; essentially typical of males of the genus; no dissection conducted because of scarcity of paratypes. Maxillae and lip: apparently typical of males of the genus. Sternum: grooves quite clearly delineated; very similar to those in D. obtina sp. nov.; fourth coxae separated by slightly more than their width. Legs: 4123 in order of length; tibial index of first leg 11, of fourth leg 9; trichobothria observed but exact number and placement not deter- mined. Leg spines: first femur with ventral spines 0-lr-lr-2-lp-0; first tibia with five pairs of long slender spines; first metatarsus with three pairs of ventral spines; second femur apparently with one weak ventral and two weak prolateral spines; second tibia with four pairs of ventral spines; second metatarsus probably with two pairs of ventral spines; these spines tend to be offset prolaterally or retrolaterally. Palp: apparently typical of males of the genus ex- cept for the tarsus whose distinctive features are shown in Figures 37-38. Abdomen: pedicel, scuta, etc., apparently typical of males 20 breviora No. 296 of the genus; genital region as shown in Figure 39. Color in alco- hoi: essentially typical of the genus with minor variations. Records. One paratype male was taken during the same period as the holotype and in the same locality. Females taken at the same time and in the same locality may also belong here but there can be no certainty at this time. Dysderina meridina sp. nov. Figures 40-4 1 Holotype. The male holotype is from San Jose, Costa Rica (En- rique Schmidt). No date oi collection is given but. presumably, the specimen was taken fairly recently. The name of the species is an arbitrary combination of letters. The holotype will be deposited in the American Museum of Natural History, New York, N. Y. The abdomen of the holotype is detached from the cephalothorax but otherwise is in a oood state of preservation. Description. Total length 2 mm. Carapace 0.94 mm long; 0.75 mm wide opposite second COXae where it is widest; about 0.4 mm tall; rises gradualk from PMF to opposite posterior border of sec- ond coxae and then descends steeply to posterior border where there is a narrow upturned shelf; surface finely granulate; median thoracic groove lacking. Eyes: si\ as usual; seen from above, posterior row gentl) recurved. Ratio of eyes ALE : PME : PLE =z 8.5 : 8 : 7.5. ALE separated from one another by slightly more than their radius; barely separated from I'll ■; separated from PME bv a little more than one-half their radius. PME contiguous for about one-fourth of their circumference: separated from PLE by a line. Height of clypeus equal to slightly less than the diameter of ALE; clypeus only moderately lobed in the middle of the ventral border. C'helieerae. maxillae and lip: all apparently typical of males of the genus from the regions under study. Sternum: also essentially typical of the genus in this general region; sternal grooves less intricate than in several other species; sternal suture procurved; fourth coxae separated by about four-thirds of their width. Legs: 42=13 in order of length; tibial index of first leg 11, of fourth leg 10; as far as observed, claws and trichobothria essentially as described for D. seclusa Chiekering (1951). Leg spines also essentially as described for D. seclusa. Palp: all seg- ments except tarsus simple and w ithout special modifications; tarsus with distinctive embolus and associated structures (Figs. 40-41). Abdomen: in general typical of the genus; dorsal and ventral scuta cover nearly the entire surface; spinnerets retracted and nearly in- visible; genital aperture typical of the genus. Color in alcohol: 1968 DYSDERINA IN C. AMERICA AND W. INDIES 21 carapace, sternum and abdominal scuta all a rich reddish brown; body of palpal tarsus light yellowish. Records. One poorly preserved male paratype accompanies the holotype. No date of collection is given. The female is unknown. Dysderina obtina sp. nov. Figures 42-45 Holotype. The male holotype is from El Volcan, Panama, Au- gust, 1950. The name of the species is an arbitrary combination of letters. Description. Total length 2.54 mm, including extended spin- nerets. Carapace 1.1 mm long; 0.88 mm wide opposite second coxae where it is widest; 0.52 mm tall; otherwise essentially typical of males of the genus. Eyes: posterior row occupies about two- thirds of width of carapace at that level; only slightly recurved as seen from above. Ratio of eyes ALE : PME : PLE = 9 : 8.5 : 8. ALE separated from one another by five-ninths of their greatest diameter; separated from PLE by about two-ninths of their di- ameter and from PME by one-third of their diameter. PME contiguous for about one-sixth of their circumference; separated from PLE by about one-eighth of the diameter of the latter. Pos- terior row of eyes wider than anterior row in ratio of about 5 : 4. Height of clypeus equal to about eleven-ninths of the diameter of ALE. Chelicerae, maxillae and lip essentially as described for D. abdita sp. nov. Sternum: only slightly longer than wide between second coxae; grooved in a distinct manner closely similar to the pattern shown by D. abdita sp. nov. and D. recondita Chickering (Fig. 42). Fourth coxae separated by nearly five-thirds of their width. Legs: 4123 in order of length; tibial index of first leg 10, of fourth leg 9; trichobothria observed on first femora and tibiae but number and placement not accurately determined. Leg spines: first femur with ventral spines 0-0-2-2-0 unevenly placed; first tibia with five pairs of ventral spines and a pair of distal bristles; first metatarsus with three pairs of ventral spines; second femur with two small ventral spines on promargin; second tibia with four pairs of ventral spines; second metatarsus as in first. Third and fourth legs devoid of true spines. Palp: general features as usual in males of the genus; specific tarsal features shown in Figures 43-44. Ab- domen 1.32 mm long exclusive of extended spinnerets; 0.92 mm wide slightly behind the middle; pedicel, scutta, etc., essentially as described for D. abdita sp. nov.; genital aperture as shown in Figure 45. Color in alcohol: also essentially as described for D. abdita sp. nov. with minor variations. ->-> HREVIORA No. 296 Records. \o paratypes have appeared in niv collection and the female is unknown. For some time the holotype was regarded as a deviate of />. recondita Chickering but careful attention to the male paip has convinced me that it must be considered to represent a new species as presented here. 47 . 42-45. Dysderina obtina sp. nov. Fig. 42. Sternum from below. I igs. 43-44. Distal end of left male palpal tarsus; nearly prolateral and retrolateral views, respectively. Fig. 45. Genital aperture from below. Figs. 4h-49. Dysderina plena O. P.-Cambridge. Figs. 46-47. Left male palpal tarsus; nearly prolateral and retrolateral views, respectively. Fig. 48. Distal end of male palpal tarsus; nearly ventral view. Fig. 49. Epigynal area from below. Figs. 50-52. Dysderina potena sp. nov. Fig. 50. Eyes from above. Fig. 51. Ventral scutum from below. Fig. 52. Epigynal area from below. 1968 DYSDERINA IN C. AMERICA AND W. INDIES 23 Dysderina plena O. P.-Cambridge Figures 46-49 Dysderina plena O. P.-Cambridge, 1894: 143; 1896: 192, pi. 17, fig. 7; pi. 24, fig. 4. Male and female syntypes from Teapa, Mexico, are in the British Museum (Natural History). F. P.-Cambridge, 1899: 44; Banks, 1909: 196; Petrunkevitch, 1911: 125; Roewer, 1933: 185; 1942: 283; Chickering, 1951: 211; Bonnet, 1956: 1638. As pointed out in 1951, the specimens reported from Panama by Banks (1929) and referred to this species really belong to other genera and species. The specimens reported by Williams (1941) and identified by Dr. W. J. Gertsch as Dysderina plena O. P.- Cambridge appear to be Dysderina silvatica Chickering (1951 ). The female from Teapa, Mexico, on loan from the British Mu- seum (Natural History), has an epigynal area (Fig. 49) very similar to that of D. silvatica Chickering. The intricate grooves on the sternum are somewhat different from those on D. silvatica Chickering but the two seem to be closely related. Male. From the British Museum (Natural History). Total length about 2.1 mm. Carapace 0.98 mm long. 0.78 mm wide opposite second coxae where it is widest; about 0.52 mm tall oppo- site interval between second and third coxae where it is tallest; descent nearly precipitous from highest point to posterior border; surface finely granulate; without a median thoracic groove or pit. Eyes: posterior row occupies a little more than three-fourths of width of cephalic region at this level; gently recurved as seen from above. Ratio of eyes ALE : PME : PLE = 10 : 9.5 : 10. All eyes are somewhat irregular in outline; PME and PLE are more oval than ALE. ALE separated from one another by nearly one-third of their diameter. PME contiguous for about one-fourth of their circumference; PLE nearly contiguous to PME posteriorly but separated anteriorly; PLE and ALE nearly contiguous; posterior row longer than anterior row in ratio of about 5:4; viewed from in front, posterior row procurved. Height of clypeus a little more than two-thirds of the diameter of ALE; border of clypeus strongly chitinized. Sternum finely granulate and intricately grooved about as usual in the genus. Genital aperture a narrow slit surrounded by a broad margin. Important features of the palp shown in Figures 46-48. Other features essentially typical of males of the genus. 24 breviora No. 296 Dysderina potena sp. nov. Figures 50-52 Holotype. The female holotype is from El Volcan, Panama. August. 1950. The name of the species is an arbitrary combination of letters. Description. Total length, including spinnerets. 2.8 mm. Cara- pace 1 .08 mm long; 0.97 mm wide opposite second coxae where it is widest: 0.4S mm tall; gradually arched from PLE to posterior border; finely but irregularly granulate; without any visible thoracic groove or pit. Eyes: seen from above, posterior row straight if measured b\ posterior borders; posterior row occupies about two- thirds of width of carapace at that level; seen from in front, pos- terior row gentl) procurved. Ratio oi eyes ALE : PME : PLE = 9.5 : 9 : 8. ALE separated from one another by a little more than their radius; barely separated from PLE (Fig. 50). Height of cly- pens equal to a little more than the diameter of ALE. Chelieerae. maxillae and lip; essential!) typical of females of the genus as ob- served in this studs. Sternum only slightly longer than wide; with the usual intricate grooves; quite convex; fourth coxae separated b\ about 1.5 times their width, legs: 4123 in order of length; tibial index of first and fourth legs 9; spines on legs essentially as described for />. silvatica Chickering (1951). Abdomen: essen- tialK typical oi females of the genus with minor variations; ventral scutum as shown in Figure 5 I . Epigynal area: appears to have cer- tain obscure but distinctive features (Fig. 52). Color in alcohol: carapace irregularly darkened along posterior declivity and lateral sides; generally a rich reddish brown. This species appears to be closely related to /). plena O. P. Cambridge but differs from that species most conspicuously with respect to its epigynal area which is in the form of a modified oval opening ( Fig. 52 | with a distinct anterior border and a series of faintly indicated granules along the posterior border. The height of the clypeus is somewhat greater than usual in the genus. Ren >nls. Two paratype females taken August, 1950, in vicinity of El Volcan, Panama. One female from Boqucte. Panama, Au- gust, 1954 is somewhat doubtfully assigned to this species. Dysderina princeps Simon Figures 53-54 Dysderina princeps Simon, 1891, 557. The female holotype from St. Vin- cent. B. W. I., is in the British Museum (Natural History). Simon, 1893: 290, 304; Petrunkevitch, 1911: 125; Roewer, 1942: 283; Bonnet, 1956: 1638. 1968 DYSDERINA IN C. AMERICA AND W. INDIES 25 So far as I have been able to determine, the species has never been correctly reported since its first recognition. In 1893, how- ever, Simon published his figure 260 which is a drawing of a male palp and it is labelled D. princeps. The figure closely resembles the original drawing (fig. 1 ) accompanying the brief description of D. principalis Simon. The following facts are derived from the study of a female from St. Vincent, B. W. I. on loan from the British Museum (Natural History) : Total length 2.66 mm (the author of the species gave the length as 4 mm). Carapace 1.17 mm long; 0.97 mm wide oppo- site second coxae where it is widest; otherwise essentially typical of females of the genus as seen during this study. Eyes in general as usual in the genus (Fig. 53) ; ratio of eyes ALE : PME : PLE = 9 : 8.5 : 9.5. Clypeus somewhat higher than that in D. spinigera Simon but otherwise closely similar. Sternum intricately but not deeply grooved. Legs 1 and 2 with the usual long ventral spines essentially typical of the genus. Abdomen with the typical scuta and related parts. Epigynal area more distinctive than usual (Fig. 54). Dysderina principalis (Keyserling) Figures 55-56 Oonops principalis Keyserling, 1882: 296, pi. 11, fig. 16. The male holotype from Colombia is in the British Museum (Natural History). Dysderina principalis,Simon, 1891, 557, pi. 42, fig. 1. Simon, 1893: 304; Petrunkevitch, 1911: 126; 1928: 87; Roewer, 1942: 283; Bonnet, 1956: 1638. (Not D. principalis Simon from St. Vincent, B. W. I.) Simon was correct in transferring Oonops principalis Keyserling to his new genus Dysderina but he was in error when he identified a species of this new genus from St. Vincent, B. W. I., as being D. principalis described from Colombia. Apparently Keyserling had only the male upon which to base his species. This specimen has been on loan from the British Museum (Natural History). It is dismembered, but the right palp is in good condition and estab- lishes Simon's error very clearly. In order to aid in clarifying this confusion I am offering two figures of the palpal tarsus (Figs. 55-56). Keyserling's small figure 16 closely resembles those I am providing. Dysderina recondita Chickering Figures 57-59 Dysderina recondita Chickering, 1951: 217, fig. 4. The male holotype and two paratypes from Boquete, Panama, July, 1939, are in the Museum of Comparative Zoology. 26 BREVIORA No. 296 During my brief visit to Boquete, Panama, in August, 1954, I collected eleven additional males. The female is unknown but it seems probable that it is among the different females now recog- nized from the mountainous regions of Panama. Figures 58-59 are added to the original drawing of the male palp; Figure 57 shows the essential features of the sternal grooves. v 53-54. Dysderina princeps Simon. Fig. 53. Eyes from in front. I ig, 54. Epigynal area from below. Figs. 55-56. Dysderina principalis < Keyserling). Two views of right m;ile palpal tarsus. Figs. 57-59. Dysderina recondita Chickering. Fig. 57. Sternum from hclow. Figs. 58-59. Two views of distal end of left male palpal tarsus. Dysderina rigida sp. nov. Figures 60-61 Holotype. The female holotype is from El Volcan, Panama, Au- gust, 1950. The name of the species is a Latin adjective referring to its strongly chitinized scuta, pedicel and other outer parts. Description. Total length 2.49 mm. Carapace 1.05 mm long; 0.88 mm wide opposite second coxae where it is widest; 0.44 mm tall; other features essentially as described for D. silvatica Chicker- ing (1951). Eyes: posterior row gently recurved, seen from 1968 DYSDERINA IN C. AMERICA AND W. INDIES 27 above; occupies about five-sevenths of width of carapace at that level. Ratio of eyes ALE : PME : PLE = 9:9:8. ALE separated from one another by slightly more than their radius; only slightly separated from PLE and from PME by about one-fourth of their diameter. PME contiguous along about one-fourth of their circum- ference and longer than wide in ratio of about 9 : 7.5; separated from PLE by one-sixth of their long diameter. Posterior row wider than anterior row in ratio of about 13 : 11. Height of clypeus slightly greater than the diameter of ALE. Chelicerae: vertical, essentially parallel; outer margin with a shallow groove at end of upper third and a slight swelling near middle; a single tooth along margin of fang groove appears to be definitely promarginal (ob- served on paratype); otherwise essentially as reported for D. sil- vatica (1951). Maxillae, lip and sternum also essentially as re- ported for D. silvatica (1951) except for the system of grooves on sternum (Fig. 60). Legs: 4123 in order of length; tibial index of first and fourth legs 9; trichobothria observed on palpal tibia and tarsus but number and placement not accurately determined. Leg spines: first femur with five ventral spines not evenly paired; first tibia with five pairs of ventral spines; first metatarsus with three pairs of ventral spines. Second leg with ventral spines nearly as on first leg; with only minor differences. Abdomen: spinnerets with a lightly chitinized ventral semicircular sclerite partially surrounding these organs; other features essentially typical of females of the genus. Epigynal area: simple as usual but apparently with obscure distinctive features (Fig. 61). Color in alcohol: essentially typical of the genus with minor variations. This species is also regarded as being rather closely related to D. plena O. P.-Cambridge. It differs from that species most con- spicuously with respect to its epigynal area which is in the shape of a modified oval region with a narrow, transverse area along its posterior border (Fig. 61). The pattern of grooves on its sternum also appears to be somewhat distinctive (Fig. 60). Records. Several female paratypes taken with the holotype, El Volcan, Panama, August, 1950. Dysderina seclusa Chickering Figures 62-64 Dysderina seclusa Chickering, 1951: 213, fig. 5. The male holotype and five male paratypes from Barro Colorado Island, Panama Canal Zone, are in the Museum of Comparative Zoology. All of these were taken in a Berlese funnel by Dr. Zetek, July, 1943 -March, 1944, and June- October, 1946. 28 HREVIORA No. 296 Since the original description was written, I have added four males to the collection as follows: June-October, 1946 (Zetek); June-August, 1949 (Zetek); August, 1954. I have been unable to match these males with females in any satisfactory manner. With the hope of further clarifying the status of the species three addi- tional figures are offered. \^ U,1 fT\ W- 1^-Ceo 63 67 •■ .->— ,- \ 62 x ^MW,/// Figs. 60-61. Dysderina rigida sp. nov. Fig. 60. Sternum from below. Fig. 61. Epigynal area from belou. Figs. 62-64. Dysderina seclusa (bickering. Fig. 62. Sternum from below. I i;^. 63. Left male palpal tarsus: ventral view. Fig. 64. Distal end of left male palpal tarsus, nearly dorsal view. Figs. 65-68. Dysderina simla sp. nov. Fig. 65. Left male palpal tarsus; retrolatcral view. F - i lt . 66. Distal end of left male palpal tarsus; nearly ventral view. Fig. 67. Genital area of male holotype. Fig. 68. Epigynal area of female paratype. Dysderina silvatica Chickering Dysderina silvatica Chickering, 1951: 217, fig. 6. The female holotype and numerous paratypes from several localities in the Panama Canal Zone and El Valle, Panama, are in the Museum of Comparative Zoology. 1968 DYSDERINA IN C. AMERICA AND W. INDIES 29 Since my report in 1951 I have added numerous females of this species to the collection from the following localities: Barro Colo- rado Island, Canal Zone Forest Preserve and Summit Gardens all in Panama Canal Zone. Although the evidence is not conclusive, I think it probable that these females belong with D. dura Chickering. Dysderina simla sp. nov. Figures 65-68 Holotype. The male holotype is from Simla, Arima Valley, Trinidad, W. I., April 5, 1964. The name of the species is a noun used in apposition after the type locality. Description. Total length 1.69 mm. Carapace 0.88 mm long; 0.75 mm wide opposite second coxae where it is widest; 0.47 mm (all; median thoracic groove lacking; smooth along broad, median area from PLE to beginning of posterior declivity; irregularly granulate along lateral sides and much of posterior declivity; fits closely to anterior end of abdomen which is unusual in this genus. Eyes: six as usual in a compact group which occupies nearly three-fourths of width of carapace at level of PE; viewed from above, posterior row slightly recurved. Ratio of eyes ALE : PME : PLE = 8:9:9. ALE separated from one another by slightly more than their radius; other relationships of eyes typical of the genus in the region under study. Height of clypeus equal to about three-fourths of diameter of ALE. Chelicerae, maxillae and lip all appear to be typical of the genus. Sternum: granulate; intricately but not deeply grooved and lobed; the usual anterior, transverse groove is complete, conspicuous and only slightly behind procurved sternal suture; only slightly longer than wide just behind second coxae; fourth coxae separated by nearly twice their width. Legs: 4123 in order of length; tibial index of first leg 10, of fourth leg 11. Spines on legs about as usual in males of the genus. Palp: all seg- ments simple and without special modifications except the tarsus which has distinctive features (Figs. 65-66). Abdomen: typical of the genus in general; with dorsal and ventral scuta less strongly chitinized than usual (all paratypes appear to be the same in this respect); genital region quite clear and rather distinctive (Fig. 67). Color in alcohol: carapace essentially as described for D. craneae sp. nov.; legs and mouth parts a variable yellowish; sternum a deeper yellowish; abdomen with dorsal and ventral scuta a light yellowish in contrast to the usual much darker coloration; epigastric scutum somewhat darker; the free areas not covered by scuta are whitish. 30 breviora No. 296 Female paratype. Total length 1.98 mm. Carapace 0.9 mm long; 0.84 mm wide opposite interval between second and third coxae where it is widest; 0.36 mm tall; surface granulate essen- tially as in male. Cephalothorax closely contiguous to abdomen as in male. Eyes essentially as in male. Chclicerae, maxillae and lip: all essentially typical of females of the genus. Sternum: essentially as in male but with granulation more extensive; fourth coxae sepa- rated b\ neark twice their width. Legs: 41 23 in order of length; tibial index of first and fourth legs 1 1; spines essentially typical of the genus. Abdomen: general appearance as usual in the genus; dorsal scutum somewhat less extensive than usual; epigastric scutum about as usual: ventral scutum much smaller than usual and reaches slightly less than half way from genital groove to spin- nerets; a narrow, ventral, semicircular sclcrite partly surrounds the spinnerets. Epigynal area: quite distinctive but difficult to repre- sent accurately in a drawing (Fig. 68). Color in alcohol: all parts except the abdomen as in male; because of the smaller scuta much more o\~ the abdomen appears nearly white than in the male; posterior to the ventral scutum are several vague darker spots which in some paratvpes are quite distinct brownish, irregular spots. Records. The described female paratype was taken with the holotype. Three male and seven female paratvpes are in the col- lection from Simla. Arima Valley, and along the side of the road from Simla to Blanchesscuse. Trinidad, W. I., March 31 -April 25, 1964. Dysderina soltina sp. nov. Figures 69-75 Dysderina principalis, -Simon, 1891: 557, pi. 42, fig. 1. Male and female syni\pes from St. Vincent B. W. I. are in the British Museum (Natural History I. (Not D. principalis (Keyserling) from Colombia.) In his study of the genus Dysderina from St. Vincent, B. W. I., Simon selected what he considered to be a male and a female of D. principalis (Keyserling), gave a brief description and furnished a figure of the male palp. As a result of a study of these specimens together with an example of Keyserling's species from Colombia on loan from the British Museum (Natural History), I am obliged to disagree completely on the identification of these specimens. They seem to me to represent a new species which is described below. Holotype. The male holotype is from St. Vincent, B. W. L, and will be returned to the British Museum (Natural History). The name of the species is an arbitrary combination of letters. 1968 DYSDERINA IN C. AMERICA AND W. INDIES 31 Description. Total length 1.91 mm (recorded as 3.5 mm in Simon's description). Carapace 0.92 mm long; 0.73 mm wide opposite second coxae; 0.35 mm tall; only gradually narrowed be- hind eyes; surface irregularly granulate as usual; otherwise typical of the genus. Eyes: six as usual; seen from above, posterior row gently recurved; ratio of eyes ALE : PME : PLE = 8 : 7 : 7 (out- lines poorly defined). ALE separated from one another by their radius. PME contiguous for about one-fourth of their circum- ference; barely separated from PLE. Height of clypeus equal to about five-eighths of the diameter of ALE. Chelicerae, maxillae and lip typical of the genus as far as observed. Sternum: typical of the genus in general; major and minor grooves and depressions moderately complex; extended as usual between fourth coxae and articulated with pedicel; fourth coxae separated by five-thirds of their width. Legs: 4123 in order of length; tibial index of first leg 10, of fourth leg 9; ventral spines on first leg as shown in Figures 71-73; fewer ventral spines on second leg. Palp: all segments ex- cept tarsus simple and unmodified as usual; tarsus considerably inflated and with distinctive terminal structures difficult to observe clearly (Figs. 69-70). Abdomen: all parts in general typical of males of the genus; scuta cover nearly the entire surface. Color in alcohol: also typical of the genus with few minor variations. Female paratype. This is apparently the specimen which Simon regarded as the female of D. principalis (Keyserling). For the present I am regarding it as the female of the new species D. sol- tina. Total length 1.98 mm (Simon gave length as 4 mm). Cara- pace 0.9 mm long; 0.73 mm wide; 0.4 mm tall; general features typical of females of the genus. Leg spines: the first femur has five ventral spines, two on promargin and three on retromargin (Fig. 74); other segments with spines much as in male. Abdomen with scuta and other parts essentially typical of the genus. Epigynal area: obscurely distinctive as usual (Fig. 75). Records. As far as I have been able to determine the two specimens described here are the only members of the species known at present. Dysderina spinigera Simon Figures 76-79 Dysderina spinigera Simon, 1891, pi. 42, figs. 2-3. Male and female syn- types from St. Vincent, B. W. I., are in the British Museum (Natural History). Simon, 1893: 304; Petrunkevitch, 1911: 126; Roewer, 1942: 283; Bonnet, 1956: 1639. 32 BREVIORA No. 296 I igs. 69-75. Dysderina soltina sp. nov. Fig. 69. Left male palpal tarsus; nearly dorsal view. Fig. 70. Distal end of left male palpal tarsus; retrolatcral view (more enlarged). Figs. 71-73. Ventral spines on first left femur, tibia and metatarsus, respectively. Hg. 74. Ventral spines on first left femur of female paratype. Hg. 75. Epigynal area of female paratype from below. Figs. 76-79. Dysderina spinigera Simon. Fig. 76. Eyes from in front. Fig. 77. Left male palp; retrolateral view. Fig. 78. Left male palpal tarsus; dorsal view. Fig. 79. Epigynal area from below. Simon reported both sexes of this species from St. Vincent and Venezuela where he regarded it as widespread. I have not seen the specimens from Venezuela but. on the basis of my study of this 1968 DYSDERINA IN C. AMERICA AND W. INDIES 33 genus, I consider it unlikely that they are the same as these from St. Vincent. I was disappointed in my failure to collect any speci- mens of the genus Dysderina during my brief visit to this island in October, 1966. The following facts relating to this species have been recorded as a result of my study of a male and two females on loan from the British Museum (Natural History). Male. Total length 1.61 mm (Simon gave length of male as 4 mm). Carapace 0.86 mm long; 0.66 mm wide opposite second coxae where it is widest; otherwise essentially typical of the genus as observed in this study. Eyes: essentially as shown in Figure 76; ratio of eyes ALE : PME : PLE = 10 : 9 : 9 (outlines irregular); ALE closer together than usual in the genus; clypeus with strongly chitinized ventral border and conspicuously lobed ventrally in front (Fig. 76). Sternum moderately grooved in fairly typical manner. Legs essentially typical in respect to spination and rela- tive lengths; trichobothria observed but exact distribution unde- termined. Palp: quite distinctive; appearance very different from Simon's figure 3, 1891; essentials shown in Figures 77-78. Abdo- men: with typical dorsal and ventral scuta; genital aperture a simple, minute slit. Female. Total length, exclusive of projecting spinnerets and slightly exposed chelicerae 2.09 mm; including posterior spin- nerets and slightly exposed chelicerae, length is 2.2 mm (Simon gave length as 4.5 mm). General form and external features essen- tially like those of male. Abdomen: with dorsal and ventral scuta somewhat less extensive than in male; epigynal area very distinc- tive; unlike that seen in other species (Fig. 79). Dysderina watina sp. nov. Figures 80-84 Holotype. The male holotype is from Turrialba, Costa Rica, July 25-Aug. 15, 1965. The name of the species is an arbitrary combination of letters. Description. Total length 2.24 mm. Carapace 1.03 mm long; 0.86 mm wide opposite second coxae where it is widest; 0.45 mm tall opposite second coxae; moderately grooved immediately be- hind PE; regularly arched from groove to beginning of moderately steep posterior declivity; surface irregularly granulate; otherwise essentially typical of the genus. Eyes: six in two rows as usual; posterior row gently recurved, seen from above; posterior row occupies about seven-ninths of width of carapace at that level. 34 breviora No. 296 Ratio of eyes ALE : PME : PLE =10:8:9. ALE separated from one another by about their radius, from PLE by nearly one- fifth of their diameter and from PME by about three-tenths of their diameter. PME barely separated from one another and from PLE by about three-sixteenths of their diameter. Height of clypeus nearly equal to long diameter of ALE. Chelicerae. maxillae and lip typical of the genus as far as observed. Sternum: clearly but less intricately grooved than in several other species; posterior coxae separated from one another bv about 1.5 times their width; otherwise essentially typical of the genus. Legs: 4123 in order of length; tibial index of first leg 8, of fourth leg 7; spines con- spicuous! v long and slender but otherwise essentially tvpical of males of the genus. Palp: all segments except tarsus simple and unmodified; tarsus with distinctive distal features as shown in Fie- ures 80-81. Abdomen: essentially typical of males of the genus. Color in alcohol: lighter than usual in the genus; carapace, pedicel and sternum a medium yellowish brown; abdomen and legs yellow- ish with variations. Female paratype. Total length 2.75 mm exclusive of slightly projecting spinnerets and chelicerae; including these parts total length is 2.9 mm. Carapace 1.1 mm long; 0.92 mm wide opposite interval between second and third coxae; about 0.44 mm tall; rather deeply grooved immediately behind PE and then gently arched to posterior declivity; otherwise typical o\' the genus. Eves: PME contiguous for neark one-fourth of their circumference; ratio of eyes ALE : PME : PLE = 10 : 10 : 9.5; other features essentially as in male. Height of clypeus immediately below ALE equal to slightly less than the diameter of these eyes; clypeus con- spicuously lobed in middle between bases of chelicerae. Cheli- cerae, maxillae and lip essentially typical of females of the genus. Sternum: with the usual pattern of major grooves and other incon- spicuous irregularities; fourth coxae separated by five-thirds of their width. Legs: 4123 in order of length; tibial index of first leg 9, of fourth leg 10; spines of first and second legs conspicuously long but essentially typical of females of the genus with minor variations. Abdomen: scuta, pedicel, spiracles, etc. nearly typical of females of the genus (Figs. 82-83); a very clear narrow sclerite partially surrounds spinnerets on ventral side; what appears to be a chitinous dot marks the position of the reduced colulus. Epigynal area: obscurely distinctive (Fig. 84). Color in alcohol: much more colorful than in male; carapace, sternum and abdominal scuta all a rich reddish brown with lateral sides of carapace irregularly dark- ened by granulations; areas of abdomen not covered by scuta clear white; legs brownish yellow with variations. 1968 DYSDERINA IN C. AMERICA AND W. INDIES 35 Records. The described female paratype together with three other females were all taken from weed and hay debris in the same general locality from which the male holotype was taken, Turrialba, Costa Rica, July 25-August 15, 1965. It seems reasonable to as- sume that the male holotype and females belong together but there can be no absolute certainty. 86 80 Figs. 80-84. Dysderina watina sp. nov. Figs. 80-81. Distal end of left male palpal tarsus; prolateral and retrolateral views, respectively. Figs. 82-83. Dorsal and ventral views of abdomen of female paratype. Fig. 84. Epigynal area of female paratype, from below. Figs. 85-86. Dysderina zinona sp. nov. Fig. 85. ALE and clypeus from in front. Fig. 86. Epigynal area from below Dysderina zinona sp. nov. Figures 85-86 Holotype. The female holotype is from St. Vincent, B. W. I., Mt. Soufriere, June 1, 1965 (R. T. Bell). The name of the species is an arbitrary combination of letters. The specimen serving here as the holotype of a new species is somewhat damaged but its most important features are still in good condition. 36 breviora No. 296 Description. Total length about 2.28 mm. Carapace 1.08 mm long; 0.86 mm wide opposite second coxae where it is widest; 0.54 mm tall; irregularly granulate as usual; otherwise essentially typical of females of the genus. Eyes: in general, as usual in the genus; viewed from above, posterior row gently recurved and occu- pies about seven-tenths of width of carapace at that level. Ratio of eyes ALE : PME : PLE = 9 : 8.5 : 8. ALE separated from one another by nearly their radius; from PLE by nearly one-half their radius and by slightly more than this from PME, which are con- tiguous for about one-fourth of their circumference and barely separated from PLE. Clypeus conspicuously lobed in front (Fig. 85); height immediately below ALE equal to nearly two-thirds of the diameter of these eves. Chelicerae. maxillae and lip typical of females of the genus as far as observed. Sternum: fairly typical also; major grooves and secondary depressions make a rather intri- cate pattern; fourth coxae separated by a little more than their width. Legs: 4123 in order oi length; tibial index of first leg 10. o\' fourth leg l); first leg with five ventral spines on femur as in Figure 74; first tibia with five ventral spines on promargin and tour on retromargin; first metatarsus with three \entral spines on promargin and two on retromargin. Second leg with somewhat fewer ventral spines than occur on first leg. Third and fourth legs without true spines. Abdomen: considerably damaged but ap- parently quite typical oi females of the genus. Epigynal area: un- damaged; rather distinctive (Fig. 86). Color in alcohol: carapace a rich reddish brown; sternum somewhat lighter; legs yellowish brown with variations; abdominal scuta probably much like cara- pace but somewhat lighter like the sternum. D. zinona sp. nov. is regarded as closely related to D. princeps Simon. The epigynal area (Fig. 86) is quite different from that region in D. princeps Simon (Fig. 54) and much simpler. The clypeus is also conspicuously lobed in front (Fig. 85). Records. No paratypes have appeared in the collection. BIBLIOGRAPHY Banks, Nathan 1909. Arachnida from Costa Rica. Proc. Acad. Natur. Sci. Phila- delphia, April, 1909: 194-234, 2 pis. 1929. Spiders from Panama. Bull. Mus. Comp. Zool.. 69: 53-96, 4 pis. Bonnet, Pierre 1956. Bibliographia Araneorum. Toulouse. Vol. 2 (2). 1968 DYSDERINA IN C. AMERICA AND W. INDIES 37 Bryant, Elizabeth 1942. Notes on the spiders of the Virgin Islands. Bull. Mus. Comp. Zool., 89: 317-363, 3 pis. 1948. The spiders of Hispaniola. Bull. Mus. Comp. Zool., 100: 331- 447, 12 pis. Cambridge, O. P.- and F. P.-Cambridge 1889- Arachnida-Araneida. Vols. I-II. In: Biologia Centrali-Ameri- 1905. cana. Dulau & Co., London. Chickering, Arthur M. 1951. The Oonopidae of Panama. Bull. Mus. Comp. Zool., 106: 207-245,31 figs. COMSTOCK, J. H. 1940. The Spider Book. Revised and edited by W. J. Gertsch. Double- day, Doran & Company, Inc., New York. Gertsch, W. J. 1941. Report on some arachnids from Barro Colorado Island, Canal Zone. Amer. Mus. Novit., No. 1146: 1-14, 4 pis. Kaston, B. J. 1948. Spiders of Connecticut. State of Connecticut. State Geological and Natural History Survey, Bull. 70: 1-874. Keyserling, Graf Eugen von 1881. Neue Spinnen aus Amerika. III. Verhandl. Zool. Bot. Ges. Wien, 31,269-314, pi. 11. Petrunkevitch, Alexander 1911. A synonymic index-catalogue of spiders of North, Central, South America, etc. Bull. Amer. Mus. Natur. Hist., 29: 1-809. 1928. Systema Araneorum. Trans. Connecticut Acad. Arts and Sci., 29: 1-270. 1939. Catalogue of American Spiders. Pt. 1. Trans. Connecticut Acad. Arts and Sci., 33: 135-338. ROEWER, C. FR. 1933. Araneen aus mexikanischen Ameisenwohnpflanzen. Zool. Anzei- ger, 102: 183-188. 1942. Katalog der Araneae. 1: 1-1040. Bremen. Simon, E. 1891. On the spiders of the island of St. Vincent. Pt. 1. Proc. Zool. Soc. of London, Nov. 17, 1891: 549-575. 1892- Histoire naturelle des Araignees. Deuxieme Edition. 2 Vols. 1903. Librarie Encyclopedique de Roret, Paris. Williams, Eliot C, Jr. 1941. An ecological study of the floor fauna of the Panama rain forest. Bull. Chicago Acad. Sci., 6: 63-124. (Received 27 September 1967.) BREVIORA Museunri of Comtipsiratlve Zoology Cambridge, Mass. 11 October, 1968 Number 297 POPULATION STRUCTURE OF THE ASTHENES FLAMMULATA SUPERSPECIES (AVES: FURNARNDAE) Francois Vuilleumier1 INTRODUCTION While conducting a survey of speciation phenomena in twenty- five genera of Andean birds, I found that the existing classifications of these birds were often insufficient for my purposes. I had to un- dertake systematic studies in each genus before I could proceed to an analysis of species formation. In view of the groundwork already laid by Hellmayr, Chapman, Zimmer, Peters, and others, it may seem surprising that a student of speciation should find it necessary to do so much systematic work. However, these earlier systematists were primarily concerned with the purely formal aspects of assign- ing samples to species and subspecies, rather than interested in the process of speciation; their works contain little information on population structure (in the sense of Mayr, 1959:294) or species relationships. My own taxonomic work on Andean birds was de- signed to fill this gap. This is the first in a proposed series of papers on these Andean birds. My purpose has been to gain an understanding of the process of speciation in each genus sampled through study of population structure and interspecies relationships, and then to synthesize this information, arriving at a general picture of speciation phenomena in Andean birds. In the present paper, an effort has been made to uncover the evolutionary relationships of the component members of the Asthenes flammulata superspecies by means of an analysis of the distribution and geographical variation of these spinetails. This study was undertaken at the Museum of Comparatize Zool- ogy of Harvard University. My investigations were based largely on 1 Research Fellow, Department of Ornithology, American Museum Of Natural History, New York, New York 10024. 2 BREVIORA No. 297 the collections of this institution, but I have also examined speci- mens from other museums. When referring to individual specimens, the following abbreviations are used: MCZ = Museum of Com- parative Zoology; ANSP = Academy of Natural Sciences of Phila- delphia; CM — Carnegie Museum. Pittsburgh; FMNH = Field Museum of Natural History, Chicago; MVZ = Museum of Vertebrate Zoology, Berkeley; MP = Museum National d'Histoire Naturelle, Paris. ACKNOWLEDGMENTS I am most indebted to Ernst Mayr for his guidance, advice, and stimulation throughout my studies of speciation of Andean birds, which were becun at his suggestion. I wish to extend mv sincere thanks to the following persons, who have helped me during mv examination of specimens, who have arranged loans for me. or who have provided me with valuable in- formation: Raymond A. Paynter, Jr. (Museum of Comparative Zoology), James Bond (Academy of Natural Sciences). Dean Ama- dou and Charles O'Brien (American Museum of Natural History), Kenneth C. Parkcs (Carnegie Museum), Emmet R. Blake (Field Museum of Natural Histor\ ), Ned K. Johnson (Museum of Ver- tebrate Zoology), I. C. G. Galbraith (British Museum [Natural History]). Jean Dorst and Jean Prevost (Museum National dllis- toire Naturelle ). Financial assistance to delra\ the costs of field work was pro- vided through the Chapman Fund of the American Museum of Natural History, National Science Foundation funds from grants G-19729 and GB-3167 to Harvard University's Committee on Evolutionary Biology, and the Bourse Federale de Voyages de la Societe Helvetique des Sciences Naturelles in Switzerland. A visit to the Academy of Natural Sciences in Philadelphia was financed by the University of Massachusetts — Boston. I am very grateful to these agencies for their generous support of my work. Permission to reproduce maps published by the Department of Geography of the University of Chicago (Goode Base Maps Series. Copyright by the University of Chicago) was obtained through the courtesy of Mrs. Carole J. Major. THE ASTHENES FLAMMULATA SPECIES GROUP The genus Asihenes Reichenbach, 1853, consists of about 18 species of small spinetails (Furnariidae) living in western and 1968 ASTHENES FLAMMULATA SUPERSPECIES 3 southern South America. At least four species groups can be rec- ognized in Asthenes. Since this arrangement of the genus will be published elsewhere, it is sufficient to mention here that what I call the Asthenes flammulata species group comprises four nominal species: Asthenes flammulata (Jardine), A. virgata (Sclater), A. maculicauda (Berlepsch), and^4. urubambensis (Chapman). These four forms were considered full species by Hellmayr (1925), Peters (1951 ), and de Schauensee (1966). They all have a diagnostic pattern of streaking — a narrow stripe along the feather shaft contrasting with the remainder of the feather. Within the A . flammulata species group, A . urubambensis can be diagnosed by the combination of a lack of streaking on the dorsum and a lack of rufescent color on the forehead and crown. Further- more, A. urubambensis is sympatric with two of the three other nominal species of the flammulata species group: with A. flammu- lata in Peru (see Zimmer, 1930:345) and with A. maculicauda in Bolivia (see Bond and de Schauensee, 1942:334). There is there- fore no doubt that A. urubambensis is a good biological species. THE ASTHENES FLAMMULATA COMPLEX The other three nominal species of the A. flammulata species group can be described collectively as the A. flammulata complex. They are morphologically rather similar to one another, and are largely allopatric, although there are two apparent exceptions: (1) in southern Peru ("contact" between virgata and maculicauda) and (2) in central Peru (alleged sympatry between flammulata and virgata ) . Sympatry Asthenes virgata was described by Sclater, in 1874, on the basis of one specimen collected by Jelski in "Junin," in the Andes of central Peru. In 1930, M. A. Carriker, Jr., secured five specimens of A. virgata in the Rock Forest area west of Lake Junin (Bond, 1945:33; 1956:245). Although Jelski did not give precise locality data, it is possible that "Junin" refers to the village of that name situated southeast of the lake at the edge of the flat depression form- ing the present-day Lake Junin and adjoining marshes (see Fig. 1 ). Junin and Rock Forest are about 50 km apart. These are the only published localities where A. virgata has been taken, at least to my knowledge. However, C. Koford collected a single female Asthenes in the Department of Puno, southern Peru, in 1951 (MVZ 124302). This bird is morphologically much closer HRI \ lORA No. 297 ^ localities cited in the text " Junin uncertain type loc. of vi rga t a .' A R G K N T I N A # Fig. 1. Collecting localities of the Asthenes flammulata complex 1968 ASTHENES FLAMMULATA SUPERSPECIES 5 to the geographically distant A. virgata from central Peru than to its nearest neighbor, A. maculicauda. As far as I know, this speci- men has not been previously reported in the literature, although it was known to Zimmer, who identified it as A. virgata (pencilled comment on the label) . "Contact" between A. virgata and A. maculicauda appears pos- sible when the collecting localities of Koford's specimen of virgata and of the northernmost record of maculicauda are examined. Ko- ford's bird was collected eight miles SSW of Limbani at 15,250 feet (label data). The northernmost specimen of maculicauda ap- pears to be a bird collected by Carriker at Huancarani, 13,000 feet (ANSP 103442). According to Bond (1945:19), Huancarani is "about ten miles above Limbani." Limbani is a village on the left (west) bank of the valley, and is accessible by only one road or trail from the southwest. Thus it is likely that "above Limbani" and "SSW Limbani" are very close to each other (Fig. 1). However, even if these two localities are not separated by a significant linear distance, they are separated by an altitudinal gap of 2250 feet (about 700 m). There is thus no direct evidence for sympatry be- tween A . virgata and A . maculicauda, but in view of their morpho- logical differences, their sympatry in southern Peru would not be surprising. Sympatry between A . virgata and A . flammulata in central Peru was made apparent by the mention of four specimens of A. flam- mulata taczanowskii collected at Rock Forest by Dorst (1956: 268). But as I shall show below, Dorst's specimens are actually referable to virgata; sympatry is therefore non-existent, since only one taxon occurs at Rock Forest. Geographical Variation Altogether about 140 specimens have been examined from lo- calities covering the entire range of the three nominal species of the A. flammulata complex (Fig. 1). Specimens were grouped into "populations," which are mapped on Figure 2. The grouping was based on several considerations, primarily (a) whether or not cer- tain localities appeared to be isolated from others, thus representing potential interruptions in gene flow, and (b) whether or not the birds from several more or less adjacent localities were morphologi- cally uniform. Thus specimens from the widely-spaced collecting locatities of the Eastern Andes of northern and central Peru were considered as one "population" (H) because of their considerable BRLVIORA No. 297 BOLIVIA r ': 5C*LC ARGENTINA I l II Eat tern Andes I ( : j j .1 m a r c a J (Jum v ilc j. I.ibcrtad K Rock Foren I. ssw Limban i, Puno \l I In. i in j r .1 n i. I'ti n 0 \ La Pa / ( !o< habamba O Aconquija — &INUMOA4. ftoJCCTON Fig. 2. The populations of the Asthenea flammulata complex. 1968 ASTHENES FLAMMULATA SUPERSPECIES 7 morphological uniformity; on the other hand, those from the West- ern Andes of northern Peru were placed in two "populations" (I and J) because of geographical variation in some characters. Despite this grouping into populations, the samples available were small. Under these circumstances, statistical treatment of the mensural characters was not attempted. TABLE 1 Geographical variation of bill length in the Asthenes flammulata complex Populations Range of mean bill length (in mm) Males Females Colombia, Ecuador northern Peru (A-L) 17.0-17.9 16.0-19.0 Southern Peru, Bolivia, northern Argentina (M-O) 15.0-16.5 15.3-15.5 TABLE 2 Geographical variation of tarsus length in the Asthenes flammulata complex Population Tarsus length of males (in mm) Tarsus length of females (in mm) Range (Mean) N Range (Mean) N A 24.0-27.0 (25.3) 3 23.0-24.0 (23.7) 3 B C D 25.0-27.0 (25.9) 11 23.5-26.0 (24.7) 6 24.0-27.0 (25.8) 12 25.0-25.5 (25.1) 4 E 25.0-26.0 (25.6) 9 23.0-26.0 (24.8) 14 F 25.0-25.5 (25.2) 3 24.0 1 G 24.5-26.0 (25.3) 2 24.0-25.0 (24.5) 2 H 23.0-24.0 (23.9) 7 23.0-24.0 (23.8) 4 I 23.0-24.0 (23.0) 2 25.0 1 J 24.0-25.0 (24.3) 3 K 23.0-27.0 (24.6) 7 23.0-24.0 (23.5) 2 L M 23.0 1 26.0 1 N 22.0-24.0 (22.8) 4 22.0-22.0 (22.0) 2 0 22.0-23.5 (22.8) 3 20.0-21.5 (20.8) 2 8 breviora No. 297 Mensural characters. Geographical variation in bill length, tar- sus length, and wing length, respectively, is summarized in Tables 1 and 2 and in Figure 3. Populations M, N, and O (from southern Peru to northern Argentina) are distinguishable from their neigh- boring populations by a gap (step) in either absolute size or in mean length of all three characters. By contrast, steps between any two of the populations from A through L (Colombia to southern Peru) are much less significant (see especially Fig. 3). Population Males (N) Females (N) « so u n is mm mm » '0 mm •— t — i — i — i I I I ) — | I | I | | l i I | l i— t — l — l I l — l t I I I I I I I \ —I- (!) + (31 B 1 — do) —I (i) C + (0 1) 1 (!l) 1 (I) I 1 (•) —I (II) I + (?) (, -4- (?) -f (I) II 1 (I) —I («) I -I- i? i + (') J -HO) + 0) k 1 (1) + (J) I. + (0 \l + (I) N (- («) -HO) () —l hi -I- in Fig. 3. Geographical variation of wing length in the Asthenes flammulata complex. From the variation in mensural characters, one might conclude that the A. flammulata complex consists of two units, each an in- terbreeding species. The first would include populations A through L (nominal species A. flammulata and A. virgata), and the second populations M through O (nominal species A. maculicauda) . Color characters. Geographical variation in the color of the gular patch, the color of the forehead, the amount of streaking on the abdomen and crissum, the color of the rump and upper tail coverts, and the markings of the median pair of rectrices is sum- marized in Table 3. Variation in wing pattern is illustrated in Figure 4. Detailed notes on variation in these characters are in- cluded in Appendix I; only the major trends will be discussed here. 1968 ASTHENES FLAMMULATA SUPERSPECIES CO LU i CO < X "Eh 6 o o « a 53 J3 03 J-i u o i- O o "o o o o a •2 cd •*H c3 > 13 o •-h o a s o *-» ca O, O ft, o < 3 ft< 3 03 00 :> z < 3 ft< 3 pq oo s s < 3 ft< 3 03 oo s hJ O 04 00 3 03 D M o oo 3 03 D e !3 ►-i >H on CO oo 8 c hH I* on 00 oo X 3 ffi >H GO -4-1 00 oo 3 0 O U CO W3 00 s Bh o ft. on ■4-J 00 CO c 13 w o ft. 00 00 00 c Q o ft. oo -4— » 00 00 3 D o o ft. oo 00 OO 3 D oq o 05* 3 pi 00 00 3 < o & CO CO 3 Ul u 4-» o ca u ca 6 o "S "3 D, ea u s '— ,3 3 ca B 00 (U 3 « .<" fcfc! •i-l 3 3 XI o ca Oh >- ja 3 O o en 3 O O ca .3 o O -2 la ft, O ft, oo 3 O OO 5^ CO 3 CO cj CO 0) <+H 3 u T3 ■a CO 3 O ca u 3 (U r o o O T3 CD ca o 3 ft< CO 00 T3 ca CO oo 3 ca E CD S a. a •3 U S3 £ 3 3 D ex. 2 c 10 breviora No. 297 Characters such as the color of the gular patch, the color of the forehead, and the intensity of streaking on the underparts vary rather gradually. There is a trend, from north to south, toward a decrease both in saturation of pigmentation and in the amount of streaking. Exceptions to this trend are provided by populations K and L (gular patch color) and populations M, N, and O (forehead color). The other characters seem to vary more discontinuously. These patterns of discontinuous variation permit one to distinguish mor- phologically several populations or groups of populations within the A. flammulata complex. The best-marked group includes popu- lations from southern Peru to northern Argentina (M, N, O). These birds arc differentiated from their neighboring populations by gular patch color, forehead color, and streaking on the rump. The steps observed ill the variation of these color characters correspond to those noted in the mensural characters. Other groups of populations can also be separated morphologi- cally, but on the basis of fewer characters. Thus the birds of populations A and B (Colombia) can be distinguished by the color of the gular patch and the color of the forehead. Similarly, birds from populations K and L (Rock Forest and SSW Limbani) are separable from others by the color of the gular patch and the gen- eral absence of streaks on the rump and upper tail coverts (if these streaks are present, they are obsolescent). However, it must be pointed out that such a neat separation of populations cannot be done using some other characters. For ex- ample, populations K and L are not distinguishable from popula- tions M, N, and O on the basis of color or streaking of the underparts. Also, populations K and L are indistinguishable from populations H through J in forehead color. When variation in color characters (as in mensural characters) is considered as a whole, the A. flammulata complex appears to comprise two units. Geographical variation within populations A through L, and within M through O, could be interpreted as that normally to be found in an interbeeding species. However, unlike what is found in mensural characters, the gap in color characters separating populations A-L and iM-O does not seem to be as well marked as the gap separating some populations within A-L. Population Structure The dual type of geographical variation (gradual and discon- tinuous) exhibited by the A. flammulata complex is often found in 1968 ASTHENES FLAMMULATA SUPERSPECIES 11 a single polytypic species with a partially continuous and partially discontinuous range. Although the number of specimens examined is not large enough to reach definite conclusions, it seems probable that the gradual size trends (especially in tarsus and wing length) detected are positively correlated with altitude. Thus the largest birds seem to live at slightly higher elevations (in north-central Ecuador and central to southern Peru) than the smallest birds (in southern Ecuador and northern Peru. Altitudinal variation in Andean birds has been demonstrated by Traylor (1950) and myself (unpublished). How- ever, the correlation of increase in size with increase in altitude is not the only possible one. In Peruvian birds, for example, the north-south size increase along the Western Andes could be due to latitude or aridity (Hamilton, 1961), rather than to altitude. The decrease in the intensity of pigmentation from north to south cited earlier is also detectable in several species belonging to other Andean genera (unpublished). Such variation may be correlated positively with the decrease in rainfall along the Andes from north to south (see, e.g., Troll, 1959); thus it corresponds to Gloger's rule. The apparent trends of gradual geographical variation are clear- est in populations A through L, which have the earmarks of repre- senting a single species, as already noted. It is, therefore, of interest that populations M-O, which are separated from populations A-L by a marked morphological gap in some characters, have a very different type of population structure. Although populations M, N, and O are apparently disjunct (see Fig. 2), the birds I have examined are all remarkably uniform and display only slight geo- graphical variation — in wing length, for example (see Fig. 3). This lack of significant geographical variation can be explained either by free gene flow, or by recency of occupation of the vast geographical range, or possibly by both. Whatever the reason, the differences in population structure would tend to confirm the pre- vious impression of specific distinctness between populations A-L (flammulata-virgata) and M-O (maculicauda) . Since, as noted previously, there is no apparent barrier between the two groups of populations in southern Peru, I would conclude that their differen- tiation does not reflect the result of evolution in situ (unless, of course, a barrier once present has subsequently been removed). This question cannot yet be answered, but the problem will be taken up in a later paper in this series. To study the possible correlation between discontinuous variation and geographical barriers in the A. flammulata complex, we must turn to populations A-L, ranging from Colombia to southern Peru. 12 hreviora No. 297 Discontinuous Variation and Geographical Barriers The presence of a geographical barrier between avian popula- tions does not ipso facto mean that gene flow between the isolates is interrupted. Similarly, the fact that geographical isolates are not differentiated morphologically, or are so only slightly, does not necessarily mean that gene flow takes place between them. Conse- quently. Statements made in the following paragraphs about dif- ferentiation and gene Bow across barriers must be taken, not as expositions of known facts, but as cautious expressions of proba- bilities. The correlations between morphological differentiation and geo- graphical barriers in populations A-L of the A. (lammulata complex will be mentioned in geographic sequence from north to south. (1) In Colombia, the birds from the Eastern Andes (multo- striata) are separated from those oi the Central Andes (quindiana) by the Magdalena Valley (Fig. 5). Morphological differences be- tween these two isolated populations (A and B on Fig. 2) include wing length, the width of the streaks on the underparts, the pres- ence or absence oi a slight bulTv wash on the undertail coverts, the extent o\' the gular patch, and the intensity of the coloration of the forehead. The two populations are certainly sufficiently differen- tiated to be recognized taxonomically. although they are more simi- lar to each other than they are (either as a group or separately) to populations farther south in Colombia and Ecuador (C-G). i 2 i Between the Eastern or the Central Andes of Colombia and the Andes of Nariho in southern Colombia is an apparent distribu- tion gap (Fig. 5 ). The only specimen from Nariho (C) I have seen (Mayasquer, ANSP 149893) appears somewhat intermediate in both postocular stripe and cheek color between Colombian birds (A-B) and Ecuadorian specimens (D-G). However, since these differences are slight, it is not possible to assess the real extent of intermediacy. In other characters, furthermore, the Narino speci- men falls fully within the range of variation of Ecuadorian speci- mens. The vast geographical gap between Colombian populations A-B and southern Colombian-Ecuadorian populations C-G is there- fore correlated with a relatively well-marked morphological gap. Taxonomic recognition of populations C-G (subspecies flammu- lata) is thus warranted. (3) The Interandean Depression of Ecuador (Fig. 5) is a low and relatively dry region separating the Eastern from the Western Andes. Only a slight difference in wing length and tarsus length 1968 ASTHENES FLAMMULATA SUPERSPECIES 13 between birds of the Eastern (D) and Western (E) Andes was de- tected, suggesting only a slight restriction in gene flow; taxonomic subdivision within populations C-G (fiammulata) would be un- justified. (4) There is a gap between the northern Peruvian populations of the Eastern (H: taczanowskii) and Western (I: pallida) Andes and those of southern Ecuador (G). For population H, the gap corresponds to the eastward bend of the Maranon River; for popu- lation I it corresponds to the generally low altitude and small area of suitable habitat of the Andes of northwestern Peru (Northern Peruvian Low: Fig. 5). The morphological differences between Ecuadorian and Peruvian birds are not large. They include the in- tensity of streaking on the underparts and the extent of rufous on tail feathers. The lack of major differences suggests either some gene flow over the gap, or that the separation of Ecuadorian from Peruvian birds is of recent origin. (5) The Upper Maranon Basin (Fig. 5) separates the northern Peruvian populations of the Eastern Andes (H: taczanowskii) from those of the Western Andes (I and J: pallida). Only slight morphological differences can be noted between them, however: these include characters such as tarsus length, wing length, and intensity of streaking of the rump and upper tail coverts. I could find no differences in the other characters examined. In view of this minor morphological differentiation, subspecific separation of the two populations may not be warranted; pallida may be consid- ered a synonym of taczanowskii. As Bond (1945:32) has re- marked, pallida "is not a very well-marked race." (6) The specimens collected in the Rock Forest area of central Peru (K: virgata) are distinguishable morphologically from the birds living in both the Eastern (H) and Western (I, J) Andes. The differences are in wing length, color of gular patch, color of central pair of rectrices, amount of streaking on rump and upper tail coverts, wing pattern, and color of the lower abdomen and crissum. These differences are better marked between Rock Forest specimens (K) and birds from the Eastern Andes (H), than be- tween Rock Forest birds and specimens from the Western Andes (I, J) . An approach to Rock Forest birds is clearly seen in Western Andean birds (especially population J) in size (Fig. 3) and in wing pattern (compare nos. 5 and 6 on Fig. 4). Geographical isolation between Rock Forest birds {virgata) and Eastern Andean birds {taczanowskii) is apparently provided by the Lake Junin Basin (Fig. 5). But since the first specimen of virgata may have come from "Junin" in the center of the basin, a more 14 BREVIORA No. 297 ® © Fig. 4. Geographical variation of wing pattern in the Asthenes flammu- lata complex. The feather illustrated is the sixth outer primary. Key: 1 = Volcan Pichincha, Ecuador; 2 = Huanuco, northern Peru; 3 = Maraynioc, central Peru; 4 and 5 = Quirivilca, Libertad, northern Peru; 6 = Rock Forest, central Peru; 7 — SSW Limbani, southern Peru; 8 and 9 = Incachaca, Cochabamba. Bolivia 1968 ASTHENES FLAMMULATA SUPERSPECIES 15 likely barrier may be the high Andes farther east. In any event, although the barrier is not clear-cut at present, morphological dif- ferentiation is fairly sharp. Isolation between Rock Forest birds (virgata) and populations found farther north along the Western Andes (J: "pallida") is not clear. The closest specimens come from Quirivilca, Libertad, about 375 km from Rock Forest. No specimens are known to me from the intervening area, although apparently suitable habitats occur there. A possible barrier might be the Santa River Canyon, which makes a cut across the Andes between the Cordillera Blanca and the Andes of Libertad. (7) The vast Andean area between central and southern Peru (about 800 km) seems to represent a distribution gap in the range of the A. flammulata complex. However, suitable habitat occurs there, and further collecting might reveal the presence of the com- plex there (? on Fig. 5). The specimen from SSW of Limbani is practically identical with Rock Forest specimens {virgata), except in the pattern of the central pair of rectrices. Taxonomic Conclusions The classification of Hellmayr (1925), Peters (1951), and de Schauensee (1966) is as follows. Colombian, Ecuadorian, and northern Peruvian populations (A-J) are treated as one species, Asthenes flammulata. The two disjunct populations from central and southern Peru (K and L) would constitute a second species, A . virgata. Finally, the populations from southern Peru to northern Argentina (M, N, O) make up a third species, A. maculicauda. A slight modification of this classification was made by Bond and de Schauensee (1942:334), who considered maculicauda a subspecies of flammulata. (Later, however, Bond, 1945:32, cited maculi- cauda as a full species without any comment.) This classification finds justification in the fact that there is a morphological and partial geographical gap between flammulata and virgata (see (6) above), and because there is a morphological gap between flammu- lata-virgata and maculicauda. I believe, however, that the evidence presented in this paper allows one to construct a classification of the A. flammulata com- plex which is at the same time a simplification and an improvement over the older scheme. The Asthenes flammulata complex constitutes a superspecies (sensu Mayr, 1963:501) having two component species. The first 16 BREVIORA No. 297 Northern Peruvian Low c Upper Mar a Eon Valley ARI \ WHERE \ Lake Jum'n Has INTER- V?, GR tDATION ? £ BETW I I N FLAMMULA1 \ WD VIRGA1 \ IS POSSIBLE PROBABLE CONTACT BET w I I N VIRGATA : ///. AM) MAC I 1. 1 CALL) A A t I j ni in ii I i i i A m j i u I u j u d .i Fig. 5. Taxonomic and evolutionary interpretation of distribution and geographic variation in the Asthenes flammulata superspecies. 1968 ASTHENES FLAMMULATA SUPERSPECIES 17 species is composed of several isolated and semi-isolated popula- tions ranging from Colombia to southern Peru (Asthenes flammu- lata). Fully isolated populations of A. flammulata which have reached a moderate to marked degree of differentiation (incipient species) are the Colombian multostriata and quindiana. These two isolates are perhaps best considered as a subspecies group with two subspecies. The specimens from Rock Forest and SSW Limbani fall within the range of variation of a single taxon (virgata).1 Some gene exchange between Rock Forest birds and birds farther north in the Western Peruvian Andes is suggested by the variation of characters such as wing length (Fig. 3). I consequently recom- mend that virgata be treated as a subspecies of A . flammulata. The second component species of the A. flammulata super- species, Asthenes maculicauda, is composed of three apparent iso- lates (M, N, O), but it does not exhibit much geographical varia- tion. The taxa of the A. flammulata superspecies and their relation- ships can be listed as follows: Asthenes flammulata superspecies (two component species which may have achieved reproductive isolation) ( 1 ) Asthenes flammulata ( Jardine, 1 850) (a) Colombian subspecies group (one or two incipient species): A. flammulata multostriata (Sclater, 1858), A. flammulata quindiana (Chapman, 1915) (b) Ecuadorian-Peruvian subspecies group (two or three incipient species): A. flammulata flammu- lata (Jardine, 1850); ,4. flammulata taczanowskii (Berlepsch and Stolzmann, 1894; includes A. f. pallida Carriker, 1933); A. flammulata virgata (Sclater, 1874) (2) Asthenes maculicauda (Berlepsch, 1901) Shows insignificant geographical variation SUMMARY The Andean spinetails of the Asthenes flammulata complex in- clude three nominal species, flammulata, virgata, and maculicauda, which, together with a fourth, A. urubambensis, constitute the A. flammulata species group. 1 The specimens reported by Dorst (1956) as A. flammulata taczanowskii from Rock Forest are therefore referable to virgata. 18 brlviora No. 297 Specific status of A. urubambensis is shown by sympatry with two nominal species of the A. fiammulata complex and bv morpho- logical distinctness. The evolutionary relationships of the three nominal species of the A. fiammulata complex were analyzed by a study of geographical \ariation and its correlation with geographical barriers. Geographi- cal variation is largely discontinuous in populations ranging from Colombia to southern Peru (nominal species fiammulata and vir- gata). By contrast, populations from southern Peru to northern Argentina show great morphological uniformity (nominal species maculicauda ) . It is concluded that the Asthenes fiammulata complex is best con- sidered as a superspecies with two component species. The first species, A. fiammulata, contains several isolated groups of popula- tions (incipient species) showing various degrees of morphological differentiation across barriers. A. virgata should be included as a subspecies of A. fiammulata. The second species, A. maculicauda, may come in contact with A. fiammulata virgata in southern Peru. Gene exchange between the two species i^ not suggested, but there- are only two specimens mi which to base a decision. REFERENCES CITED Bond, J. 1945. Notes on Peruvian Fumariidae. Proc. Acad. Nat. Sci. Philadel- phia, 97: 17-39. 1956. Additional notes on Peruvian birds II. Proc. Acad. Nat. Sci. Philadelphia. 108: 227-247. Bond, J., and r. m. df. Schalensee 1942. The birds of Bolivia. Part I Proc. Acad. Nat Sci. Philadelphia. 94: 307-391. Carriker, M. A., Jr. 1933. Descriptions of new birds from Peru with notes on other little known species. Proc. Acad. Nat. Sci. Philadelphia, 83: 1-38. DORST, J. 1956. Etude d*une collection d'oiseaux rapportee du Perou central. Bull. Mus. Paris, 2e Ser., 28: 265-272. Hamilton, t. h. 1961. The adapti\c significance of intraspecific trends of variation in wing length and body size among bird species. Evolution, 15: 180-195. Hellmayr, C. E. 1925. Catalogue of birds of the Americas. Part IV. Furnariidae- Dendrocolaptidae. Field Mus. Nat. Hist. Zool. Ser., 13: 1-390. 1968 ASTHENES FLAMMULATA SUPERSPECIES 19 MAYR, E. 1959. Trends in avian systematics. Ibis, 101: 293-302. 1963. Animal Species and Evolution. Harvard Univ. Press, Cam- bridge. Peters, J. L. 1951. Check-list of Birds of the World. Vol. VII. Mus. Comp. Zool., Cambridge. SCHAUENSEE, R. M. DE 1966. The Species of Birds of South America and their Distribution. Livingston Publ. Co., Narberth, Pennsylvania. SCLATER, P. L. 1874. Descriptions of three new species of the genus Synallaxis. Proc. Zool. Soc. London, 1874: 445-447. Traylor, M. A. 1950. Altitudinal variation in Bolivian birds. Condor, 52: 123-126. Troll, C. 1959. Die tropischen Gebirge. Bonner Geogr. Abhandl., Heft 25: 1-93. ZlMMER, J. T. 1930. Birds of the Marshall Field Peruvian expedition, 1922-1923. Field Mus. Nat. Hist., Zool. Ser., 17: 233-480. (Received 23 November 1967.) APPENDIX I: NOTES ON GEOGRAPHICAL VARIATION IN NON- MENSURAL CHARACTERS Gular patch. The gular patch varies in color intensity from whitish and yellowish to rich ochraceous, and in extent from a small, roughly tri- angular patch more or less restricted to the chin, to a wider patch com- prising also the throat and upper chest. Colombian specimens (populations A and B) have a richly ochraceous gular patch. Central Andean birds (B) have a rather broad patch, while in Eastern Andean birds (A) it is more restricted. Ecuadorian specimens and the Nariho specimen from Mayasquer (popu- lations C-G) have relatively narrow and more or less triangular gular patches, varying from whitish-yellow to pale ochraceous. Peruvian specimens (populations H-J) have a broad, ill-defined gular patch merging into the upper chest. The patch varies in color from dirty white to yellowish-cream. Usually the chin area is somewhat paler than the throat. Some specimens of Ecuadorian birds (e.g. MCZ 199104, female, and MCZ 199099, male) have a more ill-defined gular patch than do most other Ecuadorian specimens, thus showing an approach to Peruvian specimens. Peruvian specimens from Rock Forest and SSW Limbani (K and L) have a relatively small, well-defined gular patch, usually of a rich ochraceous color. 20 BREVIORA No. 297 Specimens from southern Peru to northern Argentina (M, N, O) lack a buffy-ochraceous gular patch: the gular area is of about the same color as the chest. Forehead color. In Colombian specimens from both the Eastern and Central Andes i A and B) the forehead is an almost solid rufous, especially in males. The forehead of most Ecuadorian specimens (C-G) is streaked with rufescent, although in some birds (e.g. MCZ 138553) it is a solid rufous. In Peruvian birds from populations H, 1. and J the forehead streaks are less rufescent. more yellowish, but here again at least one specimen i I MNH 65849, male) has a rufous forehead, where the streaks almost join to form a solid color. In Peruvian specimens from Rock Forest and SSW LimbanJ (K-L) the streaks are relative!) yellowish, not verv rufescent. Finally, specimens from southern Peru, Bolivia, and northern Argentina (M, N. O) have rufescent foreheads, distinctly more pigmented than those of Peruvian birds in general, and of Rock Forest-SSW I imbani birds in particular. Coloration of under parts. Colombian specimens are the most beavil) streaked; Bolivian and Argentine specimens are almost unstreaked. The birds from Rock forest. SSW 1 imbani. Huanearani. Bolivia, and northern Argentina (k-Ot are the least streaked ol all populations within the Asthenes ftammulata complex I be additional fact that in the birds from these same populations the lower abdomen and crissum are buffy with obsolescent streaks, rather than whitish with rather conspicuous streaks, permits one to distinguish them immediately from all other populations (A-J). There is. in particular, a rather well marked difference in streaking intensity between the crissum of Peruvian birds from the I astern (H) and Western Andes i I. J t. and that of birds from Rock Forest iki. Specimens fiom southern Peru to northern Argentina (M, N, O) have, alone among the populations of the A. tlammuhitu complex, a series of ill-marked streaks on the chest converging toward the center to form a ring. Color of rump and upper rail coverts. I he birds from Rock Forest and ssW Limbani (K and 1.) have slight or obsolescent streaking on the rump .mA upper tail coverts. (This was already noticed by Bond [1956: 245] for Rock Forest specimens.) By contrast, specimens from all other populations have generally much heavier streaking on the rump and upper tail coverts. These streaks are relatively well marked in Peruvian populations H, [, and J, as well as in Peruvian, Bolivian, and Argentine populations M, N, and O. Rock Forest and SSW Limbani ( K.-E ) specimens are thus distinguishable as a group from neighboring populations. Wing pattern. The primaries of birds in all populations are more or less rufous with a dark brown to blackish tip. I have studied interpopulation variation in pattern by comparing the patterns of the sixth outer primary. which was chosen arbitrarily. Except for a few Colombian specimens, which have only a relatively narrow rufous edge to both vanes of the sixth primary, in most other speci- mens (including those from Colombia) the tip of the feather is more or 1968 ASTHENES FLAMMULATA SUPERSPECIES 21 less wholly dark brown, while its base is rufous (see Fig. 4, Nos. 1, 2, 3, 4, and 9). The most significant population variation is in the presence or absence of a thin dark brown stripe along the outer vane of the primary (com- pare Nos. 4 and 5 of Fig. 4, for example). This pattern is seen in some specimens from Quirivilca, Libertad (population J), every specimen from Rock Forest (K), the single specimen from SSW Limbani (L), and most specimens from southern Peru to northern Argentina (M, N, O). The presence of the two major patterns in the Libertad and Cochabamba popu- lations is illustrated in Fig. 4 (Nos. 4 and 5, and Nos. 8 and 9, respectively). Marking of rectrices. Analysis of variation in this character is difficult because a rather large number of specimens have one or several rectrices in molt or missing and because many specimens have extremely worn rectrices. Nevertheless, the detectable variation appears as follows. There is a tendency for Peruvian specimens (including the specimen from SSW Limbani, but excluding Rock Forest specimens) to have more rufous on outer tail feathers than do Ecuadorian or Colombian specimens. Rock Forest specimens lack entirely or almost entirely the narrow rufescent or pale brownish outer edge to the outer vane of the central pair of rectrices, which is present in specimens from all other Peruvian localities including SSW Limbani. In birds from southern Peru to northern Argentina (M, N, O), the central pair of rectrices is marked with irregular dark brown designs on a rufous or rufescent background. No other specimen from anywhere in the range of the Asthenes flammulata complex has similar markings on the median rectrices, with one exception: in the central pair of rectrices of the specimen from SSW Limbani, the outer part of the outer vane is rufescent, with a thin and irregular dark brown line running alongside the rufescent color. This pattern, although reminiscent of that found in birds from populations M, N, and O, is not really similar to it. In rectrices other than the central pair, a pattern reminiscent of the one found in specimens from southern Peru to northern Argentina (M, N, O) is noticeable in specimens from Rock Forest (ANSP 92428, ANSP 92427, MP CG1956 No. 942, MP CG1956 No. 940), as well as in other populations. But the number of specimens in which this pattern occurs tends to become smaller north of central Peru. BREVIORA Museum of Conn para tive Zoology Cambridge, Mass. 11 October, 1968 Number 298 MONOGRAPH OF THE CUBAN GENUS VIANA (MOLLUSCA: ARCHAEOGASTROPODA: HELICINIDAE) William J. Clench and Morris K. Jacobson Abstract. The genus Viana in the family Helicinidae is limited in its distribution to the Sierra de los Organos in Pinar del Rio, the westernmost province of Cuba. All members are found only on limestone substrate. The genus consists of one species, regina Morelet, with two subspecies, laevigata Pfeiffer and subunguiculata Poey, that form many races which are not clearly discrete morphologically and which show no coherent geographical distribution. INTRODUCTION This study of the genus Viana is based upon specimens housed in the Museum of Comparative Zoology which include the very large collections made by Carlos de la Torre, Pedro Bermudez and Julio Armenteros. Much additional material was contributed by Thomas Barbour, William J. Clench, Paul Bartsch, Carlos G. Aguayo, Miguel L. Jaume, Morris K. Jacobson, and numerous other Cuban and American collectors. The strikingly handsome shells of the helicinid genus Viana are confined to the Sierra de los Organos, Pinar del Rio Province, in the western end of Cuba. Their range is limited in the west by the sierras and mogotes1 about Mendoza, and in the east by the Sierra Rangel, about 75 miles away. The species of Viana are purely calciphilous, living exclusively on the limestone mogotes and sierras which emerge from the surrounding red, iron-bearing soil. In such locations during the rainy season they can be collected as readily as littorinids on rocky sea coasts. Unlike the related genus Emoda, they are found in no other part of the island. Within their range they inhabit practically every outcrop of limestone of any size which enjoys sufficient moisture and shade. 1 In western Cuba a mogote is any isolated limestone block separated from the main sierra. HRI-VIORA NO. 298 Thus they are common on high sierras like those of Guane, Paso Real, Vinales, San Carlos, San Andres, Guajaibon, and Rangel, and on the small mogotes in the \alle\ oi \ ihales. the Laguna de la Piedra, and around Luis La/o. Isabel Maria, and between the Estrechura of San Carlos and Guane ( Teneria. Punta de la Sierra. 1 .1 Muralla, and Los Portales). Map I Showing the greater part of the Province of Pinar del Rio, Cuba, outlining the areas containing ihe sierras and mogotes which are occupied In the three subspecies of Viana regina Morclet. A. liana retina .uilutnguiculata (Poev). B, V. r. laevigata (Pfeiffer). C. V. r. regina (Morelet). (hies and towns Sierras and Mogotes 1. Guanaja) X. Sierra Rangel 2. Artemisa 9. Pan de c ruajaibon 3. ( onsolacion del Norte 10. Sierra San Andres 4. \ ihales 1 1. Sic i ra \ ihales 5. Consolacion del Sur 12. Mogotes at "Km. 14" ft. Pinar del Rio 13. Ski ra del Quemado 7. Guane 14. Sierra Paso Real The genus Viana appears in a series of differentiated forms. These vary in sculpture from smooth and shining to dull and spi- rally grooved; in color from vcllow to reddish and purple, orna- mented at times with red. yellow, or diaphanous bands, or with CUBAN GENUS VIANA whitish mottling; and in size from the large forms of Sierra Rangel, (29.5 mm in width) to the small forms (18.2 mm in width) which inhabit small mogotes such as those at Kilometer 14, Vinales. In most cases, specimens of Viana collected on small mogotes are dwarf forms about one-half the size of typical specimens. It is apparent that the size of individual specimens is a function of the area of the habitat and may very well be of selective value for survival, based upon the supply of food. In addition to other variations, the margin of the peristome may be simple and unre- flected as in V. regina regina and V. regina laevigata, or more or less strongly reflected as in V. regina subunguiculata. From the scant field notes we assembled we find that vianas are largely satisfied with the presence of limestone, shade, and mois- ture. We have never collected a live specimen other than on lime- stone rock. Nothing has been published about their food, but they probably eat the lichens found on the rocks on which they live. They have an enemy in the larval form of the large Cuban fire- fly, Alecton sp.; pictures have been taken of such a larva devouring a Viana through the aperture (PI. 1). Probably the firefly larva PLATE 1 Viana regina (Morelet) Viana regina Morelet being attacked by a lampyrid beetle larva (Alecton sp.). Photo by F. de Zayas (about 6 X). 4 BREVIORA No. 2C>K attacks its pre) when the snail is relaxed. The predator may wait for the Viana to relax the operculum and then attack, much like the marine predators of the genus Melongena, among others. Baker described the radula of Viana ( ll)22) and its anatoim (1926). We could find nothing in print regarding its natural history. The following notes are translated from a personal com- munication from Oscar Alcalde l.edon who. until he left the countr) in 1963 was in charge of the Division oi Malacology in the r.cwk formed Acadcnna de Ciencias de la Rcpublica de Cuba. Alcalde tells us that he unfortunately left his notes behind and writes from recollection only. I I he eggs] are subglobular in shape, the size of a capital letter ( )' m normal typewriter print, or perhaps a hit larger. I found them under a slab of roek ami the) were covered with the reddish soil on which the rock rested Vfter being washed, the) proved to be wax colored [whitish]. I did not know at tit si thai the) were Viana eggs, but later upon breaking them. I found the ncpionic whorls of a snail which under microscopic examination proved to be iden'ical to the nepionic whorls of Viana. The operculum was ver) small and thin."' According to Thiele ( 1929:81 ). all helicinids lack a male copu- lator) organ, and the manner of transfer of sperm from male to female is unknovt n. Several of the forms of Viana mentioned above were given fames b\ the late Dr. Carlos de hi I orre. but almost all remain in manuscript. Under these manuscript names the) have been widck distributed b\ Cuban collectors, and one of the purposes of the present Stud) was to examine the validity i^i these forms. After carefull) localizing a large number of Viana specimens on an en- larged map of the Sierra de los O.ganos region, we find that we are amply justified in rejecting these names in i<>to. The confusion in this genus is very much like that of Polymita and Liguus. That is. unless the localities of the various forms are carefull) plotted, the worker would find it easy to attach names to the many color forms and forms with only minor morphological differences. A review of the literature covering Polymita and / iguiu has demonstrated amply the confusion which results from such a proliferation of names. We choose to follow a more con- servative course and to refer to striking varieties, which blend into one another, bv vernacular names such as marbled, liratc. smooth, banded, roseate, yellow, dwarf, shining, dull. etc.. instead of using Latin infraspecific taxa which might be misunderstood as tax- onomic designations. CUBAN GENUS VIANA 5 In doing so we follow the procedure generally adopted for such a variable species as the intertidal marine mollusk from the north- east coast of North America, Thais lapillus (Linnaeus). The analogy between the two groups mentioned is quite close, since in both cases we are dealing with prosobranch mollusks living in more or less isolated colonies on rocks, and hence displaying a rich variety of sculpture, size, and color. The common practice today is to designate all the lapillus varieties under the simple name Thais lapillus. We propose to do the same for Viana regina (Morelet). The distribution of the various populations as revealed on our maps nevertheless seems to justify the recognition of sub- unguiculata Poey and laevigata Pfeiffer as valid subspecies. V. subunguiculata extends southwestward from a line running between Sierra del Quemado Sierra Vihales (with a very strong salient reaching into the Sierra Celadas branch of the Sierra Vinales complex) and Sierra Cabezas. Southwestward from this line, sub- unguiculata is found reaching to the extreme western limit of the distribution of the genus Viana at Sierra de Guane and Sierra Paso Real near Guane. This range includes the sierras around Luis Lazo (San Carlos, Los Acostas), at Isabel Maria and Cabezas, and at the isolated mogotes between Luis Lazo and Guane. The subspecies laevigata is centered about the complex of mountain ranges which go under the name of Sierra de Vihales,1 with disconnected outrunners at the Cayos de San Felipe, La- gunillas, Matahambre, Pan de Azucar, Cabezas, Isabel Maria, and some colonies on the Sierra del Quemado. The line separating laevi- gata from regina, sensu stricto, appears to run south between the Banos de San Vicente on the west and the Sierra San Andres (with La Jagua) to the east. East of this line, and reaching to the eastern limit of the range of the genus at Rangel, the nominate subspecies regina regina can be found. The mogotes at Kilometer 1 4 and El Guama also are inhabited by regina regina, as are many mogotes in the Vinales valley (Dos Hermanos, Mogote La Vega, Hoyo de los Santos, Mogote de los Muertos), whereas others in the same area (Mogote Pequeno, Mogote Alto, Hato Morales, El Cuajani) have populations of laevigata. We realize, of course, that certain populations are sufficiently distinct to enable the student to recognize the locality from which 1 Sierra del Infierno, Sierra Penitencia, Sierra Pan de Azucar, Sierra Martillo, and others. 6 breviora No. 298 they come. Nevertheless, there arc many specimens for which this is not possible, nor did we find it easy to establish fixed dividing lines between the various characteristics, with the exceptions noted below. Throughout the range of the genus there is much dupli- cation o^ the characteristics upon which de la Torre based his manuscript names. This separation of the genus into three subspecies and numer- ous, if not clearly defined, races presupposes an early continuous limestone area which, bv subsequent lifting and corrosion, was broken up into a series of discontinuous ranges (sierras) and iso- lated blocks (mogotes). There is a small amount of territory at the southwestern end of Sierra de Vihalcs and at Sierra del Oue- mado where laevigata and mbunguiculata overlap with very little fusion of characters. However, mure collecting is needed in these two areas. I his is also true of the small territorial overlap in Vihalcs at El Ouecjue between laevigata and regina. Mechanical dispersal may be responsible for this overlap. Gene How. how- ever, between members of the same subspecies must be vary rare, particularly between isolated populations, such as those oi Tan de Guajaibon and Sierra Rangel. Hurricanes could be the means of such mechanical dispersal, carrying specimens or their eggs. Writing about a large mogote near Vihalcs. called previously T.I I umbadero but now El Quequc. Henderson ( 1916:234) states: "The rock is discolored and considerabl) altered bj metamorphic process, it is |um possible thai the limestone of this southern range of the Organos Mountains is ;i shade older than that of the northern system (the ( ostanera), hut the folding upward of the two was cer- tainl) contemporaneous. All the mogotes of the Viriales valley, in- eluding the large Tumbadcro. the Dos Hermanos. and the many smaller ones scattered about the valle\ floor are clearly but fragments of the main sierra now wholly detached and isolated by erosion [corrosion]. The ammonites, which Dr. Torre has been collecting for several /ears, come principally from the rocks of this southern range and indicate probably Jurassic origin of the limestone.1 As hereto- fore noted, the mogotes about Kilometer 14' do not belong to the Vinales range, but together with the Cerro de Cabras hills farther west, represent either remnants of a third line of Jurassic sierra that once followed a course parallel with the existing ranges, or possiblv a later Tertiary deposit upon the flanks of the older mountains." 'According to We\l (1966: 42-47), the Vinales limestone (Kalk) is of Upper Jurassic age and has come by three overthrusts from the north. CUBAN GENUS VIANA 7 Some of the localities cited here must be taken provisionally. There are no reliable detailed maps of the Sierra de los Organos region. Even the map composed by the Cuban army engineers in 1933-1941 is too general, not always accurate, and in several in- stances illegible. Coupled with this difficulty is the haphazard way in which the Cuban peasant assigns names to the various features of his local geography. Many mogotes, cerros, vegas, hoyos, and arroyos have the same name, but on the other hand, many of these features have several. Frequently the name of a mogote is that of the farmer near or on whose land it stands; hence, when this land changes hands, these names frequently change. There is no certainty that such names were ever officially recognized or published, or even that they appear on local land records. A final difficulty lies in the matter of spelling. The unique Cuban pro- nunciation of Castillian is reflected at times in the orthography. Thus, ayua, the local name of a tree {F agar a sp.), which is also associated with the name of a hill and cove in the Sierra San Andres, appears as ayuda, allua and alluda on field labels. In one instance a mogote called muela de la vieja (Old Woman's Tooth) also appears on a collecting label as colmillo de la vieja (Old Woman's Fang). The most useful local maps where those published by de la Torre and Bartsch (1938) and field maps prepared by Oscar Alcalde Ledon and Julio Armenteros. The gazetteer on Cuba published by the United States Board on Geographic Names (1957) was also very useful. In some cases, however, we have had to be satisfied with reporting localities as they appear on the collection labels. We do so in the hope that local inquiry at a later date will enable the collector to find these localities. ACKNOWLEDGMENTS We are deeply indebted to Ruth D. Turner for the dissection of the soft anatomy and of the radula, and to Kenneth J. Boss and Ruth D. Turner for having read the manuscript and having offered many valuable suggestions. This work was supported largely by National Science Foun- dation grant GB-1004. SYSTEMATIC DESCRIPTION Genus Viana H. and A. Adams Viana H. and A. Adams, March 1854, The Genera of Recent Mollusca, 2: 305 [as subgenus of Trochatella Swainson] (type-species, Helicina regina Morelet 1849, subsequent designation, Pilsbry and Brown 1910, 8 BREVIORA No. 2l)S Proc. Acad. Nat. Sci. Philadelphia, f>2 525); non Walker 1869 [Lepi- doptera] Hd/uini Gray, November 1856, Ann. Mag. Nat. Hist., (2) IK: 414 (type- species, Trochatella regina (Pfeiffer) [sic, Morelet], original designa- tion ). Rhynchocheila Shuttleworth IS77. Notitiae Malacologicae, 2: 15 (type- species, Helicina regina Morelet 1849, monotypic). Fitzia Guppj 1895, Proc. Victoria Ihm rrinidad, I. pt. 2: 74 (type-species, Helicina regina Morelel 1849, monotypic). Description. Shell large, reaching 29 mm in diameter, imper- forate, trochiform, generally solid, taint 1 \ to rather strongl) sculp- tured, and lacking periostracum in adult specimens. Slid Is uni- eolored. or ornamented with bands of varying width, or with a marbled or mottled design; a thin reddish subsutural line is present in some specimens. I ip simple and occasional!) sinuous or Baring, male shell with a notch mid\va\ between the outer lip and its insertion in the parietal area. ProtOCOnch \Vl whorls, smooth or irregularl) pitted, sharpl) raised above the succeeding whorls as in Eutrochatella, generall) lighter in color than the rest of the shell except the base Operculum calcareous, auriculate, white externall) with a thin, light brown, internal corneous layer and darker at the periphery. I he outer margin is widel) rounded, the columellar margin con- cave above, generall) straighl and descending almost vertical!) below. It is rounded at the base, generall) on a single plane, but sharpl) raised basall) and less so along the- columellar margin. I here is a long, rounded, low. more or less vertical protuberance along the inner margin, thickest centrally. Outer margin very slightK thickened; a thin, opaque, transverse area reaches internally across the mid-line. Nucleus lateral and central near the columellar margin, with concentric lines of growth which follow the outline o\ the shell. Remarks, The notch on the peristome of the male, which we hereafter call the peristome notch, was one cause o\' the repeated renaming which this genus underwent. Morelet ( 1849: 19) writes "Peristome supero. . . profunde emarginato." Gray ( 1856: 414) likewise thought the notch was characteristic of the entire group and created his genus Hapata. H. and A. Adams (1854: 305) also thought this to be true of their genus Viana. Fischer (1885: 796), citing Pfeiffer, says "les coquillcs d'une mcnic especc ont leur labre tantot echancre. tantot a peine sinueux et cette difference serait en rapport avec la sexe." Arango (1878: 41 ) took the notched shells to be females. Wagner ( 1908: 129) correct!) stated that it was the male which possessed the notch, but then changed his opinion (1910: 1X5). He was corrected by CUBAN GENUS VIANA 9 Baker (1926: 52), and the dissections performed for this study by Turner l support Baker's contention that the notched shells are male. Athough Thiele (1929: 83), correctly describes the situa- tion, Wenz (1938: 438) and Keen (1960: 288) state that the entire genus is characterized by the notch. Wagner (1910: 184) noted other features of sexual dimorphism in the shell which our examinations did not seem to justify. Baker (1922: 62) considered Ampullina De Blainville a syn- onym of Eutrochatella sensu stricto, a Jamaican group. However, the only helicinids of Jamaica that compare in size with De Blain- ville's figure of the type, A. striata (1827: pi. 35, fig. 4a-c), are Eutrochatella tankervillei (Gray) and Alcadia major (Gray). But neither the description (1824: 441) nor the rather poor figure indicate either of these. There is no mention of the strong sculp- ture of tankervillei, and the aperture of Alcadia major is quite different. Ampullina striata De Blainville (1824: 567) (locality un- known) has been considered to be in the genus Viana, but a re-ex- amination of the figure published by De Blainville in his Manuel de Malacologie (1827, pi. 35, fig. 4a-c) indicates that it is not a member of this genus. The operculum figured (4b-c) is quite dif- ferent from that of Viana. De Blainville shows a strong ridge on the inner face from the center margin at the parietal area and extend- ing horizontally to the opposite margin. This morphological struc- ture does not appear in any of several hundred opercula examined in the three subspecies of Viana. The whereabouts of the figured specimen (type) is unknown to us. The range of Viana extends in a broad arc from the Sierra Rangel near San Cristobal, west and south to the immediate vicin- ity of Guane, all within the Province of Pinar del Rio, Cuba. Viana regina regina (Morelet) Plate 2, figures 1 -4 Helicina virginea d'Orbigny 1842 [in] Sagra, Hist. Tile de Cuba, Mol- lusques, 1: 241 (Interieur de l'lle de Cuba); non Lea 1834. Helicina regina Morelet 1849, Testacea Novissima Insulae Cubanae et Americae Centralis, Paris, 1: 19 (Ad Montes Guajaibon, Insulae Cuba). [Paratypes, MCZ 146706, ex Putzey's collection.] Helicina maculosa 'Newcomb' Jay 1850, A Catalogue of the Shells Con- tained in the Collection of John C Jay, 4th ed., New York, p. 261 [nomen nudum]. 1 Dr. Ruth D. Turner is in the process of preparing a detailed anatomical investigation of Viana. which will appear later in this series of reports. 10 BREVIORA No. 298 Helicina multistriata 'Velasquez' J;i\ 1850 ibid., p. 2M [nomen nudum]. Eutrochatella {Hapatd) regina Morelet. Wagner l('i!S, [in] Conchylien- ( abinet, (2) I. Sect. is. pt. 2: 130, pi. 25, figs. 12. 13, IS. 19. Viana regina (Morelet). Baker ll>22. Proc. ^cad Nat. Sci. Philadelphia, 74: 63, pi. 7, figs. 34-36. liana regina marmorata (lone MS) Webb I'MS. Foreign I and and Fresh Water shelN. St. Petersburg, Florida, p. 143. pi. 65, fig. 6. Description. Shell moderate!) large, turbinate, sculptured with spiral cords oi varying degrees of strength, from deeply lirate to cjuite smooth, crossed by weak, oblique, irregular growth lines. Color basicalK yellow, whitish, or light reddish purple, at times with faintly outlined spiral bands. Shell with some degree o'i mot- tling, occasionalK covering the entire surface or else being confined to the spue. When such a shell is held to the light, the whitish or yellowish markings are seen to be more opaque than the base color o\' the shell, and the) apparent!) constitute an added layer o\' shell matter on the surface. I hese markings, however, are SO closel) bound to the rest o\ the shell that they cannot be removed even by the most diligent scraping. Whorls 6V£, moderately inflated, regu- larl\ widening; bod\ whorl rounded. Aperture widely lunate, white near the edge, yellowish or light reddish purple internally. Outer lip entire, not reflected and not flaring, except vcr\ weakly near the insertion with the columella, and deflected downward at the base ol the aperture. Peristome notch rather deep. Y-shaped, narrowly rounded at the base. Parietal lip smooth, without folds, gcntl\ convex above, sharply and almost verticall) descending below. barel) rounded at the juncture with the basal part of the lip. Colu- mella without a fold but with a low. oblique, rounded and slightly raised margin at the umbilical area, appearing as a slight tubercle on the outer basal edge. Shell imperforate, but with a shallow excavation just above the base o\ the columella. Sutures moder- ately impressed, faintly and irregularis scalloped by the edges of the growth lines. Operculum as in the generic description. Pan de (iuajaibon. Syntypc. Sierra Rangel. Pico Grande. Sierra San Andres. Sierra de Guacama\a. San Andres. Kilometer 14, Vinales. Height Width mm mm 20.5 23.0 9 24.5 29.5 6 21.5 23.5 i 23.0 25.0 9 15.5 IS. 2 6 15.2 17.5 S 16.5 18.5 9 CUBAN GENUS VIANA 1 1 Remarks. There are many intergrading forms throughout the range of this subspecies. Some populations are composed largely of a single variety, while others are a mixture of several forms. At Guajaibon, the type locality, there exists a pale yellow form, occasionally ornamented with a more or less distinct reddish sub- sutural line. The shell here is moderately lirate, dull, and rather coarse to the touch. The characteristic marbling is confined to the upper whorls. At San Andres a rather spectacular form is found. The early whorls are pale yellow but this soon gives way to a light reddish purple or roseate ground color, variously ornamented with irregular white or pale yellow marbling. This marbling is occasion- ally arranged in broad spiral bands distinguished by the differing nature of the spots. The ground color, as well as the marbling de- signs, becomes obsolete near the aperture and at the base, both of which are white. The lirations are very faint, not perceptible to the touch and barely visible under a magnifying glass, but the shell, though smooth, does not possess the brilliant sheen which occurs in some of the populations of the subspecies laevigata. In some col- onies in the San Andres region (e.g., El Aserradero near Caigua- nabo and Puerto Escondido), yellow shells occur together with pur- plish ones, as well as some that have varying mixtures of purple and yellow. The marbling in both color forms covers practically the entire shell. The bands sometimes consist of a solid color and hence resemble the prominent spiral bands of the subspecies laevigata. Occasional populations, as at La Sierra, have both the yellow and purple colors, which, however, are very pale and appear faded, and the marbling is widely scattered over the surface. At some isolated mogotes (Bella Maria and Talavera near La Palma, Hoyo de los Mogotes near Vinales, La Catalina near San Diego de los Banos, Mogote del Rojero between Cayo San Felipe and Isabel Maria, and some of the mogotes about Kilometer 14) a dwarf form occurs, generally in some shade of yellow. These remarks are not to be construed as having at all ex- hausted the range of variations of regina. Variations of all sorts occur, in sculpture, color, and size, sometimes in localities quite close to one another (as near Isabel Maria), sometimes even on a single mogote (as El Queque). It is not the purpose of this study to give a full account of all the forms in which regina or either of the other two subspecies occurs, or to give a detailed description of each population. Such an undertaking would lead us far be- yond our original purpose. Specimens examined. CUBA: Pinar del Rio. Vinales: Mogote W of Kilometer 14; Mogote E of Kilometer 14; Abra de la Colmena; 12 MRI-MORA No. 298 \rrovo Melindre; Encinar Alto; La Laguna, Potrero Constancia; Mogote de Jose Maria Garcia; Hoyo de la Sierra; Hato Morales; Hoyo de los Mogotes; \\o\o do los Santos; Mogote la Canona; Constancia; Mogote Capon; Mogote Dos Hermanns. Vinalet (/'tc las Cuevitas; Hoyo Paige de San Antonio. San Andres (Can- ulac). Los Hoyos; Mogote del Abra de Canalete; Ensenada de Borges; I nsenada de los Colorados; Mogote de Abraham; Ense- nada de Canalete. San .-Indies (Caiguanabo) : El Aserradero; Mogote Large Mogote Pucrtccitas; Mogote Andres Diaz; Mogote de Caiguanabo; Abra de Caiguanabo. s ^ -3 _ ' ' o" L. .-3 ^-« b_ Urn j) C , s '-I o O -3 J -_> E •* ■j H o H i/j -"y •— s O o L^ 0 '-< " 73 o 3C — uemad dc los pd u 3 X C 2 •J E ■■ L. t— U o j 1 K — y ~* fc a « u i ii Si los. |> — — «• s M d Im B u u « E y. E u 5 - / d rt « y 'j H u. _W rra d Siei i 175 E u U 7 < ' (il j r**t a. *- CUBAN GENUS VIANA 19 20 HRKVIORA No. 29S Map 4. Siena del Quemado to Sierra Guacamava. H, Sierra del Quemado. I. Siena Celadas; 11, Santo Tom&s (town). J. Sierra Pan de Azucar. K, Siena Martillo. L, Siena del Infierno; 12, Cayos de San Felipe'; 13. El Guama; 14, Mogotes de Km 14. M. Sierra Penitencia. N, Sierra El Abra. O, Sierra Vinales; 24. Mogote Palmarito; 25, Hoyo Magdalena. P, El Queque; 17. Mogote Vigil; IS. Mogote Rojas; 19, Abascal; 20. Dos Hermanos; 21. Vinales (town); 22, Hoyo de los Santos; 23. Ensenada Miranda; 26, Puertecites; 27, Mogote Capon. Q, 1 a Chorrera; 28, Hoyo de Jaruca; 29. Laguna de la Piedra; 30, MogOte Jutia; 31, Fogon de los Negros; 32, West ridge of the Siena San Vicente; 33. East ridge of the Siena San Vicente; 34. Ratios de S.m Vicente. R. Sierra Cialeras; 15. Pan de Azucar. S. Sierra Ancon; 61, San Cavetano (town). T, Sierra San Andres; 35, Mogote Mina; 36, Rinconada; 37, La Jagua: 38, AsientO de la lamia; 39, Mogote de la Jagua; 40. Hoyo Corto de San Antonio; 41, Canalete (town); 42, San \ndres ( town ) . 43, Mogote funic; 44. Mogote de Canalete; 45. MogOtes de Canalete; 46, Ensenada de la Ayiia; 47, Pasada de las Escaleras; 48, Puerto de San Andres; 50. Pico Grande; 51, c olmillo de la Vieja; 53, Sitio de la Sierra; 54, Mogote Bella Maria. U. Sierra Gaucamava; 55, Mogote Grande; 56, Caiguan- abo; 57, Pinalito; 58, Mogote I argo; 59, Mogote Angelena; 60. Mogote Mamey. I ocalities aumbered 49 and 52 represent place names that could not be located on an\ maps available to us. 1 Not to be confused with the small islands "Cayos de San Felipe" off the northwest coast of the Isla de Pinos. CUBAN GENUS VIANA 21 ~>-> HRF.VIORA No. 298 N i sKM 6« »7 M • »4 Map S Sierra Guacamaya to Sierra Rangel. U. Sierra Guacamaya. V, Sierra Cialalon; <> I . Caracoles; r>2. Mogote Colmena de Piedra; 63, I os Portales. W, Sierra La Giiira; '4. San Diego de los Banos (town); 65, Los Cayitos de Catalina. 66. Mogote Bosque; 67. I a < timbre. X, Pan de Guajaibon. \ . Sierra RangeL CUBAN GENUS VIANA 23 PLATE 2 Viana regina regina (Morelet) Fig. 1. Pan de Guajaibon, Consolacion del Norte, Pinar del Rio (para- type, MCZ 146706 9 ). Figs. 2, 3. Pico Grande, Sierra de San Andres, Consolacion del Norte Pinar del Rio (MCZ 256282(4 ). Fig. 4. La Catalina, San Diego de los Banos, Consolacion del Sur, Pinar del Rio (MCZ 126684 9 ) (all 2 X). 24 HRHVIORA No. 298 PI ME 3 I mini regina laevigata (Pfciffer) I k,s i. : ( ,i\ds de San Felipe, Vinales, Pinar (.Id Km (Topo types, \K / 126716 ). Fic ; sun., Celadas, Vinales, Pinar del Rio I \l( / 256173 |. lh, 4. Pan de Azucar, Vinales, Pinar del Rio (MCZ 126703 I (all : \). CUBAN GENUS VIANA 25 PLATE 4 Viuna regina subunguiculata (Poey) Fig. 1. Siena Celadas, Santo Tomas, Vinales, Pinar del Rio (MCZ 256342 & ). Fig. 2. Sierra Celadas, Santo Tomas. Vinales. Pinar del Rio (MCZ 256342 6 ), to show the outer peristome notch in the male. Fig. 3. Valle San Carlos, Luis Lazo, San Juan y Martinez, Pinar del Rio (MCZ 256335^ ). Fig. 4. Sierra de Guane, Guane, Pinar del Rio (Lectotype MCZ 73672 6 ) (all 2 X). BREVIORA Museum of Coeip&irative Zoology Cambridge, Mass. 11 October, 1968 Number 299 WHAT IS LUMBRICUS EISENI LEVINSEN, 1884 (LUMBRICIDAE, OLIGOCHAETA)?1 G. E. Gates2 Abstract. Levinsen's species, eiseni, after being in the synonymy of an unrelated form and in three lumbricid genera, is excluded from Bimastos, as defined by characters less liable to rapid evolutionary modification than those of the genitalia on which the classical system of the Oligochaeta (Michaelsen, 1900 — Stephenson, 1930) was mainly based. The species, now more adequately characterized, cannot go into any other lumbricid genus because of current ignorance of somatic anatomy in their type species. Relationships suggested by available anatomical data are with Allolobophora chlorotica and, less closely, with Dendrobaena rubida and Lumbricus spp. As the references below show, no agreement has been reached as to the status of Levinsen's species and as to its generic affilia- tions. The taxon, like other more or less widely spread lumbricid anthropochores as well as most endemics of the family, never was adequately characterized. Although much less material could be procured than was desired, study of new material has enabled the following contribution toward a solution of the problem. "Lumbricus eiseni" Levinsen, 1884 Lumbricus eiseni Levinsen, 1884, Vidensk. Meddel. Naturhist. Forhandl. Copenhagen, (4) 5:311, 241. (Type locality, Deer Garden, Copenhagen. Types, 5, in the Copenhagen Museum.) Allolobophora (Dendrobaena) mew'.-Friend, 1892, Jour. Linnean Soc. Lon- don, (Zool.), 24:302. Helodrilus (Bimastus) mem.-Michaelsen, 1900, Das Tierreich, 10:503. Bimastus eiseni, Friend, 1911, Zoologist, (4) 15:145. Eisenia parva f. typica (part) Pop, 1950, An. Acad. Romane, Sect. Sti. Geol. Geogr. Biol., (A) 1 (9):89. Eisenia eiseni, -Graff, 1953, Regenwiirmer Deutschlands, p. 27. 1 From research financed by the National Science Foundation. 2 Zoology Department, University of Maine, Orono, Maine, U.S.A. 2 breviora No. 299 SPECIMENS EXAMINED Channel Islands. Herm. under rock above tide mark, June 24, 1925. 1-0-0. G. E. Pickford. South Africa. Cape Province. Cape Peninsula. Kirstenbosch, rotting stump, October 29, 1925, 5-0-4. Table Mountain, Window Gorge, rotting log at 400 feet, July 30, 1926, 0-0-9. Brockenhurst. Queen's Bower, April 2, 1925.0-0-1. G. E. Pickford. Interceptions by U.S. Bureau of Plant Quarantine i ire. From soil with twenty-nine narcissus bulbs and two bundles of lily plants, in baggage on plane, arriving at Boston, June 22, 1964, 0-0- 1. Scotland. From heather plants, in plane, arriving at New York, October 26, 1949.0-0-1. Germany. From cabbage, in ships stores on Norwegian M S I ancing, ar- riving at Savannah, Georgia. October 2l>. 1964, 0-0 1. Portugal. From soil with five azaleas, Bve ferns, two orchids, in baggage on plane, arming at Boston. M,i\ 21, 1965, 1-0-0. Azores. From soil with one begonia, in baggage on plane, arriving at Bos- ton, June 18, 1965,0-0-1. From roots of plains o\ ( 'Una sp.. in baggage on plane, arriving at Bos- ton. June 18. 1965. 2-0-0 DESCRIPTION External characteristics. Segments. 79-1 12 (cf. Table 1 ). Color, brown, slate to almost black but never red, restricted to dorsum except in ii-vi or vii, most obvious anterior to the clitcllum, usually lacking or very sparse in buccal and anal segments. Body sub- circular in cross section posteriorly. Prostomium, tanylobous (all). Setae, all present in ii. the d setae at or below mL anteriorly but posteriorly above that level. BC usually < AA throughout post- clitellar region where DD < AC. Nephropores, obvious. On one South African worm they are above B on the left sides of x-xiii, xv, xvi, xix-xx, xxvi, on the right sides of ix-xi. xv, xxii-xxvi, xxix. Other pores of that worm, in the region of iv-xxix. are well above D. Pores of xv-xvi, except on one other worm (left pore of xv not found), were lateral to B. Pores may be at the same level on one side for three, four, or five consecutive segments, but on the opposite side asymmetry is oc- casional. Pores of xii have been seen (rarely) just below C. The arrangement throughout most of the axis can be characterized as: irregularly alternating, usually between two levels, one slightly lateral to B and another about midway between D and wD.with occasional intrasegmental asymmetry. 1968 LUMBRICUS EISENI LEVINSEN 3 First dorsal pore, at 5/6 (21 specimens), occasionally smaller than the one at 6/7. Female pores, slightly lateral to B, equatorial in xiv. Male pores, minute, at or median to mBC, each laterally in a small cleft at eq/xv. Male tumescences, confined to xv and to median half of BC. Clitellum, red (formalin preservation), saddle-shaped, reaching ventrally to or nearly to A, or mostly annular, in xxiv-xxxii but restricted to DD in xxiv and xxxii (13), xxiv-xxxiii/n (1), xxv- xxxi (2, possibly not fully developed). Even at apparent maximal development the clitellum is thinner in AA than laterally and inter- segmental furrows still are recognizable ventrally from mBC. Genital tumescences, often conspicuous, without distinct boun- daries, transversely elliptical, including a,b, in xvi (27), xxiv (2), xxiv-xxv (l),xxv (26). Internal anatomy. Septa, none thickly muscular, 10/11 bulged anteriorly and 11/12 posteriorly by the calciferous gland, the coelomic cavity of xii, xiii and xiv quite small. Pigment, lacking in peritoneum, present anteriorly in the longitudinal muscle band at mD but disappearing behind the clitellum, associated with or in circular muscle layer, apparently brown, yellow brown, or dark brownish red. Pharyngeal glands to 8/9. Brain, circumpharyngeal connectives and subpharyngeal gan- glion all left in iv (19), sometimes well posteriorly, by a transverse section exactly along 3/4. Ventral bundles of the longitudinal musculature, according to Prof. Harman (in litt.), can be characterized as fasciculate, the pinnate arrangement so characteristic of some species of Lumbricus being lacking. The cells form compact "Kastchen" ventrally and a compact band dorsally. Ventrally the bundles resemble those of A. chlorotica. Calciferous sacs, in x, elongated horizontally rather than ver- tically, anteriorly or anterolaterally directed, reaching to or nearly to 9/10, apparently opening posteriorly into gut lumen at level of insertion of 10/11. Calciferous lamellae, extending nearly to anterior end of sac but in x small and of squarish shape. Gut much widened, rather bead-shaped and with vertically slit-like lumen in xi, narrower (half as thick as in xi) and of nearly uni- form calibre through xii-xiv. A deep internal constriction (or vertical groove) on one or both sides at or slightly behind insertion of 11/12. Calciferous lamellae, large and rectangular in xi, but behind insertion of 11/12 quite small, not reaching 12/13 (if present posteriorly too small to be recognized in dissections). The inner lining of the gut was rotted away in slightly macerated speci- mens so as to reveal lamellae with diagrammatic clarity. Intestinal 4 breviora No. 299 origin, in x\ (20). Gizzard, in xvii (20). the layei with brilliant muscular sheen in horizontal sections narrowing behind insertion of 17/18 and not reaching 17/18. A postgizzard valve, it" present, is relaxed in each worm but the gut is somewhat narrowed at 18/19 or in xix. Tvphlosole. beginning gradually in region of xx- \\i\. at first a low horizontal band, then becoming vertically and rather thickly lamelliform. A slight longitudinal groove may be recognizable on the ventral face at mV. or, if not. a slight longi- tudinal flange ma) be visible on each side. (Condition does not allow a more precise characterization.) The tvphlosole ends, abruptly in unamputated specimens, in xc-xevii (Table 1 ), leaving 8-15 segments (proctodeal?) atyphlosolate. Dorsal blood vessel, continued onto pharyngeal bulb. Sub- neural, adherent to nerve cord, l:\tra-oesophageal trunks, median to hearts, posteriori) in xii passing up to dorsal trunk (21). Hearts, present in viii-xi (22). Quite slender vessels at sites (in other species) of hearts o\ vii were traceable to ventral trunk only in three worms in each of which onlv the dorsalmost quarter is heart-like. Nephridia, vesiculate. Bladders, each in shape oi a long U- shaped loop, from A well into DD, both limbs of the loop of about the same length, posterior limb passing into parietes near H. The one other major loon is longer than the bladder loop. Holandric. Testes, flattened horizontal discs sometimes rather fan-shaped or saccular and pyriform. Male funnels, crenellate. large. Male deferent ducts, rather slender, almost straight, slightly sinuous, with several, very short, u-shaped loops, once a single long hairpin-loop, or coiled into a ball, loops or coils up to a millimeter awa) from funnel septum. Ducts of a side, uniting posteriorly in xii. continued to cq w across top of atrial gland (latter developed after penetration of duct into parietes'.') or disappearing from sight at edge oi gland. Seminal vesicles, rather small to small but not vestigial, vertical or dorsal, soft, without obvious lobing, in xi-xii (15). Anterior vesicles, flattened in the space between septum 10 11 and the anterior face of the calciferous gland. Ovaries, often large and then reaching up to level of dorsal face of gut. filled except in proximal quarter or so with large ova, each with a single egg string containing one to three ova. Ovisacs, with one to five ova which were easily released by teasing, especially those in a distal portion. Oviducts, rather thick passing into parietes lateral to D. Atrial glands (14 clitellate worms), confined to median portion of BC or reaching nearly to C, flat or of a rather low dome-shape, usually with a cleft at eq/xv and another at insertion of 15/16. 1968 LUMBRICUS EISENI LEVINSEN 5 reaching to or well toward 16/17. TP glands, lacking (all). GS follicles, large, conspicuously protuberant into coelomic cavities, genital setae elongated and otherwise modified. Follicles of ventral couple of xv not as much enlarged as the others. Supraparietal glands around GS follicles, lacking (all). Follicles of all ventral couples, from the clitellum anteriorly, larger than those of the lateral setae. Variant specimen. Clitellum, light pink rather than red, reach- ing little below C, but of near maximal thickness in region of mD except at intersegmental furrows which are still obvious even there. Male funnels, smaller than usual but crenellate, with very slight iridescence. Male deferent ducts pass into parietes at eq/xv. Atrial glands lacking (no rudiments recognizable in longitudinal musculature). GS follicles smaller than usual. Distal two-thirds of ovaries with apparently mature ova. No ova in ovisacs. The clitellum probably had not attained full development. Absence of atrial glands does not, however, appear to be attrib- utable to immaturity. Hence the worm may be of a strain (possibly with obligatory parthenogenesis and markedly reduced spermato- genesis) in which the atrial glands already have been lost and in which genital setae as well as tumescences are disappearing. Elim- ination of genital tumescences and genital setae would result in a state similar to that of Bimastos species. Reproduction. Spermatophores were present externally in clitellar region of two worms but no iridescence was recognized in the opaque matter within the transparent cases. Spermatozoal iridescence was lacking on funnels of one worm but on those of other clitellate worms does not seem to be as brilliant as in species with obligatory cross-breeding, and was not recognized in male deferent ducts. Sparse maturation of sperm is associated, in some lumbricids, with parthenogenesis. Pyriform testes are unusually soft, much of the interior filled with a watery fluid in which there are many morulae but only a very few sperm. Uniparental repro- duction has been demonstrated for this species by Evans and Guild (1948). Bimastos eiseni is diploid, 2n = 32, according to Muldal (1952:61). Parthenogenesis is not, however, contraindicated by the diploidy, that method of uniparental reproduction being obliga- tory (idem:66) in diploid individuals of Octolasion tyrtaeum (Orley, 1881). Remarks. Soil particles of any sort were completely absent from guts of each of these worms, the lumen being filled with brown organic matter. 6 BREVIORA No. 299 Distribution. Ireland, Scotland, England, Channel Islands. Sweden. Denmark. Germany. Poland. Belgium, Czechoslovakia, I ranee, Switzerland, Austria. Italy (including Sicily). Jugoslavia. Albania. Bulgaria. Portugal. Azores, Madeira. Canary Islands, South Africa, St. Helena. India (Western Himalayas only), New Zealand (including Stewart Island ). The extra-European records show that the species has been fre- quently transported by man with man) successful colonizations. Absence from the Americas where so main other lumbricid anthro- pochorcs are well established is puzzling. In Europe, presence above the southern limit of Ouaternar\ glaciation may also be a result of unintentional human carriage. 1 he original home of the species is unknown. It must have been somewhere in Eurasia but how much oi the distribution in southern Europe is fortuitous, remains to be learned. I ( »cies has been called cosmopolitan < Omodco, 1950:5) and Euro-American (Omodeo, 1961, table). The first adjective hardl) seems applicable to a species absent from all Americas. most o\' Africa and all oi Asia excepl for a couple oi localities in the western Himalayas. The second characterization is in- applicable. Systematic relationships. The clitellum is shorter posteriorly than in English definitions (Cernosvitov and Evans, L947; (icrard, 1964). Paucity oi data about individual variation in eiseni at present ob\ iates further discussion of the differences. The species docs not belong in Bimastos as that genus now ((iates. in press) is defined. The calciferous sacs of segment x suggest relationships with / umbricus, Dendrobaena rubida and related species (but certainly not 1). octaedra and related species), as well as Allolobophora chlorotica. Further evidence is provided by the similarity, in each taxon, of nephridial vesicles and ncphro- pore locations. Main characters provided by those and other somatic systems now seem resistant to rapid evolutionary modi- fication and so are oi greater systematic importance than the more easily changed genitalia on which classical specialists so much relied. The purple red pigment by which Dendrobaena, Eisenia and Lumbricui were defined (Pop, 1941 ) is lacking in chlorotica and eiseni. The longitudinal musculature of Lumbricui spp. and of D. rubida (also closely related species?) is pinnate in cross-section rather than fasciculate as in eiseni and chlorotica. Atrial glands (ignored by classical authorities) are indeed present in D. rubida 1968 LUMBRICUS EISENI LEVINSEN 7 and related species just as in eiseni and chlorotica but are lacking in all species of Lumbricus studied by the author. Those differ- ences seem to suggest that chlorotica and eiseni may be more closely related to each other than either is to any of the other above-mentioned taxa. Then, should eiseni be transferred to Allolobophora of which chlorotica recently was made the type species? If the two species do belong in the same genus, that is where eiseni seemingly should go. A. chlorotica, however, differs from every one of the common allolobophoras by its calciferous sacs, by the presence of well developed atrial glands (lacking in every one of the common allolobophoras), as well as by other characters. Information now available indicates that: 1) Enterion chloroticum Savigny, 1826, should not have been designated the type of Allolobophora. 2) All of the common allolobophoras, if that type designation in some way cannot be nullified, must go into another and possibly new genus. The common species of the classical Allolobophora are now widely distributed throughout much of the world. They success- fully colonized many areas to which they were introduced and even now are spreading, and being spread both accidentally and deliberately, through considerable areas of Australia and New Zealand. Those same species also have been widely used in experi- ments. A. chlorotica and B. eiseni, on the contrary, were much less often transported, have not colonized so extensively, are much less common, and only rarely have been studied experimentally. Attention was directed on two previous occasions (Gates, 1958a, b) to the fact that eiseni does not belong in the American genus Bimastos, primarily in the hope that some interest in the problem would thereby be aroused on the part of those who have access to more plentiful material. Because of our ignorance of so much essential data for a large majority of lumbricid species, no nomenclatural changes are made herein. The single North American anthropochore, Bimastos parvus (Eisen, 1874), is not of the same genus as eiseni and just as ob- viously does not belong in Eisenia. What Eisenia parva of sundry authors was, probably can only be determined from an examination of the specimens. Parasites. Nematodes from coelomic cavities of the Scotch worm, according to Dr. S. Prudhoe of the British Museum, prob- ably are Dicelis filaria Dujardin 1845. Nematodes from the ventral blood vessel in x-xiv of a Channel Islands juvenile were identified by Dr. Maybelle Chitwood as larvae of Spiroptera turdi Molin by Cori, 1898. 8 breviora No. 299 REFERENCES CITED Cernosvitox . I ... and A. C. Evans 1947. Lumbricidae. Linnean Societj of l ondon Synopses of British Fauna, No. 6: 1-36. Evans, A. C. and W. J. di ild ll>48. Some notes on reproduction in British earthworms. Ann. Mag. Nat. Hist . (1MI 1:654-659. Gatks. G. E. 1958a. On endemicity of earthworms in the British Isles \nn. Mag Nat. Hist . ( 13) 1:33-44. 1 958b. Contribution to a revision of the earthworm family Lum- bricidae. II Indian species. Breviora, Mus. Comp. Zool., No. <>l:l-16. In press. On tWO American genera of the earthworm family I um- bricidae. Jour. Nat. Hist (,l RAKD, B. M 1^64 Lumbricidae. linnean Societj of London, Synopses oi British i auna, 2nd ed . No. 6:1-58. Ml I DAI . S 1952 I he chromosomes of the earthworms. I. I he evolution of poly- ploidy. Heredity, 6:55-76. ( )M()DI (). P. 1950. Oligocheti. \nn. Inst Mus. Zool. Univ. Napoli, 2(10): 1-12. 1961. l e peuplemenl des grandes iles de la Mediterranee par lesoligo- chetes terricoles, Colloques Internatl. Centre Natl. Rech. Sci., No. 94:127-133. Pop. \. 1941. /ur phylogenie und Systematik der Lumbriciden. Zool. Jahrb. Sysl Okol.Geogr. Tiere, 74:487-522. (Received 16 January 1968.) 968 LUMBRICUS EISENI LEVINSEN TABLE 1 Typhlosole termination and segment number in "Lumbricus eiseni" Atyphlo- Number Serial Typhlosole solate of Number ends in segments Segments Remarks 1 70 9 79 Posterior amputee* 2 74 7 81 Posterior amputee* 3 82-85 3 88 Posterior amputee* 4 80-86 86 Posterior amputee* 5 88 11 99 Channel Islands 6 90 9 99 7 90 13 103 8 91 13 104 9 92 9 101 10 92 12 104 Posterior amputee* 11 92 15 107 12 93 12 105 13 95 8 103 Posterior amputee* 14 96 13 109 15 97 15 112 Specimens from South Africa unless otherwise indicated. * After loss of the posterior end, the presently terminal metamere had been reorganized and now looks much like a normal anal segment. Proof of its original, preterminal position is provided by vestiges of former apertures of setal follicles. The latter were lysed after dehiscence of their setae. BREVIORA Museumi of Comparative Zoology Cambridge, Mass. 11 October, 1968 Number 300 CONTRIBUTIONS TO A REVISION OF THE LUMBRICIDAE. III. EISENIA HORTENSIS (MICHAELSEN) (1890).1 G. E. Gates2 Abstract. The classical system of the Oligochaeta (Michaelsen, 1900- Stephenson, 1930) and proposed neoclassical revisions, did not permit defini- tive conclusions as to disposition of a taxon variously known as var. or /. hortensis which has been in each of five lumbricid genera. Characters less liable to rapid evolutionary modification than those of the genitalia, on which previous systematics primarily had been based, allow inclusion of the taxon, as a distinct species, in a genus recently redefined in terms of formerly neglected somatic anatomy. Parthenogenesis may be facultative in some lines. Evolutionary developments now under way in hortensis parallel some of the degradations in genital anatomy associated, in D. rubida, with parthe- nogenesis and male sterility. Characteristics of an undegraded amphimictic morph are predicted. Abnormalities are characterized. To the distribution, as previously known, are added Ireland and Greece. Disagreements as to systematic status and generic affiliations of various species (one example provided by the synonymy below) indicate a need for revision of lumbricid classification. Even widely spread anthropochores, that usually, though mistakenly, are be- lieved to be adequately characterized systematically, need a search- ing reinvestigation. A single Irish specimen that arrived on St. Patrick's day provided a clue to a possible solution of some of the problems hitherto asso- ciated with "dendroidea." LUMBRICIDAE Eisenia Malm, 1877 Eisenia hortensis (Michaelsen, 1890) Allolobophora subrubicunda f. hortensis Michaelsen, 1890, lahrb. Ham- burgischen Wiss. Anst., 7: 15. (Type locality, Hamburg. Types, orig- inally in the Hamburg Museum.) 1 From research financed by the National Science Foundation. - Zoology Department, University of Maine, Orono, Maine. 2 BREVIORA No. 300 Allolohophora (Notogama) veneta f. hortensis Michaclsen. 1900, Abhandl. Naturwiss. Ver. Hamburg, 16: 12. Helodriliu (Eisenia) venetus \ar. hortensis.-Michaelstn, 1901. Ann. Mus. Zool. Acad. Sci. St. Petersbourg, 15: 37. Allolohophora {Eisenia) veneta dendroidea Friend. 1909, Gardener's Chron- icle. 46: 243 (Type locality, St. James Garden. Malvern, Worcestershire, England. T\ pes. in the British Museum.) Allolohophora ilu.se/iiii) veneta var. hortensis Michaelscn, 1922. Cap. Zool. 1 (3): 13. Eisenia veneta var. dendroidea Friend. Il>23, British Earthworms, London, p. 30. Eisenia veneta van dendroidea 4- /■.". v. var. hibernica f. dendroidea Cer- aosvitov, 1M42. Proc. Zool. Soc. London. Ill: 240. 241, 274. / isenia veneta \ar. hibernica f. dendroidea 4- E. V. var. hortensis Cernosvitov and Evans, 1947, Linnean Soc London, Synopses of the British Fauna. No. 6: 25. Dendrobaena hortensis (part '.'» 4- D. h. dendroidea Omodeo. 1955, Ann. 1st. Zool. Univ. N.ipoli. 7: 6. 8. (Excluding />. hibernica.) I tenia veneta ^ar. hibernica f. dendroidea dates. 1958, Ann. Mag. Nat. Hist., (13) 1: 34. (Name Of the ta\on should have been enclosed by quotation marks' ) Dendrobaena veneta var. hibernica f. dendroidea 4- D. v. var. hortensis Gerald. 1V64. linnean Soc. 1 ondon. Synopses of the British fauna. No. 6 (2d ed.): 19 Bimastos veneta Causey. 1953. Proc. Arkansas Acad. Sci. 6: 47. SPECIMENS EXAMINED Oregon, Yamhill County. McMinnville, kitchen drain, August 1 1. 1944, 0-o_f3. Backyard, March 12. 1945, 0-1-1. Backyard, brought out bv wash water. October 31, 1945. 0-0-1. Chicken yard, under cow manure, November 14. 1945, 0-1-0, Novem- ber 23. 1945. 0-4-10. Under wet paper carton on back porch, January 10. 1946, several specimens. Backyard, December 1,1947.0-1-7. D. McKey-Fender. California. San Francisco County. Arboretum of Golden Gate Park, March 28, 1951, 0-0-2. H. B. Leech per D. McKey- Fender. Ohio. Trumbull County. Masury, greenhouse, 0-0-2. W. R. Mur- chie. Virginia. Giles County. Goldbond, sawdust, August 21, 1955, a number of specimens. W. A. Harman. Italy. Naples, November 23, 1925. 0-0-2. Posillipo, October 7, 1925, 0-0-7. G. E. Hutchinson per G. E. Pickford. India. Darjiling, Northpoint, at ca. 6250 feet, June 1945 0-1-38. 1968 EISENIA HORTENSIS MICHAELSEN 3 Interceptions by U.S. Bureau of Plant Quarantine. Ireland. From 25 pounds of wood duff with seeds of Crataegus oxycanthus in mail arriving at Hoboken, November 1, 1963, (?)-l-0. Greece. From soil with 25 unidentified herbs in baggage on plane arriving at Boston, August 28, 1962, 6-0-0. DESCRIPTION External characteristics. Size, 26 by IV2 mm (Ireland), 26-32 by 2-2^ mm (Maine, New York), to 22 by V-A mm (Greece), to 37 by 2+ mm (Ohio), 42-54 by 2*/2-3 mm (Naples), width measured in the clitellar region not always at maximal tumescence, thickness of adults near hind end W2 mm. The body is dorso- ventrally compressed behind the clitellum and almost oblong in transverse section, ventral and lateral sides flat, the dorsum slightly convex. The b and d setae are at the four corners of the body. Segments, 42, 50, 55, 56, 60 (2), 64 (2), 69 (2), 70 (2), 74, 76, 77, 81, 82, 83, 84 (2), 85, 86 (2), 88, 90 (3), 92 (3), 94 (3), 95 (7), 96 (4), 97 (5), 98 (4), 99 (2), 100 (3), 101 (2), 102 (3), 103 (4), 104 (2), 105, 108, 115 (2), 124, 126, 130. The mean number of segments for 69 of the specimens is 90.7536, with a standard deviation of 17.4266 and a standard deviation of the mean of 2.0979. A large majority of unamputated worms have segments in a range of 90-105 which may be more useful systemati- cally than the computerized figures. Worms with 1 15-130 segments were from Maine greenhouses in which they might almost be said to have been force-fed. Color, unrecognizable after alcohol preser- vation, otherwise light to bright red, reddish, restricted to dorsum and lacking below C except in front of xv and near the hind end where the ventrum may also be colored. In some Greek worms, color is in discontinuous unpatterned areas. Prostomium, epilobous, tongue open (all). Setae, present from ii where none usually are lacking though sometimes hard to recognize, not closely paired, width of AB and CD somewhat variable but CD slightly < or > AB < BC < or > A A < DD which is of course < Y2C, the c setae often seeming to be about at mL. In some worms, AB is ca. = V2AA and slightly < BC. Nephropores, inconspicuous, in xv-xvi at or just above B ( 39 specimens ) , one pore of xv at D ( 1 ) , one pore of xvi at D ( 1 ) , both pores of xiv at D (4), at B (10), one of xiv at B the other at D (22), usually unrecognizable in iii-vi or vii. Nephropores, at B on left sides of 17th, 19th, 21st, 24th, 26th, 29th, 30th, 33d, 34th, and on the right sides of the 17th, 23d, 25th, 27th-29th, 31st-33d 4 breviora No. 300 segments (No. 1), on left sides of 14th-20th, right sides of 15th- 17th. 19th. 20th segments (No. 2). On the leftside of No. 3 the following locations were noted, at B, 35th-40th, 52d. 57th. 60th. 61st, 64th, 65th, at /) in the 34th. 42d-51st. 53d-56th, 58th-59th, 62d-63d. Pores on the opposite sides of those same segments more often than not were in the alternate location. Both pores of vi were in CD, one Italian worm on which the right pores of 7th, 8th. and 1 Ith segments were near B, of 12th at D, but those of 1 3th- 1 7th in BC. Nephropores. just above B, on left sides of 7th. 8th. 11th, 14th-16th. lSth-22nd. 24th-32d. on the right side of 8th, 9th, 12th-15th. 17th-34th (Naples, No. 4). on the left of 7th- 10th. 12th. 13th. 15th-22d. 24th. 27th, right sides of 7th-8th, 10th-l 2th. 15th-19th. 22d. 24th. 26th (Naples. No. 5). At pres- ent, about the only seemingly feasible characterization is: pores usually alternate irregularly and with asymmetry between two ma- jor locations along most of the anteroposterior axis. First dorsal pore, at 5 6 (22 specimens), at 7 8(2). Spermathecal pores, minute, superficial, at 9/10-10/1 1, close to mD in an area where coloi often is faint or quite unrecognizable. I emale pores, minute, superficial, equatorial in xiv and just lateral to B. Male pores, minute, not superficial, within a cleft laterally and seemingly about at ////>'( Male tumescences, almost confined to median half of BC but obliterating 14 15-15/16 and reaching well into w and xvi. Clitcllum, saddle-shaped, at maximal tumescence reaching to or nearly to B, dorsal pores occluded, intersegmental furrows oblit- erated, setae unrecognizable, xxvii-xxxii (15 specimens), xxvii- xx.xiii (14). eq xxvii-xxxiii, eq (2). eq 'xxvii-xxxiii (2), xxviii- x.xxii (29), xxviii-xxxiii (3), eq xx\ iii-xx.xiv/cq (9). Tubercula pubertatis, usually longitudinal bands of greyish transluccnce just lateral to B, with straight median margins but with lateral margins slightly convex, in xxx-xxxi but occasionally extending slightly into \\\ii. rarely reaching eq/xxxii. A deep groove occasionally de- marcates a tubcrculum laterally. Even when a red coloration (post-preservation artifact) of the clitellum is best developed, none is recognizable in the tubercula. Genital tumescences, often rather indistinct and especially so in relaxed material, include setal couples as follows: a, b of xi ( 1 specimen), xii (4), xvi (2), xxii (1 ), xxiii (7), xxiv (7), xxv (1 ). xxvi (1), xxix (26), xxx (35), xxxi (35), xxxii (9), c,d of x ( 1 ), xi (22). Condition obviated recording locations on a number of worms but dissections indicate that ventral setae of xxx and xxxi always were associated with tumescences. Follicles opening 1968 EISENIA HORTENSIS MICHAELSEN 5 through tumescences are thickened and have slightly enlarged apertures. The b,c and d follicles project conspicuously into the coelomic cavities but a follicles obviously are shorter and protrude only slightly above the parietes. Internal anatomy. Septa, 13/14-14/15 often thickest and quite muscular, 10/11 near the gut separated into two lamellae the an- terior of which passes straight to the gut, the other inserted on it more posteriorly so as to leave a space filled by a watery fluid with- out corpuscles or other solid matter. This curious situation pre- sumably arises as a result of an ontogenetically late elongation of a small section of the esophagus. The anterior lamella seems to be at the front end of the calciferous gland. Setal gaps in longi- tudinal musculature, obviously eight. Special longitudinal muscle band at mD, present from 5/6. Pigment, red, in circular muscle layer, lacking immediately under intersegmental furrows. Pigment also is visible on or in the coelomic face of the parietes, dorsally throughout but also ventrally in front of xv, in flecks, or spots or rarely even in larger patches. Location of the brain is variable, a section exactly along 3/4 sometimes left the brain anteriorly in iii, sometimes anteriorly in iv. Pharyngeal glands, through iv-v to 5/6. Esophagus of x has the shape of a truncate cone, narrower an- teriorly. The wall is dark (gorged with blood?) and un thickened (no sacs or lamellae). The calciferous gland, insofar as could be learned from dissection, is between 10/11 and 12/13, sometimes not reaching the latter, usually not constricted at 11/12 but of the same thickness through both segments. Rarely the portion in xi is moniliform. Intestinal origin, in xv (62). Gizzard, superficially appearing to extend through xvii-xix. Actually, the muscular layer becomes much thinner just behind apparent level of insertion on gut of 17/18. Gut often valvular in region of insertion of 19/20. Typhlosole, beginning rather abruptly in region of xxi-xxv, thickly lamelliform, height variable, when gut is empty sometimes reaching or nearly touching intestinal floor, ending abruptly, as shown in Table 1. The typhlosole does not end abruptly in specimens 1-5, 8-9, and 33 (cf. Table 1) but becomes much lower or even vestigial in the indicated regions, i.e., in lxi-lxv of No. 9. Posterior amputation had been involved in each of those cases. The typhlo- sole, in a very large majority of the unamputated individuals, ended in a region comprising lxxiv-xcv. The typhlosole of the above- mentioned specimens, on the contrary, ends in a region comprising xxxix-xlv, leaving 1-10 intestinal segments atyphiosolate. Very 6 BREVIORA No. 300 probably, the typhlosole had completely disappeared in some seg- ments of each oi those worms. In unamputated worms with abrupt typhlosolar termination. 10-15 segments were atyphlosolate, 10 (3), 11 (5), 12 (7). 13 (13), 14(6), 15 (2). [f, as some have thought, the atyphlosolate portion of the gut is of proctodeal origin, then that part of the gut extends through more segments than the embr\ ologistS indicated. Extra-esophageal trunk, passing up to dorsal \essel in \ii. but often unrecognizable (because empty) behind 10 II. In \ii (one specimen) the trunk bifurcated, the larger branch joining the dorsal \esse!. the other branch passing \entrall\ and shortly disappearing. \ branch from the trunk OCCasionall) passes up to the dorsal trunk in \. When that connective is distended with blood ami the pos- terior portion o( the trunk is empt\ the cxtra-csophagcal could be thought to end in x. I he ventral trunk is complete, bifurcating over the subpharyngeal ganglion. The dorsal trunk is also com- plete, having been traced forward to the brain when blood is present. I he subneural trunk probabl) always is complete but USUall) is not traceable throughout because of lack o\ blood. Hearts, present onl\ in \u-\i (62 specimens). Ncphridia. present from in . presence or absence in ii not de- mined), vesiculate. Bladders, sausage-shaped (ellipsoidal), transversel) placed, reaching to or nearb to ( or even beyond ( . joined lateralis by the tubular portion of the ncphridium. passing into p.: it or near B. restes, fan-shaped. Male funnels, plicate, size normal with ref- erence to that oi the body. Sperm ducts, without epididymis. Seminal vesicles, in ix. xi and \ii. Atrial glands, large, dcepb cleft at ei| x\. extending through xi\ as well as x\i. TP and atrial glands, acinous, supraparietal. CiS glands, acinous, each lobe narrow ing to a slender thread-like duct that passes straight down in the parietes. associated only with the .' Mesodont, relatively long; lophs and lophids rather stout, tending to form transverse lamellae; flexi and flexids (or transverse valleys) fairly persistent. Echimys, Diplomys, Central and South America, hothrix, South America. Recent. Dactylomyinae late 1 *-> 3 5 . Brachyodont, wide, large relative to e oi skull: lophs and lophids rather thin, sharp and with ten- dencies toward acquisition k)\ lamellae; flexi and flexids persistent. Dactylomys, Thrinacodus, Kannabateomys, South America. Recent. Heteropsomyinae Anthom 1917. Brachyodont to hypsodont, with some unilateral hypsodonty; upper cheek teeth tending to become subcircular to quadrate in outline; flexi and flexids not \er\ peis, stent, converting to Eossettes and fossettids after moderate wear; small antcroinicrnal fossettid may be present in lower molars. Proechimys, Central and South America. Recent. Puerto Rico, uncertain age; Hoplomys, Central and South America, Re- cent; Euryzygomatomys, Clyomys, Mesomys, Lonchothrix, South America. Recent; Carterodon, South America. Pleistocene to Re- cent; Cercomys, South America. Pliocene to Recent; Heterop- somys, Homopsomys, Puerto Rico, latest Pleistocene or early Re- cent; Brotomys, Hispaniola, latest Pleistocene or early Recent; Boromys, Cuba, iatest Pleistocene or early Recent; Eumysops, South America, late Pliocene to Pleistocene; Protacaremys, South America, late Oligocene. Protadelphomys, South America, late Oiigocene. is tentatively placed here, and there is an undescribed genus in the Santa Cruz Miocene of Patagonia. Myocastorinae Ameghino 1904.- Brachyodont to hypsodont; labial flexi and lingual flexids of later forms converting to fossettes and fossettids after considerable wear; later forms showing an in- crease in size and in obliquity of lophs and lophids from front to 'As Echinnina ( = Echymyna Gray 1825, Echsmidae Bonaparte 1X45). -'Proposed as a family ( = Myopotamini Tullhcrg 1900. Myiopotamyina Bonaparte 1850). 1968 OLIGOCENE ECHIMYID RODENTS 7 back; drn4 with a small anterior fossettid, with five lophids in later forms; M3 usually with four lophids, metalophid either retained or a third lophid reacquired. Myocastor, South America, Pliocene?, Pleistocene to Recent (and currently doing only too well in the feral state in Europe and North America); Paramyocastor, Iso- myopotamus, South America, late Pliocene; Spaniomys, South America, early Miocene; Prospaniomys, South America, late Oli- gocene. Tramyocastor does not appear to be separable from Myocastor. The new genera to be described, one of them Deseadan, the other Colhuehuapian, are both adelphomyines and appear to be more closely related to each other than to any of the previously known forms. DESCRIPTION OF NEW FORMS Xylechimys1 gen. nov. Type species: X. obliquus1 sp. nov. Distribution: Deseadan, Patagonia. Diagnosis: Reduced metalophid present on all molars, joined to metaconid to isolate antero-fossettid but continuing on into meso- flexid to define small anteroflexid, metalophid smallest in M3, sub- equal in Mi_2; hypolophid and posterolophid very oblique, espe- cially on M^; posterolophid terminating lingually near median line of tooth row; principal flexids deep, all of nearly equal depth on M2_3, metaflexid shallower on Mi; metaconid sharply angulate posteriorly. Xylechimys obliquus sp. nov. Type: Museo de La Plata No. 59. II. 26. 81, portion of right ramus with Mi_3, base of I and alveoli of P4. Hypodigm: Type only. Horizon and locality: Deseadan Oligocene; Laguna de los Machos, Campo Belisario Escobar, Departamento Paso de los Indios, Chubut, Argentina.2 Diagnosis: As for the genus. 1 £v\ot, wood, plus Echimys. Named for Professor Albert E. Wood in rec- ognition of his contributions to our knowledge of the order. The specific name refers to the obliquity of the posterior lophids. 2 This is a new locality that has yielded a small faunule. Found with Xylechimys were specimens of Asteromys, Archaeohyrax, Plagiarthrus, Pro- pachyrucos, Prohegetotherium, Trachytheriinae indet., which are sufficient to reveal the age. 8 BREVIORA No. 301 Description: The anterior cheek tooth is unfortunately repre- sented only by its broken alveoli. X-rays reveal no trace of a tooth below these alveoli. It is virtually certain that the missing tooth was P4, as in Deseadomys. The molars are mesodont and exhibit no tendency toward uni- lateral hypsodonty. ML. is the largest, and M:, the smallest of the scries, but the size differences are not great. All are broader rela- tive to length than in Deseadomys and are wider anteriorly than posteriorly, most aotabl) as regards M:t. The anterior portions of the molars are short anteroposterior^. The anterolophids are Dearly transverse. A reduced metalophid is present on all teeth, that of M:: being the shortest, those of Mi.2 sub-equal. Near the middle of its course each metalophid is united to the metaconid to isolate a small anterofossettid, and then con- tinues linguallv beyond this junction for a short distance. A short B I ig. 2. Xylechimys obliquus gen. et sp. nov. x 10. Type. Museo de La Plata No. 59. II. 26. 81. A, RM, ,, crown view; B, RM, . lingual view. 1968 OLIGOCENE ECHIMYID RODENTS 9 and shallow anteroflexid is thus present in addition to the antero- fossettid, a marked contrast to Deseadomys arambourgi. The hypolophid is decidedly oblique, being almost in a direct line with the ectolophid. On Mi.2, but not on M3, this crest is slightly swollen lingually, the swelling marking the position of the ento- conid. The posterolophid is essentially parallel to the hypolophid and terminates posteriorly a short distance lingual to the mid-line of each tooth. The metaconid is sharply angulate posteriorly. Hypoflexid and mesoflexid are of equal depth, extending nearly to the base of the enamel. The metaflexid progressively deepens posteriorly through the series until it is as deep as the hypoflexid and mesoflexid on M3. The walls of these three valleys are steep. In Mo.3, especially in the latter, these flexids deepen toward the mid-line. The anterofossettid is shallower than the principal flexids, and the anteroflexid is little more than a conspicuous groove. The mesoflexid and anteroflexid together constitute the largest valley in Mi.o. On M3 the connection between the posterolophid and the ectolophid had not yet been fully established by wear, which sug- gests that in the unworn state a gap existed between these lophids. On M3 a slight notch in the anterior wall of the anterofossettid had not yet been obliterated by wear. The base of the incisor is poorly preserved. All that can be said concerning it is that the enamel extends about half way around the lateral face. The mandibular ramus, so far as preserved, is robust and deep, about as in Deseadomys. As in that form, the ascending ramus arises near the front of M3. The masseteric crest begins, as a small shelf, at a point beneath the anterior portion of Mx. Paradelphomys1 gen. nov. Type species: Paradelphomys fissus1 sp. nov. Distribution: Colhuehuapian, Oligocene, Patagonia. Diagnosis: Lophids very oblique, posterolophid forming an iso- lated lamella; metalophid vestigial on dm4, absent on molars; anterofossettid of dmt elongate. Paradelphomys fissus sp. nov. Type: Museo municipal de Mar del Plata No. 125, fragment of right mandible with dm4 — Mx. 1 7rapa, beside, plus Adelphomys. The specific name alludes to the isola- tion of the posterolophid. 10 BREVIORA No. 301 Hypodigm: Type only. Horizon and locality: Colhue-Huapi Formation; barranca south of Lake Colhue-Huapi, Chubut. Argentina. Diagnosis: As for the genus. Description: Dm, is nearly oval in outline and longer than M,. The anterolophid is gently convex anteriorly and very obliquely directed posterolinguallv. Near its extremity it is connected to the hypolophid by a short bridge, a vestige o3714. collected by natives lor Fred Parker at Mutahi. Bougainville ( elevation 2200- 32(H) feet) on IS Ma\ 1966. Paratypes (7 specimens): NIC'/. 8761 I. 93710, 93712, Natur- historiska Riksmuseet (Stockholm) 67-0001, same data as holo- type. but collected in the period 10-19 Ma\ 1966. MCZ 93713, Melilup, Bougainville (elev. 3000 - 500 feet) on 14 Ma\ 1966. MCZ93711,Ramazon River, Bougainville (elev. L 600-2400 feet) on 19 May 1966. MCZ 93708, Turiboiru, Bougainville (elev. 500 feet) on 21 March 1966. Generic diagnosis: The genus is similar to an assemblage of lygosomine skinks inhabiting the Australian Region and subsaharan Africa. This assemblage mav be characterized by the following suite ol skull characters: palatal rami of pterygoids somewhat ex- panded and deeply emarginated posteriori) and separated mediall) bv medioposterior processes from the palatines, which project into the interpterygoid vacuity (beta palatal pattern, see p. 10 and Figure 4); 11-15 teeth on the premaxillae; no postorbital bone; SUpratemporal fenestra absent or minute; and Meckel's groove obliterated bv the overlapping and fusion of the dentary. I he genus Geomyersia mav be distinguished from all other ta\a in this assemblage (see p. II) bv its depressed body form, the complete absence o\' prefrontal scales, and the presence of a single median scale between the single pair of nuchal scales. All other taxa in the assemblage possess well-defined prefrontals and lack an "internuchal" scale. No other skink, in fact, is known to have such an internuchal scale as a normal feature (Figure 1 ). Species diagnosis: The species diagnosis is the same as that for the genus. Description (Figure 1): In general appearance Geomyersia glabra is a small (32-36 mm in snout-vent length), very dark, brownish black skink with a markedlv depressed body form. The 1 The genus is named for Professor George Sprague Myers of the Division of Systematic Bioloi:\. Stanford University, who stimulated the early interest of the senior author in systematic and evolutionary hiology. The specific name (glabra) refers to the extremely smooth appearance of the species. 1968 NEW SKINK FROM BOUGAINVILLE 3 limbs are pentadactyl and fail to meet when adpressed to the body. Head noticeably depressed; rostral about twice as wide as deep, forming an almost straight suture with the very large frontonasal; external naris centered in a rectangular nasal; a single loreal be- tween the nasal and two superposed preoculars; prefrontals entirely lacking; frontal short, as long as the medial longitudinal length of the fused interparietals, and forming a shallow, but broad, con- cave suture with the frontonasal; frontal in contact with the two anteriormost of the four supraoculars; lower eyelid movable with a large translucent spectacle; small scales of eyelid separated from supralabial scales by complete row of suboculars; 6-6 supra- labials, 4th supralabial below center of eye; frontoparietals fused into a single scale; interparietal distinct, shorter than medial longi- tudinal length of the single frontoparietal; parietals large, forming a suture approximately two-thirds the length of interparietal; each parietal bordered posteriorly by a single large nuchal and temporal; a small, median scale separates the pair of nuchals. Figure 1. Dorsal view of the head of the holotype of Geomyersia glabra (MCZ 93714). Notice the absence of prefrontal scales and the presence of the single frontoparietal and internuchal. Tympanum sunk below level of epidermis; external ear opening present but minute and without auricular lobes; body scales smooth, slightly iridescent, and disposed in 22-24 longitudinal rows around midbody; a median pair of enlarged preanal scales bordered laterally 4 BREVIORA No. 302 by two subcqual scales; median row of subcaudal scales transversely enlarged. Dorsal surface of digits covered by a single row of scales (Group 1 of Brongersma, 1942); 11-15 subdigital lamellae beneath 4th (longest) toe; lamellae folded into a median and two lateral ridges. (Dior: The dorsum and sides are dark grayish brown to brownish black; a median series of anteriorly-directed dark brownish black chevrons beszin on the base of the tail and extend forward to the nape; three or four horizontal streaks composed oi long dark dashes extend along the centers of the lateral scale rows between the fore and hind legs. Color does not appear to alter with preser- vation. The head is as dark as the dorsum, except for light cream stripes along the sutures between the supralabials. The venter is light with thickly clustered dark spots which become more confluent and dense posteriorly. The underside of the tail is so heavily spotted as to be almost uniformly dark. Distribution (Figure 2): The species is known only from middle elevations (500-3500 feet above sea level) in the highlands of northern and southern Bougainville. Reproduction: Geomyersia glabra is oviparous and apparently lays only a single egg at a time. Three o\ the seven individuals known were gra\id. MCZ 87611 and MCZ 93712 (snout-vent lengths 35.5 and 34 mm respectively) each contained 1 oblong. leathery shelled egg. MCZ 93714, with a snout-vent length of 33 mm. contained a single, large volkv egg in the oviduct. Ecological observations: (icomxersiu glabra has been collected between 500-3500 feet. Only one specimen (MCZ 93708) has any precise ecological information associated with it. This indi- vidual was found in decayed vegetable matter in secondary forest. All the specimens of G. glabra were brought in by native collectors along with large numbers of Sphenomorphus (especially solomonis and tanncri). Apparently the natives do not distinguish Geomyersia glabra from the small svmpatric Sphenomorphus (see Morphologi- eal Comparisons with Other Small Solomon Islands Skinks). The few (six) specimens of Geomyersia glabra obtained in the Mutahi-Melilup area, in proportion to the thousands of individuals of other species collected in the same area, indicate that the species is either extremely rare or is more or less restricted to a peculiar habitat that was not widely sampled during the overall collecting. 1968 NEW SKINK FROM BOUGAINVILLE •6°S 7°S 156° E 30 miles o 155 E SHORTLAND MONO I (^J> 6°S 156° E Figure 2. Bougainville and neighboring islands showing the known locali- ties from which Geomyersia glabra has been collected. The apparent rarity is not solely the result of size, as many juveniles of the small skinks are collected and the natives were offered in- centives to collect the species. (Interestingly enough the largest skink in the Mutahi-Melilup area, Sphenomorphus taylori, is as infrequently collected as is Geomyersia glabra.) Morphological Comparisons with Other Small Solomon Islands Skinks: No close relatives of the monotypic Geomyersia are known 6 breviora No. 302 to occur naturally in the Solomon Islands,1 although there are three small to medium sized skinks with a more or less uniform dark dorsal coloration in the Solomons with which the new species might be confused. Two of these three, Sphenomorphus solomonis and S. tanneri, occur together with G. glabra on Bougainville, while the third. S. bignelli, is known at present only from the southern Solo- mon Islands. The distinguishing features of the four species are outlined in Table 1. Discussion: Among lygosomine skinks there is a large group cen- tering on the genus Leiolopisma, as used in its broadest sense, that may be distinguished on the basis of ( I ) a single pair of nuchal scales behind the parietals. each o\' which is in contact with a single large temporal scale laterally, (2) the upper sides of the digits covered by a single row of scales (Group I of Brongersma, 1942), ( 3 ) the lower eyelid usually, but not invariably, with a clear spec- tacle, and (4) preanal scales often subequal or at least not greatly enlarged. Correlated with these external characters are the following skull characters: ( 1 ) palatal rami of pterygoids separated on the midline — often by medioposterior projections from the palatines into the interpterygoid vacuity. (2) II or more premaxillarv teeth. (3) a small or minute supratemporal fenestra. (4) no postorbital bone. and (5 ) Meckel's groove obliterated by the overlapping and fusion of the dentary. Within this large group, two subgroups may be recognized on the basis of the presence or absence of a medial, posteriorly pro- jecting process from the palatal ramus of the pterygoid and a concomitant emargination in the palatal ramus (Figures 3 and 4). One group, the alpha group for the purposes of this discus- sion, has the palatal rami of the pterygoids gradually and smoothly diverging postcriorlv from the midline (Figure 3). In the other ( arlia fusca, a close relati\c of Geomyersia glabra (see below), is known to occur in the Solomon Islands only on tiny Sohano Island in the strait be- tween Buka and Bougainville, on I-aisi Island and at Buin, Bougainville. The species is as yet unknown from any other localities in the Solomons, and it has almost certainly been introduced in the known localities. C. fusca is very common around the town of Rabaul, New Britain, the port from which Bougainville receives most of its supplies. The ecology of the species in Rabaul. the Solomons, and New Guinea is similar. It is a grass-dweller, pre- ferring the thick tussock grass of open areas. 1968 NEW SKINK FROM BOUGAINVILLE _l 00 < .5 «5 T3 C Si C C O CO o & > • i-h as ■*— » -*-J -> w 03 (ty ortl 0) o c j>^ (0 .c 3- CO eul, I da, M Oh co 5 CO 2 3° O "§ 03 O -O r- PnfflU^ Ti- ts ts ts ts £ o 73 C — 73 > "> D w I 3° CO £ 73 C 3 O 03 CO 13 O CO *J 73 03 i> Ofl O •-H -^H I5 CO w c o 03 Oh O 3 O >-H Uh u ?^ u u o H-l c o ■a, 3 X) Q 8 HRl.VIORA No. 302 Figure 3. The alpha palatal pattern (ventral view): (A) Mabiiya poly- tropi.s (MCZ B103), Kribi. Cameroon; (B) Leiolopisma me tallica (MCZ 67129), Mt. Toolbewong, Healesville, Victoria; (C) Emoia samoense (MCZ 16931 ), Fiji Islands; (D) Eumecia anchictac (MCZ 41562), Kaimosi, Kenya. Not drawn to scale. Abbreviations: E, ectopterygoid; P, palatine; PT, pterygoid. 968 NEW SKINK FROM BOUGAINVILLE Figure 4. The beta palatal pattern (ventral view): (A) Carlia bicari- nata (MCZ 64315), Port Moresby, New Guinea; (B) Leptosiaphos bloch- manni (MCZ untagged). Upper Mulinga, Idjwi Island, Democratic Republic of the Congo; (C) "Ablepharus" smithi (MCZ 42880, paratype), Nyonga, Democratic Republic of the Congo; (D) Geomyersia glabra (MCZ 87611, paratype), Mutahi, Bougainville. Not drawn to scale. Abbreviations as in Figure 3. 10 BREVIORA No. 302 group, here called the beta group,1 the palatal rami of the pterygoids have medial, posteriorly projecting processes set off bv emargina- tions lateral to them. In addition, whereas medial, posteriorly pro- jecting processes from the palatines are present only in some alphas, they are a constant feature of the beta palatal pattern (Figure 4). The functional significance of the deeply emarginated palatal ramus oi the pterygoid in the beta group (Figure 4) is not pcr- fectl) clear. The emarginatioa may serve as a "catch" for the basipterygoid process as the quadratal ramus o\' the pterygoid slides backward along the outer edge of the process. Such a movement on the part oi the quadratal ramus is evident in Creshl) prepared skulls when the forward part of the skull is depressed (hinging at the transverse frontal-parietal suture), and the whole palatal complex is displaced backward. In freshly prepared skulls, the basipterygoid process can catch in the emargination, stopping the posterior displacement o\' the palatal complex, thus limiting the amount o{ strain put on the skull. 1 he skull is probably capable o\' such movement during active feeding, but the differences in feeding behavior or habits that would make this "catch" mechanism an advantage to the skinks o{ the beta group are unknown. The taxonom) of the alpha and beta groups is extremely com- plex and will be dealt with in future papers. It is of interest for the present discussion, however, to list, using current generic concepts, those skinks. with their distributions, which belong to the alpha and beta groups. I he number of species in each genus in each area is given in parenthesis. I he beta skinks ma) be of mono- or diphvletic origin. Present evidence. largelj geographic, suggests to us th;it the betas of subsaharan Africa and of the Australian Region base probably independent!) originated from an alpha ancestry, i.e.. they are diphyletic. if so, use of the word "assemblage," which carries no connotations of close relationship for zoologists, would perhaps be more appropriate than the word "group," which does connote close rela- tionships, when discussing the beta skinks as a unit. In the diagnosis of Geomyersia the word "assemblage" was used for the beta skinks. hut in the following discussion, largeh as a matter of stylistic balance with the term "alpha gioup" (on present evidence the alphas are monoph\letic in origin), the term "beta group" will be used. It should be borne in mind, however, that the mono- or diphvletic origin of the beta skinks is still a problem needing further research. 1968 NEW SKINK FROM BOUGAINVILLE 11 alpha Ablepharus Australia (part, 11); Asia (8) Anotis New Caledonia (3) Didosaurus mauritianus, subfossil on Mauritius Emoia Indo-Australian archipel- ago, N. Australia, and Pacific islands (40) Eugongylus New Guinea, N. Aus- tralia, and Solomon Islands (5) Eumecia subsaharan Africa (1 or2) Leiolopisma Asia (17); North America (6); Indo-Australian archipelago (20); Australia (part, 8) ; Lord Howe Is. ( 1 ) ; New Cale- donia (6); New Zealand (16); Mauritius (1) Mabuya Central and South America; to Indo-Australian archipelago (66) Sphenomorphns New Guinea (mi- nutus); Solomon Islands (bignelli); New Zealand (pseudornatus) beta Ablepharus Africa (6); Australia (greyi and burnetii) Carlia Australian Region (21 ) Geomyersia Solomon Islands ( 1 ) Leiolopisma Africa (7); Australia {weeksae, challengeri, mustelina, guichenoti, delicata) Leptosiaphos Africa (7 Panaspis Africa (7) Within the beta group, the group to which Geomyersia belongs, relationships are complex and still incompletely worked out. For the moment, therefore, our purpose will only be to discuss some of the clearest groupings within the beta group and with this informa- tion assess the relationships of Geomyersia. (1) It is implicit in the list of alpha and beta skinks given above that the genus Ablepharus, characterized by "no movable eyelids, [and] a transparent disk covering the eye" (Boulenger, 1887), is not monophyletic; indeed, the genus is an assemblage of species, derived from diverse stocks, with a clear spectacle in the movable lower eyelid (Smith, 1935 and 1937; Greer, 1967a; Fuhn, in an unpublished discussion of the polyphyletic origin of Able- pharus). 12 BREVIORA No. 302 The Australian A. greyi and A. burnetii (beta group) arc the only "Ablepharus" with a reduced digital formula (4 fingers and 5 tecs), and significantly, they occur in the same zoogeographic region as the only other group of beta skinks with a similarly re- duced digital formula, i.e.. Carlia. The fused frontoparietal and spiny auricular lobules of A. burnetti are shared with Carlia and suggest that A. burnetii is simplv a Carlia in which the movable lower eyelid with its clear spectacle has fused almost completely1 to the upper edge o( the orbit. The 4-5 digital formula and fused frontoparietals of I. greyi also argue for the close relationship o\ this species with Carlia. but the unique supraocular arrangement found in A. greyi would sug- gest separate generic rank. The generic name Menetia Gray, 1845, originall) proposed for this species, is available. (2) I he other beta skinks in the Australian region, in addition to Carlia and its two ablepharine derivatives discussed above, are Geomyersia glabra and five species referred to Leiolopisma: week- sue, challenger i, mustelina, guichenoti, and delicata. In possessing 5 lingers these five species of Leiolopisma, as well as Geomyersia, displa) the primitive morphological condition from which the 4-lingered condition oi Carlia and its two ablepharine relatives must have been derived. Three of the 5 beta Australian Leiolopisma displa) yet another primitive character from which a more advanced character state displayed by all other beta Australian skinks must have evolved: Leiolopisma weeksae, chaUengeri and mustelina have paired fronto- parietals while all other Australian betas have fused frontoparie- tals.3 1 he onl\ other skinks with B reduced number of fingers ami iocs pre viously considered to be in the genus Ablepharus are six Australian species: elegans, distinguenda, orient alis, muelleri, timida, lineata. These species are now considered to be congeneric (genua Lerista) with the Australian skinks formerly referred to Rhodona (Greer. 1967a). and are only distantly related to any of the alpha or beta skinks discussed in this paper. In the single specimen of Ablepharus burnetii available to us (MCZ 6486 ) there is a small palpebral slit remaining along the dorsal edge of the lower eyelid. The e\e is effective!) "ablepharine," however, as this free dor- s.il edge is surely too short to allow much movement of the eyelid. 'Geomyersia grabra may be distinguished from the 2 beta Australian Leiolopisma with fused frontoparietals and 5 fingers {guichtnoti and delicata) by the complete absence of prefrontal scales and the depressed body form. 1968 NEW SKINK FROM BOUGAINVILLE 13 These 3 primitive beta Australian Leiolopisma cannot, at this stage of the study, be distinguished morphologically as a group from the African beta Leiolopisma, all of which have 5 fingers and separated frontoparietals. This similarity could be the result of either convergence or close relationship. For the present the widely disjunct distribution of the two groups implies to us that the groups have arisen independently in each area, but, admittedly, this is an unsolved problem and will be dealt with elsewhere. (3) Taken as a group the African betas are themselves taxo- nomically difficult at an infragroup level. Several characters might be used to distinguish taxa within the African betas: the presence of supranasals, presumably a primitive condition, serves to diagnose Panaspis; a scaly lower eyelid, minute ear opening and a group tendency toward an elongate body and reduced number of digits distinguishes Leptosiaphos; the ablepharine eye and reduced num- ber of supraocular scales (3 instead of 4) defines the African "Ablepharus," while an African group of Leiolopisma could be "diagnosed" negatively on the basis of showing none of these char- acters. Certain species, however, bridge these apparent taxonomic gaps. For example, Panaspis cabindae may have 3 instead of 4 supraoculars (Parker, 1936), and in P. breviceps the supranasal may be fused to the nasal (Smith, 1937) ; in addition, Leptosiaphos kilimense and L. rhodurus are atypical of other Leptosiaphos in displaying a transparent window in the movable lower eyelid. Such difficulties suggest that on the basis of these characters, it might be better to resort to, at most, a subgeneric scheme of ranking taxa. (4) On the basis of geography, Geomyersia glabra would seem more closely related to Australian betas than to African betas. In addition the fused frontoparietals of G. glabra may be a morpho- logical indication of relationship with the Australian betas which possess this advanced character in 26 of the 29 species. The Afri- can betas have divided frontoparietals, presumably the more primi- tive condition, in all but one of the 27 species. Only in the African beta Ablepharus wahlbergi are the frontoparietals normally fused,1 a condition probably derived independently from that of the Aus- tralian betas with fused frontoparietals. 1 The fusion of the frontoparietals and their fusion with the interparietal is a variable condition in A. wahlbergi (personal observation and de Witte, 1953). 14 hreviora No. 302 Geomyersia glabra docs, however, have 5 fingers — a character which is predominant in the African betas and which might, there- tore, be thought to indicate greater likelihood of relationship with African rather than Australian beta skinks. However. 5 fingers is the primitive condition for both the African and Australian betas, and although in the Australian betas the 4 fingered species are a conspicuous element in that group's radiation, the 5 lingered G. glabra can be readily interpreted as a relic of an early stage in the evolutionary history of Australian betas. It would presumably represent a stage later than that of Leiolopisma weeksae, /.. chal- lengeri and /.. mustelina and approximating that of I., guichenoti and /.. delicata, but with specializations of its own. Certainly the island of Bougainville on the periphery of the Australian region would be a logical refuge in which to find a relic of an early stage of a radiation taking place in this region. A dendrogram of the Australian beta skinks. assuming on the basis of present evidence — mainl) geographic — that they are monophyletic, is presented in figure 5. I he radiations of the Australian and African beta skinks will be discussed further in forthcoming papers. ( mparisons with Morphologically Similar Species: It is desir- able to mention brief!) those species in the Australian Region that are confusing!) similar to Geomyersia glabra on the basis of exter- nal morphology. Only those skinks displaying the external charac- ters of the alpha and beta skinks are discussed. Leiolopisma mccooyi (Lucas and frost. 1894) from Victoria lacks prefrontals and has a minute external ear opening as does G. glabra, but the frontoparietals of mccooyi are divided, the body is not depressed, and the palate is of definitely alpha type. / ygosoma graciloides Lonnberg and Anderson, 1913 (=L\go- soma scharffi Boulenger, 1915) has been collected twice and is known from 4 specimens. Although described twice under the genus / ■/// 1 3 ) and Grouvelle ( 1919) placed it in C"r\ptophagidac. and Arrow | 1 l>2c>a ) proposed that Hapalips, together with all Cryptophagidae having lobed tarsi. be plaeed in Languriidae. Rymer Roberts | 1939) placed Hapalips with Boleriu (=Platycladoxena) angulosus in Languriidae. Cla- doxeninae, alter studying larvae oi the two genera. Bruce (1951) noted that the aedeagus of Hapalips is similar to those oi Leucohimatium, Loberus, and Toramus. Crowson (1955) re- tained Hapalips in the Languriidae and considered it as a tran- sitional form between that family and Erotylidae. This is the second largest genus of the tribe Loberini. Reitter (1877) de- scribed eight species. Schenkling ( 1923) listed 47, to which Arrow (1927) added another one, and more recently, Bruce (1952, 1963) added five more species and excluded one of his older species. //. spegazzini, because of its different type of aedeagus. Thus the genus at present consists of 54 valid species. There ap- pears to have been no previous designation of a type for this genus, so I here designate as type H. mexicanus, one of the original species described by Reitter ( 1877) and quite common in Mexico. 1968 GENERA OF LOBERINI 7 Description. With general characters of Loberinae, Loberini. General appearance more or less as in Xenoscelis Wollaston; form narrow, elongate, rather flattened, and more or less parallel sided. Head (Fig. 23) with broad clypeus, its front margin evenly rounded; antennal insertions completely hidden by the sides of frons. Stridulatory files rarely present and, if so, paired and well separated; sometimes files represented by a pair of longitudinal ridges without any striations (e.g. H. juscus and H. acaciae). Transverse groove on anterior part of gular region sometimes re- placed by transverse cavity (Fig. 23). Antenna of moderate length, scape small but longer than pedicel, which is often shorter than segment 3, segments 4-8 usually equal in length, segments 9 and 10 transverse and equal in size, terminal segment equal in breadth to segment 10 and more or less rounded at apex. Pro- thorax (Fig. 24) usually elongate with side margins smooth, more or less parallel sided, front and hind margins equal in width, all the angles rather obtuse; prebasal impressions on pronotum often indistinct. Front coxal cavities usually with narrow, sometimes slitlike (Fig. 24) opening behind; prosternal process with its apical margin straight. Sometimes shape of pronotum differs between male and female. Elytra with regular rows of strial punctures, usually with a scutellary striole, and pubescence absent or fine and recumbent. Wing with (Fig. 18) or without (Fig. 16) anal cell, radial cell rarely without spur of radial sector (Fig. 17). Meso- coxae closely situated; mesepisternal pockets often obscured; meta- sternum elongate or as broad as long; median impressed line of metasternum usually extending % of its length. Tarsi short and compact, first three segments rather broad and equal in length, segment 3 lobed below, segment 4 minute, received in the lobe of segment 3, segment 5 about equal in length to first three together; tibiae rather short, broad at apex, which is obliquely truncate with two normal spurs. Ventrite 1 with intercoxal process pointed at apex. Ovipositor and aedeagus as figured (Figs. 12, 10). Species examined. The species H. mexicanus Reitter, H. eichel- baumi Grouvelle, H. cribricollis Gorham, H. grouvellei Gorham, H. nigriceps Reitter, H. nitidulus Champion, H. filum Reitter, H. scotti Grouvelle, H. prolixus Sharp, H. juscus Reitter, H. acaciae sp. n., and also H. taprobanae Grouvelle (external characters only), have been studied in detail for generic characters. Habitat. Little is known about the habitat of members of this genus. H. filum has been recorded from Cuba in corn (maize) stalks, and H. annulosus Grouvelle from Guadeloupe in flowers of the cactus Cereus triangularis. H. prolixus has been recorded from 8 breviora No. 303 New Zealand in tree ferns, and more recently R. A. Crowson col- lected both larvae and adults of this species under the sheathing bases of dead palm leaves. //. championi Grouvelle and //. scotti Grouvelle are recorded from the Seychelles, both from the bases of palm leaves. Geographical distribution. This genus is widely distributed in both the New and Old Worlds, mainly in tropical and subtropical regions; a few species extend into warm temperate areas. No spe- cies are yet recorded from Australia, and onlv one species. H. proli.xus, is known from New Zealand. Subgenus Hapalips \cnsit str'n to Description. With the general characters of Hapalips. Form more convex, less parallel-sided; elytra with scutellarv striole; wing (Fig. 18) with anal cell and radial cell with spur of Rs. Head sometimes with a pair o( stridulatoiy tiles; a transverse groove pres- ent on gular region | Fig. 20); eyes large, facets moderately coarse. Pronotum more coin ex. length and breadth more or less equal. Mctasternum not elongate, more or less equal in length and breadth. This subgenus includes the majority of the species. ( \\ OPHORUS subgenus novum Type species. Hapalips fuscus Reitter Description. With the general characters oi Hapalips. General appearance as in Hapalips sensu stricto. Head with vestiges of Stridulatory tiles, represented by two longitudinal ridges without striatum (Fig. 23). On the gular region, in place of the usual transverse line, there is a large trilobed cavity (Fig. 23). opening ventrally and protected by hairs. Eyes large, as in Hapalips sensu stricto. Elytra, wing venation, shape of pronotum and mctasternum as in Hapalips sensu strict". According to present knowledge, this subgenus includes only two Central American species, H. fuscus Reitter and //. acaciae sp. n. Xenosceloides subgenus novum Type species. Hapalips proli.xus Sharp Description. With the general characters of Hapalips. Form more flattened and parallel-sided. Wing without anal ceil (1 16), radial cell sometimes without spur of Rs (Fig. 17). Elytra 1968 GENERA OF LOBERINI 9 often without stridulatory files; gular region with a transverse groove as in Hapalips sensu stricto. This subgenus includes Hapalips prolixus Sharp, H. fdum Reit- ter, H. scotti Grouvelle, and possibly H. taprobanae Grouvelle. I have not been able to check in detail the characters of H. taproba- nae, which is less flattened than the other species; if it truly belongs in this subgenus, then Grouvelle's name Loberina will have priority over Xenosceloides. Key to the Subgenera of Hapalips 1. Anterior part of gular region with a large cavity (Fig. 23) . .Cavophorus Anterior part of gular region without cavity but with a transverse groove (Fig. 20) 2 2. Wing with anal cell (Fig. 18); body shape more convex and less parallel sided; elytra with scutellary striole. Prothorax and metasternum more or less equal in length and breadth. Stridulatory files on head sometimes present; eyes large with moderately coarse facets Hapalips Wing without anal cell (Fig. 16); body shape more flattened and parallel sided; elytra often without scutellary striole. Prosternum (Fig. 24) and metasternum longer than broad. Stridulatory files on head absent; eyes smaller with finer facets Xenosceloides Hapalips acaciae species novum Holotype, 9 , and para type $ , Cotaxtla Exp. Sta., Veracruz, MEXICO, VIII- 15-1 962, D. H. Janzen coll., seed pods of Acacia cornigera, deposited in the Museum of Comparative Zoology, Cambridge, Mass. (holotype MCZ No. 31604). Paratype (on slide), same data, deposited in the Department of Zoology, The University, Glasgow, Scotland. Measurements of holotype. Total length: 5.58 mm; width of head across eyes: 1.18 mm; length of antenna: 1.26 mm; length of prothorax along midline: 1.23 mm, width across middle: 1.40 mm; length of elytra: 3.60 mm, width across middle: 1.45 mm. Description. With the general characters of Loberini, Hapalips (Cavophorus). Larger and more elongate than H. fuscus Reitter; upper surface uniformly reddish brown. Head (Fig. 23) trans- verse; eyes large and rather coarsely faceted. Vestiges of stridula- tory files represented by a pair of widely separated longitudinal ridges without transverse striations. Occipital region glabrous, an- terior half of dorsal side of head sparsely clothed with recumbent setae. A large transverse trilobed cavity (Fig. 23) present on anterior part of gular region, opening ventrally and protected by 10 breviora No. 303 fine hairs. Antenna with scape about double the length of pedicel, which is shorter than segment 3. segments 4-8 equal in length. segments 9 and 10 roughly semicircular, segment 9 slightly less wide than segment 10. terminal segment asymmetrically rounded in apical part and longer than preceding two segments. Prothorax slightly wider than its length, side margins more or less parallel sided, bordered by fine hairs, posterior angles pointed and an- terior angles obtuse. Punctation on pronotum coarser and closer than on head, setae minute and directed toward center. Elytra uniformly reddish brown, stria! punctures in rows, interstices with sparse fine punctures, scutellarv striole rather irregular and indis- tinct, pubescence short and recumbent. Scutellum minute, nar- rowed in front and weakly angulate on posterior margin, pubescent. Wing with anal cell. Mesocoxal lines present but very short; mesepisternal pockets obscured. Tarsi densely hairy on underside, first three tarsal segments more or less equal in length, segment 5 about as long as first three segments together. Femoral lines on ventrite 1 very short; intercoxal process narrow and pointed. Key to the Species of Cavophorus l. Species larger and more elongate, ahum 5.80 nun in length; prothorax more or less equal in length and hreadth. Femoral lines on ventrite 1 very short //.(<-.) acaciae sp. n. Species smaller and less elongate, aboul 3.80 mm in length; prothorax distinct]) transverse. Femoral lines on ventrite l fairly long //. (<) fuscus Reittei Description of a Larva op Hapalips prolixus Sharp One larva recorded from New Zealand under the base of dead palm leaves {Rhopalostylis sapidd) along with the adults, by R. A. Crowson, deposited in the Glasgow University Zoolog\ Depart- ment. Total length 3.48 mm; length of head including labrum 0.64 mm. width across the middle 0.56 mm; width of prothorax across the middle 0.64 mm; width of 9th abdominal segment across the front margin 0.48 mm. General appearance narrow, elongated, somewhat flattened. tapered in front and behind. Dorsal surfaces bearing many gran- ules symmetrically arranged in a definite pattern of lines, except on pronotum and head, wartlike setiferous tubercles on cither side of each tergite. setae near middle line minute and blunt, becoming 1968 GENERA OF LOBERINI 11 longer towards sides. Setae on ventral surface rather short, fine and pointed. Head rather elongate, shape of head and arrangement of setae on dorsal surface as figured (Fig. 27), dorsal surface granulated. Frontal suture rather indistinct, as figured (Fig. 27); endocarina absent. At the base on either side of occipital foramen there are three microscopic peglike setae. Ocelli not distinct. Antennae short, narrow, length of the segments 1:2:2, sensory appendage lying ventrally, % of the length of segment 3. Mandible (Fig. 29) with two equal teeth, mola well developed bearing transverse ridges and asperites. Between the apical teeth and mola the pros- theca translucent and pointed at apex. Ventral crushing tubercle well developed; single sensory pit present in the middle of dorsal side of mandible, another one on ventral side anterior to dorsal one. Maxillary mala (Fig. 31) acute at apex, with three apical spines as in Hapalips sp. larva described by Rymer Roberts (1939), a row of seven setae present on dorsal side of inner mar- gin of mala; at the base of these setae on dorsal side a group of small denticles, similar denticles present at the base of palpiger (Fig. 31). Cardo rather long, narrow, at right angle to stipes; maxillary articulating area well developed and oval. Labium (Fig. 32) with two jointed palpi, palpiger not distinguishable; ligula rather indistinct; hypopharynx with well developed hypopharyngeal bracon. Pronotum slightly wider than head; granulation on dorsal sur- face irregular. Meso- and metathorax slightly shorter and progres- sively wider than prothorax; granulation on dorsal side in a regular symmetrical pattern as in abdominal tergites (Fig. 33). Abdomi- nal segments 1-6 equal in length and breadth and equal to meta- sternum, segments 7-9 progressively narrower. Arrangement of setae and granulation are similar on meso- and metanotum and first 8 abdominal tergites, on tergite 9 granules and setae are ar- ranged as in Figure 33. Each segment with two transverse rows of four minute blunt setae; two pairs of tubercles on either side of each segment, anterior pair carrying two blunt and comparatively short setae, posterior pair with single long and pointed seta (Fig. 33). Urogomphi well developed, as figured (Fig. 33), not hooked or upturned, projecting posteriorly. A pair of setiferous tubercles (pregomphal process) present anterior to urogomphi. Pygopod small, rounded, and not projecting. All spiracles are bicameral (Fig. 34), lying on body surface, lateral air tubes directed posterolaterally. Legs fairly long, coxae closely situated, claws simple with two tarsungular setae. 12 breviora No. 303 This larva may be distinguished from that of Hapalips sp. de- scribed by Rymer Roberts ( 1939) by the following key: Urogomphi long, not hooked (Fig. 33). Mandibular prostheca not narrow or hooked at apex (Fig. 29). Ocelli obscured. Tubercles on each segment with pointed and blunt setae (Fig. 33) .... H. prolixus Sharp Urogomphi short, hooked. Mandibular prostheca rather narrow and hooked at apex. Ocelli 5 posterior to antenna. Tubercles on each segment with single long pointed seta Hapalips sp. Discussion. Although all the species of Hapalips are restricted to the warmer climatic zones, the two larger subgenera are repre- sented in both the New and Old Worlds; even within these sub- genera there are no obvious general differences between the New and Old World forms. The subgenus Cavophorus, with only two known species, both restricted to the New World, appears to repre- sent a specialized development from Hapalips sensu stricto, at least in respect to the vestigial stridulatorv files and the big cavity in the anterior part o( the gular region. apparentl\ developed from the transverse groove which is present in other Hapalips in the same position. The subgenus Xenosceloides is another group which may represent a specialized off-shoot o[ Hapalips sensu stricto; the main differences between Xenosceloides and Hapalips sensu stricto parallel those between the genus Hemipeplus Berth and its relatives in the Mvclcridac ( Heteromera ). Typical Hemi- peplus spp. as far as known occur under leaf bases of palms, as do Xenosceloides adults. I hus the distinguishing features ol Xenosce- loides ma) be adaptive to this mode of life and may have originated independently in the Old and New Worlds, in which case Xenosce- loides would not be a natural subgenus. Further research will be needed to establish whether this is in fact the case. The only spe- cificallv identified larva of this genus is that of //. prolixus from New Zealand, which shows considerable differences, as well as similarities, when compared with the larva of Hapalips sp. de- scribed bv Rymer Roberts (1939) from Mexico; the differences are such that the two larvae would be expected to represent at least different subgenera. If the Mexican larva is really of a Hapalips sensu stricto, then larval characters support the separa- tion of Hapalips sensu stricto and Xenosceloides as subgenera; if, on the other hand, the Mexican larva proves to be of a Xenosce- loides, this will strongly support the theory that the New and Old World forms of the subgenus are not really related. 1968 GENERA OF LOBERINI 13 Genus Truquiella Champion Truquiella Champion, 1913: 87. Type species, by monotypy, Truquiella gibbifera Champion. This monotypic genus was established by Champion (1913) un- der Cryptophagidae; I have found no subsequent references to it. Champion described it as having tetramerous tarsi, but I found its tarsi to be pseudotetramerous, as in other Loberini. The genus seems to be very closely related to Hapalips. Description. With general characters of Loberinae, Loberini. General fades more or less as in Hapalips. Head with fairly large and coarsely faceted eyes. Stridulatory files apparently absent. A pair of protuberances present on anterodorsal side of eyes (Fig. 3) in both sexes, more prominent in male; clypeus broad with rounded apical margin as in Hapalips; antennal insertions com- pletely hidden by frons. Antenna moderately long, with scape small but larger than pedicel, which is shorter than segment 3; segments 4-8 equal in length and shorter than segment 3; segments 9 and 10 weakly transverse, terminal one elongate with rounded apex. Prothorax weakly transverse, side margins smooth, front and hind margins more or less equal in breadth, front angles slightly projecting forward and weakly acute, hind angles obtuse. Prebasal impressions on pronotum obscured; front coxal cavities rather narrowly open behind; prosternal process narrow and truncated at apex. Elytra with regular rows of punctures, scutellary striole present. Wing with closed anal cell, venation as in Hapa- lips eichelbaumi Grouvelle (Fig. 18). Mesocoxae closely situated; mesepisternal pockets weakly developed; metasternum weakly transverse; median impressed line extending % of its length. Tarsi with first three segments equal in length, segment 3 lobed below, segment 4 minute and segment 5 about as long as first four together; tibiae weakly broadened at apex with two normal spurs. Intercoxal process of ventrite 1 narrow and pointed at apex. Habitat. Unknown, larva undescribed. Geographical distribution. Mexico. Genus Pseudhapalips Champion Pseudhapalips Champion, 1913: 112. Type species, by monotypy, Pseud- hapalips lamellifer Champion. Champion (1913) established this genus under Cryptophagidae, and described it as closely related to Hapalips, although he noted several dissimilarities from Hapalips, e.g. the extraordinary form 14 BREVIORA No. 303 of head, very prominent eyes and thorax, as in Platoberus, etc. He described the front coxal cavities as closed behind, but careful study of a slide preparation reveals that these cavities are distinctly open behind ( 1 ig. 6). Arrow (1929a) described Pseudhapalips as having stridulatory files on the head, but neither I nor Mr. R. D. Pope of the British Museum could find stridulatory tiles on the head of P. lamellijer. It seems that the genus may be related to Hapalips, as suggested by Champion and Arrow, but not very closely. Since Champion's description only one species has been added to this genus, and that was by Grouvelle (1919) from French Guiana. Description. With general characters of Loberinae, Loberini. I lead transverse, in male with a strong ridge between the eves; in female this ridge is less distinct and Battened in front; ebpeus broad with rounded apical margin. Eyes large, markedly project- ing and moderatel) coarsely faceted. Antenna moderately long with scape slightly longer than pedicel, which is slightly shorter than segment 3. segments 3-8 equal in length, club loose, segment 9 very slightly wider than segment 10. terminal segment rather transverse and rounded at ape\. Prothorax (Fig. 6) strongly transverse, slightly narrowed be- hind, side margins weakl) undulate or dentate, front angles slightly projecting and obtuse, hind angles acute. Prebasal impressions on pronotum strongl\ marked; prosternal process weakly broadened posteriori) with straight apical margin (Fig. 6). Elytra glabrous, strial punctures in regular rows and with a scutellary striole. Wing as figured (Fig. 15), without anal cell. Mesepistcrnal pockets weakly developed; metasternum elongate and median impressed line extending to % of its length. Tarsi are unlike Hapalips, first three segments lobed below, segment 5 equal to length of first two segments together; tibiae broadened at apex. \ entrite 1 with inter- coxal process narrow and pointed at apex Habitat. Unknown, larva undescribed. Geographical distribution. Amazon. Surinam, and French Guiana. Genus Pseudhenoticus Sharp Pseudhenoticus Slmrp, 1900: 596. Type species, by monotypy, Pseudheno- ticus parallelus Sharp. Sharp ( 1900) established this genus under Cryptophagidae and placed it just before Henoticus, Grouvelle (1919) added 10 spe- cies to it, and more recently. Bruce (1943) described another 1968 GENERA OF LOBERINI 15 species from Madagascar. From Bruce's figure and description it seems very doubtful whether this specimen is a true Pseudhenoticus or even a member of Loberini; if it is a Pseudhenoticus, then the genus will manifest a distribution pattern unusual in Clavicornia, though it is known in some other organisms. Description. With general characters of Loberiinae, Loberini. Head transverse, clypeus less broad than in Hapalips, broad at base, narrowed in front, its apical margin rounded. Unlike Hapa- lips, frons scarcely projecting over antennal insertions. Stridulatory files apparently absent; eyes of moderate size and very finely faceted. Antenna rather short and stout, scape, pedicel and seg- ment 3 more or less equal in length, segments 4-8 slightly shorter than segment 3 and equal in size, segments 9 and 10 somewhat semicircular, and terminal segment elongated, about double the length of segment 10. Prothorax (Fig. 5) transverse, weakly nar- rowed in front, side margins undulate or more or less dentate, pre- basal impressions on pronotum present. Front coxal cavities rather narrowly open behind; prosternal process broad at apex as in Bolerus (cf. Fig. 2). Elytra with regular rows of punctures and without scutellary striole, pubescent. Meso- and metacoxae mod- erately widely separated, sternal fitting between the mesocoxae in a straight line. Metasternum transverse, narrowed in front, median impressed line extending % of its length. First three tarsal seg- ments more or less equal in length, segment 2 weakly and segment 3 strongly lobed below and narrow, segment 5 almost equal in length to first three segments together; tibiae slightly broadened at apex. Ventrite 1 with intercoxal process moderately broad and its apical margin rounded. Habitat. Unknown, larva undescribed. Geographical distribution. Panama, Bolivia,? Madagascar. Genus Bolerus Grouvelle Bolerus Grouvelle, 1919: 93. Type species, by present designation, Crotchia minuta Fleutiaux, 1887: 68. Thallisellodes Arrow, 1925: 257. Type species, by original designation, Thallis transversus Gorham, 1895: 325. Platycladoxena Kraatz (partim), — Arrow, 1929a: 316. Crotchia Fowler (partim), — Fleutiaux, 1887: 68. The nomenclature of this genus is very complicated. The genus Platycladoxena (type P. castanea Kraatz) was established by Kraatz (1899), in the Languriidae. Arrow (1925) synonymized it with Microlanguria Lewis and erected a new genus Thallisellodes with four species. He later (1929a) stated that P. castanea and 16 BREVIORA No. 303 P . javanica of Kraatz are congeneric with Thallisellodes and sank the latter name. In the same paper, he listed 13 valid species in this genus. Grouvelle (1919) described the genus Bolerus, which he considered to be related to Hapalips and Loberus, and trans- ferred Crotchia minuta Fleuiiaux to the genus Boletus. Arrow ( 1929a) pointed out that the species of Bolerus are congeneric with Thallisellodes. \ illiers ( 1961 ). having seen the type material of Platycladoxena castanea Kraal/, stated that the two specimens represented two distinct species, both of the genus Microlanguria; the lectotype should be named Microlanguria castanea (Kraatz), while the other specimen was described as M. angulosa Villiers. The above facts indicate that the genus Bolerus Grouvelle ( 1919) must have priority over Thallisellodes Arrow (1925). Unfortu- nately. Grouvelle did not specify a type o\ his genus Bolerus; there- fore I here designate Crotchia minuta Fleutiaux as type species. I Ins is one of the species originally included b\ (irouvelle ( 1919). The species Bolerus minimis, which 1 have studied in detail, seems to be \er\ similar to Thallisellodes angulosus Arrow and T. transversus (Gorham). I have found its characters more similar to those of the Loberini than to those of Cladoxenini (e.g. front coxal cavities internally open behind and ovipositor like other Loberini with st\li attached at the apex of coxites). Rymer Rob- erts (1939) described the larva of Bolerus ( -.Platycladoxena) angulosus Arrow and stated that it is very similar to the larva of Hapalips. The present study supports the view of Grouvelle and Rymer Roberts and shows that the genus is actually related to Hapalips and probabl) Pseudhenoticus. Description. With the general characters of Loberinae, Lobe- rini. General appearance less linear than in Microlanguria and prothorax Strongl) transverse. Head transverse, with a pair of closely situated stndulatory files; eyes fairly large and moderately coarsely faceted. Clypeus broad at base and narrowed in front, with straight or weakly rounded apical margin; antennal insertions less hidden under Irons than in Hapalips. Transverse groove on anterior gular region weak but distinguishable. Antenna with scape and segment 3 longer than pedicel, club rather compact and segments 9. 10. and 1 1 transverse and equal in length, apical seg- ment rounded at apex and slightly less transverse than segment 10, sometimes club loose and apical segment elongate and pointed at apex. Prothorax (Fig. 2) parallel sided, side margins more or less smooth or finely dentate, front angles rather rounded, hind angles somewhat acute, prebasal impressions on pronotum strongly 1968 GENERA OF LOBERINI 17 marked. Front coxal cavities rather narrowly open behind, pro- sternal process as figured (Fig. 2), sometimes sinuate at apical margin. Elytra glabrous, strial punctures in regular rows and with scutellary stride. Wing as figured (Fig. 19), with an anal cell. Mesocoxae moderately widely separated; mesepisternal pockets well developed; metasternum weakly transverse; mesocoxal lines short; median impressed line extending about % of its length. Metendosternite broad and short, as figured (Fig. 21). Tarsi as in Hapalips; tibiae broad at apex and obliquely truncated. Ventrite 1 with intercoxal process slightly broad, shape as figured (Fig. 1); sometimes a second pair of lines present on outer sides of the nor- mal femora] lines (Fig. 1 ). Habitat. Adults and larvae of B. angulosus have been found on a lichen-covered rock in a cave in Malaya. Geographical distribution. Indo-Malayan region. Key to the Genera of the Tribe Loberini 1. Trochanters short and broad; tarsi simple, except in Xenoscelis, where segment 3 is very slightly lobed below. Femoral lines on first ventrite absent; anterior part of gular region without transverse groove. Front coxal cavities usually narrowly open, rarely completely closed behind {Xenoscelis) Tribe Pharaxonothini ■ — trochanters broadly elongate (Fig. 26); tarsi distinctly lobed below, with minute segment 4. Femoral lines on first ventrite present; anterior part of gular region with a transverse groove (Fig. 20) or a cavity (Fig. 23). Front coxal cavities usually fairly widely, more rarely narrowly open behind Tribe Loberini .... 2 2. Mesocoxae more widely separated and sternal fitting between them in a straight line. Prothorax (Fig. 5) with anterior part of side margins undulated. Intercoxal process of first ventrite broad with more or less rounded apical margin Pseudhenoticus — Mesocoxae more closely situated and sternal fitting between them with single knob (Fig. 9). Prothorax not as above. Intercoxal process of first ventrite narrow and pointed at apex (except in Bolerus) 3 3. Prothorax as figured (Fig. 2), with prosternal process broad at apex. Elytra glabrous; mesocoxal line on metasternum present. Intercoxal process of first ventrite broad, as figured (Fig. 1 ) Bolerus — • Prothorax with prosternal process not as above. Elytra usually hairy; mesocoxal lines on metasternum absent. Intercoxal process of first ventrite narrow and pointed 4 18 breviora No. 303 4. Species larger, narrow and elongated, less Cryptophagidae-like in form. Elytra with scutellary striole (except in Hapalips scotti Grouvelle); uing often with anal cell (Fig. 18). Tibiae broad and truncate at apex; tarsal lobes broad. Metastcrnum less transverse 5 — Species smaller, more elliptical and Cryptophagidae-like in form. Elytra without scutellary striole; wing without anal cell. Tibiae slender, not broadened at apex; tarsal lobes narrow. Metasternum more trans- verse 7 5. Prothorax as figured (Fig. 6). side margins dentate or undulate, front angles projecting forward. Tarsal segments 2 and 3 lobed below. Anterior part of dorsal side of the head with transverse ridge .... Pseudhapalips — Prothorax not as above, side margin not dentate or undulate, front angles not projecting forward. Only tarsal segment 3 lobed below. WuAd not as abo\e 6 I- Head with a pair of humps on anlerodorsal sides of eves dig. 3). Pre- basal impressions on pronotum indistinct. I lytra pubescent. Truquiella — Head without humps .is above. Prebasal impressions on pronotum iisualh distinct Elytra glabrous or pubescent Hapalips 1. Prothorax narrowed in front, shape as figured (Fig. 7). Antenna with segmenl 9 considerably smaller than segment 10, which is markedly broad Telmatoscius — Prothorax not narrowed in front, shape as figured (big. 7). Antennal segment l> \er\ little smaller than segment 10. which is less trans- verse Loberus ACKNOWLEDGMENTS This research was carried out in the Department of Zoology, University of Glasgow. I am grateful to Prof. D. R. Newth for laboratory facilities. I am deeply indebted to Dr. R. A. Crowson of Glasgow University for his valuable advice and also for the larva oi Hapalips prolixus, and to Dr. J. F. Lawrence of the Mu- seum of Comparative Zoolog) for reading the manuscript and for the adults of Hapalips acaciae. I would like to thank the stall oi the Coleoptera section of the British Museum (Natural History), particularly Mr. R. D. Pope, who have made available to me the facilities to study material in the Museum collection. 1968 GENERA OF LOBERINI 19 LITERATURE CITED Arrow, G. J. 1925. Fauna of British India, including Ceylon and Burma. Coleop- tera: Clavicornia: Erotylidae, Languriidae and Endomychidae. London, Taylor and Francis, viii + 416 pp. 1927. Clavicornia and Lamellicornia. In: Insects of Samoa. Part 4, Coleoptera. Brit. Mus. (Nat. Hist.), pp. 35-66. 1929a. On the families of Coleoptera related to the Erotylidae, with descriptions of a new family, two new genera and a few new species. Ann. Mag. Nat. Hist., (10) 4: 305-322. 1929b. A revision of the African Coleoptera belonging to the family Languriidae. Proc. Zool. Soc. London, 1929, Part 1: 1-15. Batra, L. R. 1963. Ecology of ambrosia fungi and their dissemination by beetles. Trans. Kansas Acad. Sci., 66(2): 213-235. Bruce, N. 1940. Coleoptera — Cryptophagidae von Juan Fernandez. In: C. Skotsberg, ed., The Natural History of Juan Fernandez and Easter Island, Vol. Ill, Zoology: 682-688. 1943. Drei neue tropische Cryptophagiden nebst Bemerkungen zur Synonymie der Cryptophagiden Familie (Coleoptera: Crypto- phagidae). Arb. Morph. Taxon. Ent. Berlin-Dahlem, 10: 56- 61. 1951. Cryptophagidae (Coleoptera: Polyphaga). Explor. Pare Nat. Albert. Mission G. F. De Witte (1933-1935). Fasc. 75, 26 pp. 1952. Revision der im Deutschen Entomologischen Institut befindli- chen Hapalips-Arten. Beitr. Z. Ent., 2: 461-473. 1963. Coleoptera-Cryptophagidae in Musee Royal de l'Afrique Cen- tral (V). Rev. Zool. Bot. Afr., 47: 203-220. Champion, G. C. 1913. Notes on various Central American Coleoptera with descriptions of new genera and species. Trans. Ent. Soc. London, 1913, Parti: 87-113. Crowson, R. A. 1955. The Natural Classification of the Families of Coleoptera. Lon- don, Nathaniel Lloyd, 187 pp. Fleutiaux, E. 1887. Descriptions des coleopteres nouveaux de l'Annam. . . . Ann. Soc. Ent. France, Ser. 6, 7: 59-68. Fowler, W. W. 1908. Languriinae. In: P. Wytsman, Genera Insectorum, 78: 1-45. Gorham, H. S. 1895. List of the Coleoptera in the collection of H. E. Andrews, Esq., from India and Burma, with descriptions of new species and notes. Families: Malacodermata — Erotylidae — Endomy- chidae. Ann. Soc. Ent. Belgique, 39: 293-330. 20 hreviora No. 303 1898. Coccinellidae (part) and supplement to Erotylidae (part). In: F. Godman and O. Salvin, eds., Biologia Centrali-Americana. Insects. Coleoptera. Vol. 7, pp. 241-256. Gro< Mill. V ll>14. The Percy Sladen Trust Expedition to the Indian Ocean. Coleoptera: Cucujidae, Cryptophagidae. Trans. I. inn. Soc. London: Zool., 17: 141-159.' 1919. Descriptions de genres et d'especes nouvelles de Cryptophagidae. In: A. Grouvelle. Memoires Entomologiques. Etudes sur les Coleopteres. Fasc. 2. pp. 70-203. Groi Mill. A., AND A. RAFFRAY 1912. Supplement a la liste des Coleopteres de la Guadeloupe. Ann. Soc. 1 m. France, 81: 289-312. HlNTON, H. E. 1945. A Monograph of the Beetles Associated with Stored Products. I ondon, British Museum (Natural History), vii f 443 pp. Kraatz, ( i. 1899, Ueber die Languriiden Arten von Kamerun nebst einigen ver- wandten l ormea Deutsch. I at Zeit, 1899: 307-313. 1 ECONTE, J. 1 . 1861. Classification of the Coleoptera of North America. Part I. i 1st Part > Smithsonian Misc. ( oil., ix\ + 208 pp. 1863 New Bpecies of North American Coleoptera. Part I. Smith- sonian Misc. c oil.. No. 167. pp. 1-86. I i NDBI BO, S. 1966. Eicolyctus brunneua GylL (< oleoptera), nagot om hi. a. hiolo- gin. Int. ridskr., 87: 47-49. \1 \RIIM .'. V. \M> A. H\RKI R\ 1966. Hallazgo de ( ryptophagidae anoftalmos y aptcros. associados a mamiferos. ( iencia, 25(1): 11-16. Mora in i sky. V. IS' ai dun catalogue des insectes de l'lle Ccylan. Bull. Soc. Imp. Nat. Moscou, 36 (1): 421-532. Rl I 1 1 1 R. I IS77. Hapalips, neue Gattung der Rhizophagidae. Verhandl. Naturf. Ver. Brunn. IS: 122-128. R\ mi R Robi are, A. w. 1939. On the taxonomy of Erotylidae (Coleoptera). with special rcf- ence to the morphological characters of larvae. Trans. Roy. Ent Soc. London, 88: 89-118. Ri mi R ROBI Ris, \. W.. \nd F. Van Emden 1958. On the taxonomy of the Erotylidae (Coleoptera I. with special reference to the morphological characters of the larvae. Trans. Roy. Ent. Soc. London. 110: 245-285. SCHENtXINO, S. 1923 < ryptophagidae. In: W. Junk and S. Schenkling, eds.. Coleop- terorum Catalogus. Vol. XV. Pars 76, 92 pp. 1968 GENERA OF LOBERINI 21 1928. Languriidae. In: W. Junk and S. Schenkling, eds., Coleop- terorum Catalogus, Vol. XV, Pars 10, 40 pp. Sen Gupta, T. 1967. A new subfamily to Languriidae based on 4 genera, with a key to the species of Toramus. Proc. Roy. Ent. Soc. London, Ser. B, 36 (11-12): 167-176. Sen Gupta, T., and R. A. Crowson 1967. The systematic position of Eicolyctus Sahlberg (Coleoptera: Languriidae). Proc. Roy. Ent. Soc. London, Ser. B, 36 (5-6): 87-93. Sharp, D. 1900. Fam. Cryptophagidae. In: F. D. Godman and O. Salvin, eds., Biologia Centrali-Americana, Insecta, Coleoptera, Vol. II, Part 1: 579-626. VlLLIERS, A. 1942. Notes sur quelques Cladoxenitae Indo-Malais (Coleoptera: Erotylidae). Arb. Morph. Taxon. Ent. Berlin-Dahlem, 9: 89-92. 1943. Etude morphologique et biologique des Languriidae (Col. Erotylidae). Publ. Mus. Nat. Hist. Natur., No. 6, 98 pp. 1961. Revision des Coleopteres Languriides Africains. Ann. Mus. Roy. Afr. Cent., Ser. 8°, Sci. Zool., No. 98, 385 pp. (Received 19 November 1967.) EXPLANATION OF LETTERING ON FIGURES ac — anal cell mt — metasternal knob or as — apical spine projection at — anterior tendon my — pocket or mycangium ( ? ) cc — mesepisternal pocket p — paramere cp — corpotentorium pa — peglike setae ct — coxite Pp — paraproct es2 — mesepisternum pr — prostheca ga — galea s2 — mesosternum hr — hypopharyngeal bracon s3 — metasternum hu — hump si — styli la — lacinia si — stridulatory file lp — lateral plate su — supratentorium Is — longitudinal line on tc — trochanter metasternum tg — transverse groove on anterior It — laminatentorium part of gular region m — mola ur — urogomphi ml — median lobe vf — valvifer mp — mesosternal pocket vp — sensory appendage ms — median strut vt — ventral crushing tubercle 22 URLVIORA No. 303 1968 GENERA OF LOBERINI 23 \c -2 3 C 00 o — hi 2= O 3 ph — C/3 "q 03 u 3 to tH o E > ■*— » C o .Br '> O a. «0 to 3 (U ~3 x" u O t-5 O > oi a. 5 ex. c o X* C3 a: x; 5 l- Ih to <*- CL O 5; o o J2 £ CA — to 3 v. t-^ to 3 4) 3. ttf to" E 5 U T3 U 3 k) s -g to < a •« c cs C s c O R %, ^t to => S «J "-) S r- o -^ <. <*-> -~ j c a o '- ■s P s- a; « .s 0 in a oq Oh 3 ^> OX) to ■*, to OO to 5 a « 3 s ■* «c 1 3 1 3 :tr o c ^ S u -Q ^ to _^ -~- .5 «> 8.J-S &i a ' o o „ S- > 2 . -3 13 5 a; ^ u P a 3 24 BREVIORA No. 303 1968 GENERA OF LOBERINI 25 o o fe .-3 &*a .a -I -s a ■ - — i to ~ n S B -•So u c i u o a> o g a u '•3 "2 -s 5 S3 > .5- ^§ £ £- ^ ^ W .a — -2 c O 60 a 5 C a > 'c £ C H 5j :- 2 i O " J?-~ « p a Si ° . a •i « § ,_ -s- ;£" - « xi "2 Q O *> n £ a a as 6 ■S-.y o *s *2 -2 .9 00 60 . <*h •S -5 S ° *> ■ .2 -g if in oo 5 ' o <5 "3 'C 33 a • *•• C <-* S • £ fc > s 3 . S.S.'S s ss al a: «> 5" ^ ■» w -5 -C 26 BREVIORA No. 303 1968 GENERA OF LOBERINI 27 X) B 4> a s o3 w> £ CD to 4_l l_, 1) .g -1 ON CO 13 si co i- .23 O "> •o , «s 03 rt u n c ■-> C (U c < 6 3 od • *^ rj 03 &T J u . > 03 • *• CO £ o ~a > „ ^ *o 03 CO u X CO O t^ « s a. ^ • 03 •5" «*} "Sl, "3 . n CO !3 s "o3 a: b a 03 6 o <4-l o XI Oh 03 > O X) 03 03 I X 4— » — v-i Tt © ro ■ •* i ^ £ cs CD .23 "> '> co' o "o3 13 £ CO CO 1- o o T3 T3 BREVIORA Mm senium of Comparative Zoology Cambridge, Mass. 31 December, 1968 Number 304 REDESCRIPTIONS OF ANACHIS AVARA (SAY) AND ANACHIS TRANSURATA (RAVENED WITH NOTES ON SOME RELATED SPECIES (PROSOBRANCHIA, COLUMBELLIDAE)1 Amelie H. Scheltema- ABSTRACT Both Anachis avara (Say) and A. translirata (Ravenel) are found from Massachusetts to southern Florida. The shell of avara varies geographically from short and broad with many fine ribs in the north to tall and slender with a few heavy ribs in the south. A. translirata does not vary in shell morphology over its range. In New England the two species are very commonly found together at mean low water and have usually been confused with each other because of a superficial similarity in ribbing on the shell. Egg capsules and veliger larvae of northern and southern populations of avara are morphologically similar. Egg capsules, larvae, and radulae of translirata are different from those of avara. In southwestern Florida, avara is replaced by A. semiplicata Stearns, which appears to be a species endemic to that area. The relationships of an Anachis species in the western Gulf of Mexico are not clear. Egg capsules of A . floriclana Rehder show it to be a species distinct from avara. A. similis (Ravenel) is considered to be a nomen dubium. 1 Contribution No. 1888 from the Woods Hole Oceanographic Institution, Woods Hole, Massachusetts. - C/o Woods Hole Oceanographic Institution, Woods Hole. Massachu- setts 02543. BRIVIORA NO. 304 INTRODUCTION [wo very common gastropod species living along the shores oi southern New England arc the columbellids Ainu his avara (Say, 1S22) and A. translirata (Ravenel, ISM). The shells and living animals of both arc quite similar, and the two species have a long histor) of being confused with each other. South of New England misidentifications have been fewer tor two reasons: first, translirata is much less common near the tidchne from New Jerse) south ( see Table I ): second, avara is highly variable in shell form, and those from southern localities are less similar to translirata than those from New England. Gould ( 1841 ). in Ins earl) work on New England invertebrates, recognized onl) one species, avara. Verrill ( 1873) noted correctl) that another species oi Anachis occurs in New England, which he called A. similh (Ravenel. 1861). llowe\er. I regard similh a nomen dubium (sec p. 12) and translirata the correct designation. \ errill figured similh with a shell of avara reproduced from Gould I 1870). From VerrhTs work until the present time, "avara" has been the name more USUall) applied to both species; when trans- lirata has been recognized .is a distinct form, the names "similis" and "translirata" have been used indiscriminately, often as varieties or subspecies ol avara (e.g.. Dall, 1889; Johnson. 1934). Abbott (1954) gave correct names and descriptions of the two species. but unfortunate!) the plate figures arc reversed and the geographic ranges are not correct. Recently, Scheltema and Scheltema (1963) described the egg capsules and \ehger larvae o\' translirata as those of avara. be- cause the adults from which egg capsules were obtained were com- pared with misidentified museum specimens. The error became evident when new ke\s to invertebrates in the Woods Hole region were being compiled (Smith, ed., 1964). Subsequently, avara were collected alive in Beaufort, North Carolina, and held in aquaria; these snails deposited egg capsules that were distinctly different from those of translirata. This fortunate circumstance made pos- sible the identification o\ similar capsules often found attached to eel >:rass m the Woods Mole area, and consequently larvae have been reared from both Beaufort and Woods Hole populations of avara. I hese larvae were morphologically the same, and distinct!) different from those of translirata in their soft external anatomy (Scheltema. MS in preparation). I hereforc. on the basis of cap- sules and larvae, it has been \erilied that the two New Faigland forms oi Anachis are distinct species, and also that the southern and northern forms of avara constitute a single species. 1968 REDESCRIPTION OF ANACHIS SPECIES 3 The necessity for redescribing the two species and for determin- ing their geographic ranges became obvious if common errors in identification were to be corrected. ACKNOWLEDGMENTS Material has been made available to me through the generosity of many individuals and museums. I would like particularly to thank Emma B. Richardson of the Charleston Museum, Joseph Rosewater and George Radwin of the U.S. National Museum (USNM), R. Tucker Abbott and Robert Robertson of the Acad- emy of Natural Sciences of Philadelphia (ANSP), and William J. Clench of the Museum of Comparative Zoology (MCZ). Robert H. Parker, Jack B. Pearce, and Edmund H. Smith, formerly of the Systematics-Ecology Program, Marine Biological Laboratory, Woods Hole, kindly allowed me to make use of their Anachis col- lections. I am extremely grateful to Louise Ridge for her many efforts and hours of collecting to provide me with live specimens of Anachis floridana, which she mailed from Florida. John R. Hall and Robert Robertson made the photographs of the shells on Plate I, for which I am very appreciative. Thanks are due also to Ruth D. Turner and Kenneth J. Boss, who gave this paper a criti- cal reading and made several helpful comments. Finally, I take pleasure in acknowledging the financial support of The Radcliffe Institute, a grant from which has made this work possible. Anachis translirata (Ravenel) Plate 1, figs. 1, 2, 6; Text-figs, lb, 2b; Table 1 Columbella translirata Ravenel 1861, Proc. Acad. Nat. Sci. Philadelphia 1861: 42 (no fig.) ("off Charleston bar" [South Carolina]). [Type specimen probably destroyed.]3 Columbella avara Say. Gould 1841, Invertebrata of Massachusetts, pp. 313-314 (in part), fig. 197; Tryon 1883, Manual of Conchology, 5: 159 (in part). 3 Much of Dr. Ravenel's collection, which lay in the line of General Sherman's march through South Carolina in 1865, was destroyed; no labeled specimens of either A. translirata or A. similis remain in his collection at the Charleston Museum. Ravenel's description of translirata, although not figured, is sufficiently precise to enable one to ascertain the species to which he was referring. UR1 VIORA No. 304 Inachis avara (Say). Perkins 1869, Proc. Boston Soc. Nat. Hist. 13: 113 (in part); Sumner. Osburn. and Cole 1913, Bull. U.S. Bur. Fish. 31(2): 710-711. 712 (in part): M. Smith 1945, last (oast Marine Shells, p. [19, pi. 46. fig. 24: Abbott 1954, American Seashells, pi. 25, lii:. ee (fig. only); Scheltema and Scheltema ll>63. Hydrobiologia 22(1-2): 85-91, 13 text-figs, (egg capsules and larvae). Anachis similis (Ravenel). Verrill 1873, Kept. IS. Comm. 1 ish. and Fisher- ies 1871-1872: 644-645 (description only, not fig. I<>1>>. Anachis avara translirata (Ravenel). Dall 1889, Bull. U.S. Nat. Mus. .'{7: 116. Anachis avara similis (Ravenel). Dall 1889, Bull. U.S. Nat. Mus. :*7: 116; M. Smith 1945, last Coasl Marine Shells, p. ||>). p|. 4(>. fig. Il> (juve- nile). h-J t=3 rext-Fig. I. Radular teeth of (a) Anachis avara and (b) Amu his transli- rata. Scale line equals 0.2 mm. Description. Shell elevated, conical, whorls llattcned and regu- larl\ ribbed. Height to 17.5 mm. width to 6.5 mm. median range in height 12.5 to 14.0 mm. Average width to length ratio 0.40. Aperture a little less than one-half total height of shell. Larval whorls (protoconch) three, smooth, rounded, and translucent. 1968 REDESCRIPTION OF ANACHIS SPECIES Post-larval whorls seven or eight, nearly flat, with distinct, regu- larly spaced, straight axial ribs and less distinct spiral striae. Suture indented. Ribs usually parallel to shell axis, and most ele- vated at the suture. Those on the spire extend from suture to suture and are about the same width as the interspaces. Ribs on the body whorl average 15 (range 11 to 20), are not as closely spaced as those on upper whorls, and extend from the suture to just below the periphery, except that those near the aperture extend only to the periphery. Periphery somewhat angulated. Faint axial ridges sometimes apparent below periphery, extending between ends of ribs of body whorl. Outer lip often thickened so as to oblit- erate one or more ribs. Spiral striae usually distinct between ribs. Text-Fig. 2. Egg capsules of (a) Anachis avara, (b) Anachis translirata, and (c) Anachis floridana. Scale line equals 1.5 mm. 6 breviora No. 304 The first spiral stria following the suture is the widest, uually the most deeply impressed, and crosses the ribs, producing the subsut- ural nodules that are the distinctive shell character of the species. Other striae may or may not cross ribs, giving a more or less can- cellate appearance: the first two post-larval whorls in particular are often beaded. Spiral striae least impressed at periphery. Below periphery, spiral striae are deeply impressed and form the dominant shell sculpture; they produce weak beading as they cross the axial ribs and ridges. Aperture elliptical. Outer lip thin or thick, re- curved anteriorly to form a short siphonal canal. When the lip is thickened it is recurved posteriorly into an anal notch. Teeth may be present on outer lip when it is thickened, and range from three to nine (average 6) in number, often indistinct, the largest posterior. Parietal callus, when present, is narrow, sharp-edged, and bears a number of teeth, which are the ends of the spiral striae. Columella is smooth. Shell is dull, color varying from straw-yellow to chestnut-brown, often with spiral streaks o\' white, especially at subsutural knobs and angulation o\ periphery, emphasizing these characters. Juveniles are without thickened lip or callus, and with the periphery sharpl) angulated. Operculum elliptical, concentric, with eccentric nucleus. The radula is rachiglossan; the median tooth is a flat plate; the lateral tooth has three cusps, with the proximal one rounded (Text- fig, lb) " Pigmentation oi head and loot similar to that of A. avara ((/.v. ). except that the white posterodorsal tip o\' the foot is small and grades into the mottling of the rest oi the loot. Egg capsule volcano-shaped, ringed b\ sculpture of concentric ridizes, with one ridge more pronounced than the rest (Text- fig.^2b ) . Remarks. The shell of A. translirata is differentiated from that of A. avara by its usually more flattened whorls and greater height, and by the spiral subsutural stria that crosses the axial ribs and is the widest stria. (This stria ma\ not be evident on all whorls, especially on an eroded or fouled specimen.) In avara, the spiral striae are usually not as pronounced and do not cross the ribs. The ribs of the first two post-larval whorls of avara are therefore smooth, whereas those of translirata are beaded. Juveniles of the two species are easily distinguished by the difference in width- length ratio, which is less variable in juveniles and therefore more obvious, and by the shape of the body whorl, which is sharply angulated in translirata but rounded in avara (PL 1. figs. 5, 6). The posterodorsal tip of the foot and the egg case of each species 1968 REDESCRIPTION OF ANACHIS SPECIES 7 also serve to differentiate them. The lateral teeth of the radulae have one small consistent difference between the two species: in avara the proximal cusp bears a denticle; in translirata it is rounded (Text-figs, la, b). The Pliocene fossil figured as Anachis (Costoanachis) avara translirata (Ravenel) by Gardner (1948, pi. 30, figs. 36, 37) is probably not this species. Habitat and distribution. This species lives on shelly bottoms, rocks, pilings, or other firm substratum, from below mean low water to 48 fathoms, from Cape Cod Bay and Georges Bank south to Florida (see Table 1). It is very common subtidally in New England, where it can also be found in beds of eel grass with A. avara. Specimens examined. The entire collections of translirata of the MCZ and the USNM were examined. Shell measurements and descriptions are based on specimens from the following localities: Nobska Point, Woods Hole, Mass. (Scheltema collection) 19 (living) Quicks Hole, Mass. (Scheltema collection) 54 (29 living) Sullivans Island, S.C. (MCZ 260887) 34 Hadley Harbor, Mass. (R. Parker collection) 30 Wellfleet, Mass. (MCZ 19372) 32 Beaufort, N.C. (MCZ 256848) 1 (living) Radulae were examined from four of the specimens taken from Quicks Hole. Descriptions of the pigmentation of the head-foot are based on the 1 9 specimens from Nobska Point. Anachis avara (Say) Plate 1, figs. 3-5, 7-10; Text-figs, la, 2a, 3 and 4; Table 1 Colombella [sic] avara Say 1822, Jour. Acad. Nat. Sci. Philadelphia 2: 230 (no fig.) ("coast of the southern states"). [Lectotype, here selected, Academy of Natural Sciences of Philadelphia No. 16887, ex Mrs. Say's collection, "Florida."] Non C. avara Duclos 1840, Histoire Naturelle Coquilles Univalves Marines, pi. 1, figs. 1, 2. Columbella avara Say. Gould 1841, Invertebrata of Massachusetts, pp. 313-314 (in part, not fig. 197); DeKay 1843, Natural History of New York 5, Mollusca: 139-140, pi. 8, fig. 179. Amycla (Astyris) avara (Say). H. and A. Adams 1858, Genera of Recent Molluscs 1: 187. Anachis avara (Say). Perkins 1869, Proc. Boston Soc. Nat. Hist. 1869- 1871, 13: 113 (in part). 8 uriviora No. 304 Anachis avara "Perkins.' Verrill 1873. Rcpi. U.S. Comm. Fish and Fish- eries 1 8 7 1 - 1 S ~ 2 : 643-644 (no fig.). Anachis similis (Ravenel). Verrill 1873. Kept. U.S. Comm. Fish and Fisheries 1871-1872: pi. 21. fig. 109 (fig. only, not description pp. 644-645). 25 I II II I ! II II [I II I N = 50 WOODS HOLE, MASS- R.PARKER COLL. 25 -I — n — n — n — n — n — n N=62 BEACH HAVEN, N.J. ANSP 71291 N=27 BEAUFORT, N.C. MCZ 81921 ii ii — 1 1 1 1 i A/--/8 * BEAUFORT, N.C. MCZ 256849 i — n — n — n — n — n — n — i 25 .111.1 A/-"/ 7 CHARLSTON.S.C MCZ 256850 i — n — ii ii — i i i i — n — r-n — i — n — n — i — i 25 r N=103 LAKE WORTH, FLA. ANSP 221 188 h — n — r I 1 I — n — n — rn CO O CM « V I \ piuiiaiA spines taken from be- tween the ambitus and the apical system and broken in half about 2 nun from the milled ring. About 10 spines were taken from each specimen. I he number ol wedges varied from II to 15. with a mean o\ 12.34 wedges (SI) 0.7683, ( V. 6.21). Pedicellariae. Globiferous, tridentate, ophicephalous, and tri- phyllous pedicellariae are present but not diagnostic (Fig. }u-\). The pedicellariae are purple, and the color is retained after clean- ing with bleach. Spicules o\ the globiferous pedicellariae are dumbbell-shaped (Fig. 3g). Large, white stalk glands are present on main globiferous pedicellariae Coloration. I he spines are USUall) deep, rich green, although two of the specimens have white spines. I he pedicellariae are purple, and tlie test is beige with a purple-brown stripe covering the median longitudinal suture o\ the ambulacra ami intcrambu- lacra. After cleaning with bleach, the test is mottled green-grey. Habitat. I he specimens were found living in niches on the top of a living, shallow coral reel at Buena Ventura, Panama. Most oi the specimens were associated with the leaf coral Agaricia agari- < itei ( Fig. 4). I he gut of one specimen was filled with small grey- green spheres, most o\ which were o\ unrecognizable origin. These urchins probably graze on algae and encrusting organisms of the reef. There was no evidence that the) were feeding on coral. Un- like L. variegatus (and most other toxopneustid echinoids), /.. willianiM docs not cover itself with bottom debris. 1968 LYTECHINUS WILLIAMSI 5 The specimens collected in December began to spawn when placed in alcohol, which might indicate a winter spawning season. Moore et al. ( 1963), however, found that L. variegatus may spawn to some degree during the entire year. Relationships. L. williamsi lives in the same depth range as L. variegatus and in a portion of its geographic distribution. Speci- mens of L. variegatus can be found in the grass areas at Buena Ventura. Although occasionally found on rocky areas or on soft sediments, L. variegatus normally occurs in grass areas (Moore et al. 1963; personal observations), while L. williamsi appears to be distributed only on living reef areas. The new species differs from L. variegatus in several features, the most obvious of which is the number of wedges per primary interambulacral spine, which averages 12.34 in the former and 24.14 in the latter (Fig. 3b, Tables 1 and 2). The relative size of the apical system and peristome is greater in the new species than in L. variegatus. The number of plates per ambulacral or IA series is much greater in L. variegatus than in L. williamsi (Fig. 5). Jackson (1912, 1914), after examining 1,443 specimens of L. variegatus, found none with all oculars exsert, whereas all of the specimens of the new species have every ocular exsert. L. williamsi can be separated from the two remaining, relatively deep-water, Atlantic species on the basis of the number of plates per series, the tuberculation, the coloring, and the spicules of the globiferous pedicellariae. L. euerces and L. callipeplus do not have naked areas in the median ambulacral and interambulacral area; the secondary tubercles are irregularly scattered on the test and are almost the same size as the poorly-developed primary tubercles. Whereas the spicules of the globiferous pedicellariae are dumbbell-shaped in L. williamsi and L. callipeplus, they are C- shaped in L. euerces. The naked test of L. callipeplus has red blotches on the dorsal side, which the new species lacks. L. williamsi is most closely allied to L. panamensis of the Bay of Panama. It differs from that species, however, in having fewer IA and ambulacral plates (Fig. 5). In addition, the pedicellariae of L. panamensis are brown, whereas they are purple in L. williamsi. Ocular I is normally insert in L. panamensis and exsert in the new species, and the milled ring of the primary spines is well developed in L. panamensis and almost absent in the new species. The other Pacific species are separated from the new species in the following key. 6 BREVIORA No. 305 KEY TO THE SPECIES OF THE GENUS LYTIC "7/ IN US 1. Secondarj tuberculation poorl) developed, leaving a naked median area in the aboral portion of the ambulacra and interambulacra; primarj tubercles large 3 Secondar\ tuberculation scattered irregularl) over the test, no naked areas, primarj tubercles small 2 2. 1 arge, red blotches dorsall) on the cleaned test, spicules of globiferous pedicellariae dumbbell shaped (Caribbean, Gulf of Guinea, deep water) /.. callipeplus Test and spines white, spicules (shaped <( aiibbean. deep Water) .... / . Cllt I i ( \ 3. Spicules of globiferous pedicellariae ( -shaped (< alifornia) .../.. pictus Spicules dumbbell shaped 4 4. Primar) I \ spines from above ambitus with less than 16 wedges .5 Spines with more than 16 wedges 6 5. Adults with less than 14 |\ plates pei senes. pedicellai iae purple (At- lantic coast of I'. m. una I L. williamsi VdultS with more than 14 I A plates pel series, pedicellariae brown i Ba) of Panama » / . panamensis 6. Ambital spines much longer than other primaries, producing a fringe: test low. mottled with purple (California) /.. anamensus Spines more or less uniform in size 7 7. Naked I A area granular, with a iow of small tubercles along the hori- zontal sutures, pedicellai iae purple, covering naked area (Galapagos, N.W. coast of South America ) /.. semituberculatus Naked I A area not pronounced, smooth, pedicellariae white, confined to ruberculated areas 8 S. Test green, mottled with white, spines green or tinged with purple i Brazil, West Indies i /.. variegatus typicus I est not greenish or whitish 9 l». Buccal membrane not densel) plated outside buccal plates K ape Verde Islands. Africa ) /-. variegatus pallidas Membrane heavily plated 10 10. Spines slender, dark purple (Bermuda) /.. variegatus atlanticus Spines reddish, stout ( U.S. east coast ) L. variegatus carolinus 1968 LYTECHINUS WILLIAMSI 7 TABLE 1 L. williamsi: equations for regression lines from plots of taxonomic features of the test. Apical system ( V-2 ax's, in mm ) : LnY = 0.8162LnX - 0.7688 Sv.x = 0.056 r = 0.89243 Peristome (V-2 axis, in mm): LnY = 0.7306LnX + 0.0646 Sv.x = 0.025 r = 0.97049 Interambulacral plates per series: LnY = 0.3832LnX+ 1.3117 Sv x = 0.039 r = 0.80026 Ambulacral plates per series: LnY = 0.7321LnX + 0.5368 Sv x = 0.038 r = 0.93362 Number of wedges per IA spine from above the ambitus (N = 250) : Mean = 12.34. S.D. = 0.7683, C.V. = 6.21 TABLE 2 L. variegatus: equations for regression lines from plots of taxo- nomic features of the test. Apical system ( V-2 axis, in mm ) : LnY = 0.7245LnX - 0.7219 Peristome (V-2 axis, in mm) : I nY = 0.824;'Lnx - 0.3291 Interambulacral plates per series: LnY - 0.3916LnX + 1.6275 Ambulacral plates per series: LnY = 0.5397LnX + 1.4525 Svx = 0.081 Sv x = 0.043 Svx = 0.042 Svx = 0.037 r = 0.97798 r = 0.99107 r = 0.97057 0.99248 Number of wedges per IA spine from above the ambitus (N = 500) Mean = 24". 14, S.D. = 1.7924, C.V. = 7.4 8 BREVIORA No. 305 TABLE 3 Material of /-. variegatus examined. \4.CZ. No. I o< ality 8179 Near Canal, Panama Buena \ entura, Panama 1252 Bahia, Brazil 3346 Bahia. Brazil 7412 Pigeon Pi., [bbago 8042 Bancs Bay, ( uba 5070 lamaica Santa Marta, c olumbia 7939 ( edai Key, N.W. I lorida 4444 I oi tugas, I loi ida I 28 1 Beaufoi i. Not ih ( arolina 3340 Bermuda Size i///> in mm) N, umber 21 to 39.6 3 :: to 27 2 10 to 41.5 5 41.5 1 30 1 15 to ;; 2 26.5 1 34.3 1 16 to 30.2 5 5 5 to 63 5 21.5 1 26 1 LITERATURE CITED C III Mil K. R II 1966. Report on the I chinoidea collected b) k \ I'll I SBURY in the Gulf of Guinea. Stud ["rop Oceanogi Miami I (1):209- 223. ( K(.\ k.n I I 1959. I lementan Statistics with Applications in Medicine and the Bio logical Sciences. Ne\» York, Dover, 375 pp. J \( kvis. K.I. 1^12. Phylogen) of the Echini with a revision ol Palaeozoic species. Mem. Boston Soc. Nat. Hist 7: 1-491. 1914. Studies of Jamaica Echini. ( arnegie Inst. Wash. Pub. 182: 139-162. \I\YR. E. 1954. Geographic speciation in tropical echinoids. Evolution 8(1): 1-18. 1968 LYTECHINUS WILLIAMSI Moore, H. B., T. Jutare, J. C. Bauer, J. A. Jones 1963. The biology of Lytechinus variegatus. Bull. Mar. Sci. Gulf and Caribb. 13 (1 ):21-53. MORTENSEN, T. 1943. A Monograph of the Echinoidea. 3(2). Copenhagen, C. A. Reitzel, 553 pp. Swan, E. 1952. Regeneration of spines by sea urchins of the genus Strongylo- centrotus. Growth 16: 27-35. 1958. Growth and variation in sea urchins of York, Maine. Jour. Mar. Res. 17: 505-522. (Received 21 March 1968.) y-.i f'\ ' \>|ff*^ s vim V M^VJjTi It 9 / ■*"• r •5T - Figure l. Dorsal view of L. williamsi (Paratype, 23 mm HD). 10 BR] V IORA No. 305 .V'y 'X • •. V' * •«,* -.« sf**tf. »• T ' -St. ■ ' Figure2. Lateral ;irul ventral view of /. wiMiamsi (Paratype, 23 mm HDl. 1968 LYTECHINUS WILLIAMSI 11 Figure 3. Spines, spicules, and pedicellariae of L. williamsi'. A, cross-section of a primary interambulacral spine 0.7 mm in diameter; B, the same, from L. variegatus 0.65 mm in diameter; C, valve of a triden- tate pedicellaria 1.09 mm long; D, valve of globiferous pedicellaria 0.8 mm long; E, valve of triphyl ous pedicellaria 0.155 mm long; F, valve of ophice- phalous pedicellaria 0.55 mm long; G, spicules of globiferous pedicellaria 0.04 mm long; H, spicules of the tube feet 0.04 to 0.047 mm long: I. base of a primary spine, 0.8 mm in diameter. 12 imi \ iora No. 305 Figure 4. Underwater photograph of /.. wiiiwmsi at the base of some le .t coral [Agarh ia agarit ih 1968 LYTECHINUS WILLIAMSI 13 30- 20- 10- 20- cv ■10- % t •I o o oo o o ooo • = L. variegatus §=L. panamensis ° L. will iamsi V • -9. § . i * »° O 10 20 30 Horizontal Test Diameter -i r 40 mm Figure 5. The number of plates in an ambulacral plate-series and an interambulacral plate-series for L. williamsi, L. variegatus, and L. pana- mensis. BREVIOIA Museum of Comnparatlve Zoology Cambridge, Mass. 31 December, 1968 Number 306 A NEW SPECIES OF ELEUTHERODACTYLUS (AMPHIBIA, SALIENTIA) FROM THE GUAYANA REGION, EDO. BOLIVAR, VENEZUELA Juan A. Rivero Abstract. A new species of Eleutherodactylus, E. pulvinatus, is de- scribed from the wet region of La Escalera, Estado Bolivar, Venezuela. The species is characterized by its small but distinct tympanum, poorly defined or absent vomerine teeth, large, transversely oval disks which are larger than the tympanum, a W-shaped marking on the occiput and non- spiny nuptial pads. The possible relationship between this frog and the genera Eupsophus, Syrrhophus, Niceforonia, Trachyphrinus and Phryno- pus is discussed and evaluated. Among the specimens recently collected by the author in the region of La Escalera, Estado Bolivar, Venezuela, there are seven specimens of a small and peculiar leptodactylid whose generic allo- cation has been somewhat difficult and problematical. In this species, the vomerine odontoids are very indistinct; in some indi- viduals they appear to be absent altogether. As in Eleutherodac- tylus, the disks are well developed and the terminal phalanges are T-shaped, but there are two well-developed glandular pads on the inner side of the first digit, a character which does not seem to have been described in that genus. It has been customary to allocate all Eleutherodactylus-like. forms without vomerine teeth to the genus Syrrhophus (type local- ity, San Antonio, Texas) and all those with vomerine teeth but without disks to Eupsophus (type locality, Chile). In 1932, Parker violated convention by describing Eupsophus wettsteini (type local- ity, Pongo, Peru, 2000 m) as "the first species that has been de- scribed as belonging to the genus Eupsophus in which vomerine teeth are lacking,, (p. 344). Even a superficial examination reveals that Andean species of Syrrhophus are not closely related to the animals that are referred 2 BREVIORA No. 306 to that genus in Central America and the United States. Whether or not they have vomerine teeth, most of the Andean animals seem to be closely related and should perhaps be called Eupsophus until a more thorough study can be undertaken and their relationships established. It is evident that the three groups of Eupsophus (Thoropa, wettsteini-peruanus, and the Chilean forms) mentioned by Cei ( I960. 1962) constitute different evolutionary lines, one of which {wettsteini-peruanus) appears to continue along the Andean Cordillera to Venezuela, where Eupsophus ginesi is known from glacial streams in the paramos oi the Merida Andes. It is now USUall) recognized that Thoropa is a separate genus, and perhaps the wettsteini-peruanus-ginesi group should be recognized as an- other, in which case. Oreobates Jimenez de la Espada, 1872. might appear to be the appropriate name. Yet. although Oreobates was described from Quijos (whence the name quixensis), and this name has been interpreted to mean Quito (Peters. 1955: 347; Gorham 1966: 117). Espada himself states: "It occurs in the Oriental province, formerly Quijos. in Ecuador, at the foot of the Cordillera" ( \ crtcbrados del \ iajc del Pacifico, 1875: 104). Specimens from the Napo-Pastaza region in the M.C.Z. correspond to the description o\' quixensis, which looks more like a Thoropa than a Eupsophus. Two new genera, each based on a single individual from the high elevations of the Colombian Andes, were described by Coin and Cochran in 1963. The first. Nicejoronia, has distinct vomerine teeth, no disks and no external tympanic disk; the other. Trachy- phrinus, has no vomerine teeth but has small disks (and T- shaped phalanges) and a well-defined tympanum. In general physiognomy, both of these genera resemble the group of frogs here considered, and Trachyphrinus does not look at all like Syr- rhophus, the genus in which it would be included if its classifica- tion were to be based on conventional morphological characters. But all kinds of combinations of characters have been described in the high elevation species from Peru to Venezuela. Thus, in 1873. Peters described Phrynopus peruanus (now called Eupso- phus, fide Gorham. 1966) with vomerine teeth and tympanum, but apparently without disks. The same combination is found in Borborocoetes columbianus Werner (now Eupsophus, fide Gor- ham. 1966). Paludicola festae Peracca (now Syrrhophus, fide Gorham. 1966). and Eupsophus ginesi Rivero. However, in Pal- udicola simonsii Blgr. (now Syrrhophus, fide Gorham, 1966). Syr- rhopus montium Shreve and Syrrhopus juninensis Shreve (both now Syrrhophus, fide Gorham. 1966) there are no vomerine teeth. 1968 NEW SPECIES OF ELEUTHERODACTYLUS 3 external tympanic disk, or disks. In Eleutherodactylus whym- peri (Blgr.) and E. surdus (Blgr.), on the other hand, vomerine teeth and disks are present but the tympanum is not apparent externally, while in Eupsophus wettsteini there are no vomerine teeth or disks but there is a small tympanic disk. Tra- chyphrinus myersi and Syrrhophus areolatus Blgr. are alike in having disks and external tympana (small and ill defined in S. areolatus) but no vomerine teeth. It is not the purpose of this paper to enter into the many prob- lems pertaining to this interesting group (or groups?) of frogs. But these digressions have been necessary because of the diffi- culty of allocating the new species described in this paper. If this group of Andean frogs is found to constitute a natural assemblage, perhaps the name Phrynopus can be used for it. It is also possible that it may constitute two or more evolutionary lines, in which case the genera suggested by Goin and Cochran may be found useful. It appears to me, however, that this group, or most of it, constitutes a latitudinal extension of Chilean and Argen- tinian Eupsophus along belts of similar climatic conditions in the Andes. If, as has been reported, some of the species are limited to paramos in the Andean Cordillera, their present diversification is to be expected. In a group where the tympanum-vomerine odon- toids-disks can occur in so many combinations, separation into various genera is hardly convenient and may actually be very confusing. A number of "Syrrhophus" have been described from the low- lands or at most, from elevations below 800 m in Ecuador. Such is the case, for example, with S. chalceus (Peters), S. calcaratus Andersson, and S. coeruleus Andersson. S. chalceus has also been reported by Lutz and Kloss, 1952, from Iuarte, along the Rio Uaupes. As the largest of the species described was 30 mm (5. coeruleus), all the others ranging from 18 to 29 mm, it is not im- probable that the lack of vomerine teeth can be attributed to immaturity in some of them. It is possible, however, that they represent another line of Eleutherodactylus without vomerine teeth (since in this genus disappearance of the vomerine teeth does not seem to be a difficult process) or that they are in some way con- nected with the Eupsophus group from Chile and/or the more northern Andes. Eleutherodactylus pulvinatus, the species described here, does not have the rotund physiognomy of the Eupsophus group, and its distribution does not suggest relationship. Nuptial pads have been described for some members of the Andean group (Parker, 1932: 4 BR! MORA No. 306 363), hut most of the species do not seem to have them. In the Chilean species (Cei, ll>o2). on the other hand, and in Thoropa (Cochran. 1955: 91), thumb and sometimes breast and axillar spines have been described for those species in which males arc known. Similar modifications have been described for Crossodac- tylodes Cochran (type localit) Macae. Rio de Janeiro), a inono- typic genus with hidden tympanum and the vomerine teeth represented onh b\ a roughened ridge. In view o\' the foregoing considerations, it has been found preferable to describe the new leptodactylid in the genus Eleutherodactylus, pending a more complete studv o\ its anatoim and relationships. I I I l llll RODA< H I. US Pill VINATUS sp. n. Type. M.C.Z. 64741, ■' , Paso del Danto, Region de la Esca- lera around 1400 m above San [sidro, road from II Dorado to Sta. Elena de Uairen. Estado Bolivar, Venezuela. Coll. J. A. Riveroand J. Pulido, :<> March 1968. Diagnosis. A small Eleutherodactylus with small but distinct tympanum, vomerine teeth poorl) defined or absent; two non-spin) nuptial pads on the inner side o\ the first digit o\ males; large trans- versel) oval disks, larger than the tympanum; first linger shorter than second; adpressed heel extending to between eye and nostril: and a W -shaped marking behind the occiput. Description. Mead as long as broad; snout short, subtriangular; tongue free and slightl) nicked behind: vomerine odontoids faintl) indicated as two irregular elevations well behind ami between the small choanae; eye diameter slightk greater than distance between eye and nostril but shorter than snout: interorbital space as broad as an upper eyelid; canthus rostralis rather indistinct. curved; loreal region onl) slightK sloping, concave; tympanum dis- tinct. ' .* the eve diameter; a Hat. indistinct supratympanic fold; a line of small, whitish tubercles from elbow to wrist; two distinct. oval, metacarpal tubercles; a glandular pad at the base o\' the inner side of the first finger and another one distal and slightK dorsad to this; first finger shorter than second, its disk much broader than the phalanx; all fingers tree: larger disks larger than the tympanum; a small. Hat. whitish tubercle at the distal portion o\ the tibial segment: a distinct and prominent inner and an indis- tinct outer metatarsal tubercle: toes free, with prominent subarticu- lar tubercles: heel of the adpressed hind limb extending to between eye and nostril. Skin above, shagreened. more coarsely tubercular in the tympanic area and anterior flanks. Below, finely granular on 1968 NEW SPECIES OF ELEUTHERODACTYLUS 5 throat and anterior part of the belly; more distinctly granular on posterior part of the belly and thighs. A very large subgular vocal sac with a triangular anterior end; a space (about 2 mm) between tip of the jaw and anterior edge of sac. A "frenulum" in the an- terior tip of the sac continues anteriorly almost to the tip of the jaw. Color. Above, brownish gray, with a dark brown interorbital bar, a black supratympanic streak, a W-shaped marking on the anterior part of the back, and some large blotches and small spots on the posterior two-thirds of the back; on the flanks the blotches continue ventrally, forming oblique, wavy bars that are separated by light gray areas; anterior and posterior part of the thighs uniform brown, this color extending as bars across the gray color of the upper part of the thighs; limbs and fingers with cross-bars and blotches; upper lip with triangular blotches on a gray background; no canthal streak. Ventral surfaces dirty white, darker on the throat and distal portion of the limbs. Measurements (in mm). Snout-vent 26.1; head breadth 9.8 (between tympana); head length 10; femur 13; tibia 14.1; foot 1 1. Paratypes. There are six paratypes (U.P.R.-M.1 02214-02219), all coming from the type locality. All are males, and the size varies from 26.1 mm to 23 mm. In some of the smaller specimens the canthus seems to be less defined than in larger specimens, and a cross-section of the snout would be more or less rounded if it were not for the slight concavities of the loreal areas. The heel of the adpressed hind limb extends to the nostril in only one speci- men, but the skin of the dorsal surfaces is more granular than in the type, at least in two examples. The belly and thighs are uni- formly granular, the throat more finely so in most specimens. The vocal sac does not form the anteriorly triangular pouch that it forms in the type, but it is possible that it has been dis- placed, as a similar condition can be noticed in several specimens. The W-shaped marking is present in all specimens, but the oblique bands of the flanks are less distinct in some, although a trace of them can be noticed in all. Vomerine odontoids cannot always be seen, but they can be felt with a needle. They seem to be irregu- lar in both size and position, and in some specimens seem to be almost totally absent. Habits and relationships. All the specimens were collected at night, calling from low bushes ( 1 to 3 ft. from the ground) along the rocky bank of the road at an elevation of approximately 1400 m. At dusk, their calls seemed to come from under the rocks 1 University of Puerto Rico — Mayagiiez. 6 BREVIORA No. 306 lining the almost vertical road bank, but none could be collected at the tune. Later at night, their calls were traced and recorded and the various specimens collected. In the dark the Erogs are yellow or cream, but they turn dark gray in the daytime. Their voices were not heard much below the elevation where all the specimens were obtained. No other species of Eleutherodactylus was col- lected or heard in this area. Only scattered low bushes, some soil bromeliads and. occasion- ally, thickets of lilaceous plants occur in the rather open areas where E. pulvinatm was collected. There were permanent and temporary streams everywhere, as the region is probablv one ol the rainiest in Venezuela.1 The forest. 50 to 60 meters from the road, is a rather open montane forest, usuallv with man) epiphytic bromeliads. orchids, and araeeans. Most of the soil is a white quartziferous sand, which is covered in the forest by a layer of dead leaves and other organic matter. Although mam Eleutherodactylus have W-shaped markings on the back, the distinct markings ol E. pulvinatUS remind one oi the condition o{ E. W-nigrum (Boettger), which, however, is a species from more than 3000 m in the Amies o\ Ecuador. Other species. from relativel) adjacent regions, are: /•.'. inguinale Parker (New River, Guyana), E. beebei Cochran (Kartabo), E. pliciferus (Blgr.) and E. ramagii ( Blgr. ) (Iguarasu, Pernambuco, Brasil), / crepitans Bockermann (San Vicente, Matto Grosso), E, car- valhoi B. Lutz (Itacoai R.. NW Brasil). E. peruvianus (Melin) (Roque, Peru). E. vilarsi (Melin) (Taracua, Rio Uaupes), / rosmelinus Gorham (Rio Uaupes). /.. melini Bokermann (Roque, Peru). /:". altamazonicus Barbour and Dunn (upper Amazon) and /■.. marmoratus (Blgr.) (Mt. Roraima). In all these species there are well-defined vomerine teeth, and nuptial pads have not been described for am. E. inguinale has on obtusely angular canthus, a broad interorbital space, much broader than the upper eyelid, and a black bordered, inguinal ocellus; E. beebei Cochran is a large species 85.5 mm in length; in E. pliciferus and E. ramagii the first linger is longer than the second; E. crepitans, from drv forest, is similar to /.. conspicillatus, and its tympanum is % the eve diameter; E. carvalhoi does not have a tympanum; E. peru- vianus and E. vilarsi are 50 and 39 to 41 mm. respectively, and may be synonyms of E. conspicillatus; E. rosmelinus has a depressed head, disks VS the size of the tympanum and first finger 1 According to a report in a local newspaper, engineers working on the road from El Dorado to Sta. Elena could only work 60 to 80 days of the year, and even those were not always sunny days uithout any rainfall. 1968 NEW SPECIES OF ELEUTHERODACTYLUS 7 longer than second; E. melini has an indistinct tympanum and small disks; E. altamazonicus has a depressed and oval snout and a barely distinguishable tympanum; and E. marmoratus has a rounded snout as long as the orbital diameter, dark-edged marb- lings on the head, and a small size (only topotypical specimen: 19 mm).1 From the lowland "Syrrhophus" mentioned before, E. pulvinatus can apparently be distinguished by the nuptial pads it has in the inner side of the first finger. Other differences are as follows: In S. chalceus (Peters), from Pastaza River, the first finger is longer than the second; in S. coeruleus Andersson, from eastern Ecuador, the canthus is distinct and there are two pairs of dorsolateral folds; and in S. calcaratus Andersson,- from near Archidona, 800 m, there is a canthal streak, and the disk of the first finger is only slightly broader than the phalanx behind it. Acknowledgments. The author wishes to acknowledge all the courtesies and attentions extended to him by the Instituto Vene- zolano de Investigaciones Cientificas of Caracas, and most espe- cially by Dr. Gilberto Rodriguez, Head of the Hydrobiology Department, who went out of his way to foster the investigations resulting in this and other papers. During the course of this study the author was employed by and received all kinds of assistance from the I VIC. For the trip to La Escalera, the author relied on an invaluable assistant, Mr. Juan Pulido, always faithful and always willing to help during the difficult situations that often arise during such trips. Major Luis Alfonso Godoy, Commander, Engineering Corps at sector A of the road from El Dorado to Sta. Elena, was extremely helpful in providing facilities and assistance. As always, the author is greatly indebted to Dr. Ernest Williams, not only for permitting the use of the facilities of the Museum of Comparative Zoology at Harvard, but also for reading the manu- script critically and for discussing various taxonomic and evolu- tionary problems. Mr. Benjamin Shreve has also read the manu- script and suggested changes. The author, however, assumes full responsibility for the shortcomings of this paper. 1 Frogs collected in Mt. Marahuaca by the author (Rivero, 1961: 80) and pronounced conspecific with E. marmoratus by Parker, do not repre- sent E. pulvinatus. However, further comparisons should be made when more specimens of E. marmoratus become available. - Among the group of frogs here discussed, the species name calcaratus was first used by Giinther, 1881, for Cacotus calcaratus, now considered a synonym of Eupsophus roseus. 8 bki vior \ No. 306 REFERENCES VNDl RSSON, 1 \KS (i \HKII 1 1945. Batrachians from Easl Ecuador collected 1937. 1938, b) Wm. Clarke-Macintyre and Kolt" Blomberg. Arkiv Zool. 37<2): 1-88. B.XRUOl R. I IIOM \s. \ND I R. l)l NN 1921. Herpetological novelties Proc Biol Soc. Wash. 34: 157-162. linkl KM \NN. \\ I RM R 1958. A preoccupied name of a neotropical frog, genus Eleuthero- dactylus. Herpetologica I 1. pt. 2: 95. 1965. Tres novos batraquios da regiao central de Matto Grosso, Brasil (Amphibia. Salientia). Rev Brasil. Biol. 25(3): 257-264 BOI l I NG1 R, Cil I iRGl \l HI Rl 1882. Catalogue ol the Batrachia Salientia in the Collection of the British Museum, ed. 2. I ondon, Kvi -♦ 503 pp. 1888. On some reptiles and batrachians from Iguarasse, Pernambuco. \.m. Mag. Nat. Hist. 6(2): 40-43. 1898 \n account of the reptiles and batrachians collected b) Mi W. I ll Rosenberg in western Ecuador. Proc. Zool. Soc l ondon 1898: 107-126. 1900. Report on a collection made h\ Messrs. I V. Mc< onuel and J. J. Quelch at \lt Roraima in British Guiana. lYans. Linn. Soc l ondon (2) 8: 51-76. I I i, losi M I960 \ surve) of the leptodactylid frogs, genus Eupsophus, in ( hile. Breviora, Mus. < omp. Zool., No. MX: 1-13. 1962. Batracios de ( hile. Santiago de Chile, 128 f cviii pp. ( I I. losi \| . \S|) 1 I Is ( MM KRI) I s 5 s . Biologia \ desarrolo de Eupsophus taeniatus Girard. Inv. Zool. ( hUenas t 159-18 DUMERIL, \SDRl' M. C, AND GaBRIEI BlBRON 1841. Erpetologie Generate, Vol. 8, Paris, ii - 792 pp. ( OCHRAN, Doris 1955. I rogs of southeastern Brasil. Bull. U. S. Nat. Mus. 206: nvi + 423 pp. 1956. A new species of frog from KartabO, British Guiana Zoologica 11. pt. I: I 1-12. Coi'i . 1 DVk w D. IX7K. A new genus of < ystignathidae from Texas. Amer. Natur. 12: 252-253. ElUmmrsm (Newspaper) 1968. I April 6) En junio sc enla/aran los ingenieros militares de la carretera El Dorado-Sta. Elena. Caracas. Venezuela. 1968 NEW SPECIES OF ELEUTHERODACTYLUS 9 Gallardo, Jose 1965. A proposito de los Leptodactylidae (Amphibia Anura). Papeis Avulsos, Sec. Agric, Sao Paulo, 17, art. 8: 77-87. Goin, Coleman, and Doris Cochran 1963. Two new genera of leptodactylid frogs from Colombia. Proc. Calif. Acad. Sci. 31(17): 499-505. Gorham, Stanley W. 1966. Das Tierreich. Liste der rezenten Amphibien und Reptilien. Lief. 85, xvi + 222 pp. Grandison, Alice 1961. Chilean species of the genus Eupsophus (Anura: Leptodacty- lidae). Bull. Brit. Mus. Nat. Hist. 8(3): 111-148. Gunther, Albert 1881. Account of the reptiles, batrachians and fishes collected during the survey of H.M.S. "Albert" in the Straits of Magellan and on the coast of Patagonia. Proc. Zool. Soc. London 1881: 18-22. Jimenez de la Espada, Marcos 1872. Nuevos batracios americanos. An. Soc. Espanola Hist. Natur. 1: 85-88. 1875. Vertebrados del Viaje al Pacifico, Batracios. Madrid, 208 pp. Lutz, Bertha, and Gertrude R. Kloss 1952. Anfibios anuros do Alto Solimoes e Rio Negro. Apontamentos sobre algunas formas e suas vicariantes. Mem. Inst. Osw. Cruz 50: 624-678. Melin, Douglas 1941. Contribution to the knowledge of Amphibia of South America. Goteborgs Kungl. Vetensk. Vitterh.-Sam. Handl., Ser. B.. 1(4): 1-71. Miranda Ribeiro, Alipio 1926. Notas para servirem ao estudo dos gymnobatrachios (Anura) brasileiros. Archiv. Mus. Nac. Rio de Janeiro 27: 1-227. Parker, H. VV. 1932. The systematic status of some frogs in the Vienna Museum. Ann. Mag. Nat. Hist. (10) 10: 341-343. Peracca, Mario Giacinto 1914. Reptiles et batraciens de Colombie. Mem. Soc. Sci. Neuchatel 5: 96-111. Peters, Wilhelm 1873a. Hr. W. Peters las fiber zwei Giftschlangen sus Afrika und fiber oder weniger bekannte Gattungen und Arten von Batrachiern. Monatsb. Akad. Wissensch. Berlin, Mai: 471-478. 1873b. Hr. W. Peters las fiber eine newe Schildkrotenart Cinosternon Effeldtii und einige andere neue oder weniger bekannte Amphi- bien. Monatsb. Akad. Wissensch. Berlin, Sept. Oct.: 603-618. 10 BREVIORA No. 306 Pi I I Rs. I Wll S 1955. Herpetologica] t\pe localities m Ecuador. Rev. Ecuador Ent. Par. 2(3-4): 335-352. Kl\l Rd, J LAN A. I9M Salientia of Venezuela. Bull. Mus. Comp. Zool. I2i> new Syrrhopus from Peru. J. Wash- ington Vcad. Sci 28 u d c 03 c ^_ x: N <4-l o U £ £ ID '> ■, ^^ u 03 a. u >~, 1) -*— • cfl r. -J =0 ct d 2 _■*. £ > a. "5 ^ C/5 ^ 1_ ^^ o Q S3 ■a < o 5 03 -c; 53 -*■. 3 >3 N a> c u > QQ uT cS < > o '—< CD eb o T3 E ca w BREVIORA Mm senium of Comparative Zoology Cambridge, Mass. 31 December, 1968 Number 307 A NEW SPECIES OF HYLA (AMPHIBIA, SALIENTIA) FROM THE VENEZUELAN GUAYANA Juan A. Rivero Abstract. A small species of Hyla, H. rodriguezi, is described from the wet region of La Escalera, Estado Bolivar, Venezuela. The new form is characterized by having a large, well defined tympanum, about % the size of the eye; nearly webless fingers; 3A webbed toes; speckled throat and orange-red areas in the thighs. It may be related to the Hyla leprieurii group but it is also similar to Hyla grandisonae Goin, from which it differs in coloration and in other details. Among the new frogs collected by the author in the region of La Escalera, in southeastern Venezuela, there is a small species of a bromelicolous form that appears to be new to science. The new species is named Hyla rodriguezi in honor of Gilberto Rodriguez, of the Instituto Venezolano de Investigaciones Cientificas, who made possible the expedition during which this and many other interesting species were collected. Dr. Rodriguez spared no effort in facilitating the author's investigations of the frogs of the Vene- zuelan Republic. During this and other investigations pertaining to the Vene- zuelan frog fauna, the author has utilized the facilities and collec- tions of the Museum of Comparative Zoology at Harvard. He is most thankful to the Curator of Herpetology, Dr. Ernest Williams, for his cooperation and unfailing willingness to be interrupted and consulted at any time. Hyla rodriguezi sp. n. Type. M.C.Z. 64740, 6 , Paso del Danto, Region de La Escalera, ca. 1400 m above San Tsidro, road from El Dorado to Sta. Elena de Uairen, Estado Bolivar, Venezuela. Coll. Juan A. Rivero and Juan Pulido, 26 March 1968. 2 BREVIORA No. 307 Diagnosis. A small Hyla with bony head; strong, straight can- thus rostralis; almost vertical, concave lorcal region: anteriorly converging vomerine teeth; large tympanum, about 7« the size of the eve; tubercular dorsal surfaces (males?); red (alive) or white (preserved) areas on the thighs and speckled throat. Description. Head slightly broader than long; snout short, rounded, concave above; nostrils forming a truncate tip; canthus rostralis strong and well defined; loreal region vertical, concave; interorbital space as broad as an upper eyelid; vomerine odontoids anteriorly converging, extending from level of palatines to about middle of inner margin o\' choanae; choanae moderate, oblique; tympanum large, distinct, about 7* the size of the eye; larger disks much smaller than tympanum; eye diameter as long as distance between eye and nostril; a line o{ whitish tubercles from elbow to wrist; metacarpal tubercles not well differentiated; subarticular tubercles well defined; fingers almost free, the first swollen at the base and with an elongated brown pad which covers :i of its inner side; first linger not extending much beyond first subarticular tubercle oi second; second finder shorter than last; a small tubercle at the heel; no tarsal fold; an elongated inner and a flattened, rounded, and less distinct outer metatarsal tubercle; subarticular tubercles o\ toes prominent; toes about % webbed (taken in order from first to fifth, the following phalanges are free of web; 2; Wi; IV6; 2V6; I1 »); heel o\ the adpressed hind limb extending to the anterior corner oi the eye. Skin above distinctly granular, most o\' the granules having a corneus tip; on the posterior half of the back, the anterior loreal region, the arms, and tibial segments, the granules become larger tubercles. Below, smooth on the chin and chest; finely granular on the throat; coarsel) granular on the belly and postero-ventral aspect of thighs; one almost imperceptible furrow on each side oi the throat. Color. Above, dark brown, with two somewhat lighter and ill- defined bands from behind the eyes and over the level of the shoul- ders to near the sacral region; a dark, ill-defined streak along the flanks below the light brown band; other longitudinal, light brown spots can be noticed on the dorsum; a whitish, ill-defined spot or line under the eye. in front of the tympanum; disk of first finger white, of second finger half white and half brown; posterior portion of arm and axillar membrane white; flanks below dark brown streak white, this color extending dorsally to the illea on the pos- terior end; thighs pink, speckled and spotted with brown; rest of hind limbs and fore limb with transverse blotches or bars; web 1968 NEW SPECIES OF HYLA 3 pink. Below, creamish white, uniform, except for the sides of the throat and lower lip, which are freckled with purplish brown. The living animal appeared to be of a solid brown color with orange areas on the thighs, white forearms (posterior part) and axilla, creamish venter with purplish specks on the throat, rosy hind limbs (underneath), and with cream points along the forearm and at the sides of the anus. Measurements (in mm). Snout-vent 34.5; head length 11.02; head breadth 11.2; femur 16; tibia 17.7; foot 12.4. Paratypes. The paratypes are U.P.R.-M.1 02207-02012, all from the same locality and with the same data as the type. They vary from 21.6 to 35.2 mm in snout-vent length. In some speci- mens the heel of the adpressed hind limb may extend to between eye and nostril; in a few the web is less extensive than in the type, and the feet may perhaps be said to be not more than % webbed. The skin may be almost smooth in some of the smaller specimens. There is some variation in coloration, although the pattern is rather similar in most specimens. In a few examples the longi- tudinal bands or spots are darker, not lighter than the background color, and in the smallest individual the background color is gray and there are many short longitudinal streaks on the dorsum. In all specimens, there is a light-colored spot or line in front of the tympanum, but in some it is very diffused, and in one it extends anteriorly along the upper lip. In preserved material, the ventral color is as described for the type, except for the fact that the freckles of the throat may not be limited to the margins and may also extend to the central area. In living animals the color is more variable, and the throat may be yellow with purplish freckles while the rest of the venter is cream or lighter yellow. Sometimes there is almost no orange on the thighs, while at others the orange be- comes red and may extend, on the ventral side, to the tibial and metatarsal segments and the foot. The lower side of the forearm may also be orange or red Habitat and relationships. The place where H. rodriguezi was collected is probably one of the rainiest in Venezuela, and when it is not raining it is usually foggy and misty. The vegetation can probably be classified as the wet montane type, and there is a great abundance of epiphytes, the Aracean Philodendron nobile being particularly evident. Hyla rodriguezi was collected in ground bromeliads (apparently of the genus Vriesea) that grow along the road bank at an elevation 1 University of Puerto Rico — Mayagiiez. 4 BREVIORA No. 307 of about 1400 in. There was never more than one specimen in any one bromeliad, and onl) once was there another species of frog (Hyla minuta) in the same plant. All were collected at dusk. At night, no sound was heard in the area except that of a new species of Eleutherodactylm (Rivero, in press), and occasionally, the whistling call of Leptodactylm rugosus. This species is well characterized, and if 1 were to relate it to an) other known species, it would probabK be to the Hyla lep- rieurii group. However, Hyla rodriguezi is a smaller species with a shorter snout, less webbing between the toes, and inconspicuous vocal sacs. It is also somewhat similar to Hyla alboguttata, but the head of Hyla rodriguezi is more bom. its canthus straighter and better defined, its feet less webbed, and its color different. The speckled coloration of the throat of //. rodriguezi seems to be \ei\ characteristic of this species. Hyla grandisonae Goin (type locality, Ma/arum Forests, Guyana) agrees in main characters with //. rodriguezi, but on closer examination it is found that the two species ma\ not even be too closet) related. In //. grandisonae, the heel of the adpressed limb reaches almost to the nostril, the skin above is smooth, the ventral color is Lira\. and the species is said to be related to Hyla parviceps Boulenger, //. rondoniae Bokermann and //. bokermanni ( loin, three species that do not seem to be close to Hyla rodriguezi. LITERATURE CITED H<'KI KM \NN. \\ I KM K 1963. Duas novas espeeies de "Hyla" de Rondonia, Brasil (Amphibia, Saliemia). Rev Brasil Biol. 23 1 \)\ 247 250. 1965. lies no\os batrachios de regiao central de Mato Grosso, Brasil. Rev. Brasil Biol. 251 3 > 257-264. ( rOIN, ( hi i \i \s I960. Description of ;i new frog of the genus Hyla from northwestern Brazil. \nn Mag Nal Hist 13(2): 721-724. 1966. \ new frog from the genus Hyla from Biitish Guiana. Quart. lour \ la. Acad. Sci. 29(1): v'-42. Rivero, J. \ 1968. A new species o/ Eleutherodactylm (Amphibia, Salientia) from the Guayana Region. Edo. Bolivar. Venezuela Breviora, \lus Comp. Zool., No. 306: I -I I. (Received 26 hme 1968.) 1968 NEW SPECIES OF HYLA Bj U ON o o -c 02 . >>" 2 TO «5 w TO TO c« a, o o c ■§-§ ? c ^0 o 41 1 Q Q o <" r r C ox) u o . OJO T3 C £ o- Ml "a ^. o a CO < E BREVIORA Museum of Comparative Zoology Cambridge, Mass. 31 December, 1968 Number 308 THE MOLLUSCAN FAUNA OF AN UNUSUAL BERMUDIAN POND: A NATURAL EXPERIMENT IN FORM AND COMPOSITION Stephen Jay Gould Abstract. Twenty-four molluscan species inhabit a landlocked marine pond of normal salinity in Hamilton Parish, Bermuda. Walsingham Pond, unusual because of its isolation and the ecologic effects thereof, can be viewed as a natural experiment. The form and composition of its mol- luscan fauna are compared with "control" samples from open marine environments. Composition of the fauna is similar to two modern and three fossil samples from localities similar in ecology to Walsingham Pond but directly exposed to the open sea. Isolation does not seem to affect the presence and absence of species. Walsingham Pond molluscs are, however, unique in their shell form. As demonstrated by plots of weight vs. size, the four major Pond gastropods have thinner shells than those of the same species from control samples. It is often said that natural history is science of a lower order than experimental biology. If description and explanation are viewed as ascending orders of sophistication, this charge is valid when natural history remains at the level of "plain story"; for the very undertaking of an experiment implies a search for explanation. Yet there are a variety of natural situations that possess the essen- tial character of experiments, even though no human manipulation of material is involved. The evolutionary phenomena of parallelism and convergence, for example, serve the same function as experi- mental replication; both provide the repeated occurrences upon which inductive generalizations are based (Harris and Morren, 1966). I am concerned here with another kind of "natural ex- periment" — the unusual situation amidst a large number of "nor- mal" occurrences that act as controls. Walsingham Pond in Hamilton Parish, Bermuda, is an unusual situation. Walsingham is a landlocked marine pond of nearly nor- mal salinity maintained by tidal interchange with Castle Harbor through underground caves. Limestone walls and mangrove roots 2 breviora No. 308 form the borders of the Pond; the bottom is covered with a soft organic mud composed primarily of decaying material, faecal pel- lets, and shell fragments. Dominant elements of the bottom biota are the alga Chaetomorpha crassa, a profusion of sponges, and. unfortunately for collectors, the jellyfish Cassiopeia; there are no hard corals in the Pond. Maximum water depth is 22 feet; the bottom mud layer may be several feet thick. Helz (unpublished manuscript) studied the water chemistry of Walsingham Pond. Oxygen increases steadily from top to bottom; values are con- sistently 1 ml 1 less than those of the normal marine waters of Castle Harbor. Nitrate and phosphate show no trend with depth; values are two to three times those of Castle Harbor. This abun- dance is presumably related to the continuing concentration of organic matter in Walsingham Pond. After several rainless days. Pond salinity was 100-170 ppm higher than that of Castle Harbor. Surface waters were 10-50 ppm above bottom waters; mixing mav occur by sinking of 'he denser surface water. Ten hours after a heavy rain, surlaee waters of the Pond were less saline than those o\' Castle 1 [arbor b\ 1 500 ppm. Two aspects of the Pond fauna — composition and form — may he studied to find and explain differences between this unusual situation and the normal open marine habitats o\' Bermuda. I have confined m\ attention to the Mollusca. due both to personal preference and to the availability of abundant comparative ma- terial in the magnificent collection o\' Mr. Arthur Haycock (Ber- muda Museum). Since Walsingham Pond's landlocked status is its most unique attribute, I shall concentrate on questions related to this point. We shall want to know if the Pond's isolation, of itself, influences the form and composition of its molluscan fauna. Failing such influence, we may still ask if a secondary result of isolation an unusual condition of water chemistry attributable to curtailed mixing of Pond with ocean, for example — can be cor- related with differences between the Pond fauna and "control" faunas from normal marine environments. COMPOSITION OF THE MOLLUSCAN FAUNA Nine thousand two hundred fifty years ago, the sea stood approxi- mately 78 feet below its present level (A. C. Neumann, personal communication, determined this figure by dating a peat bed cored at the bottom of Harrington Sound). With a maximum depth of 22 feet, Walsingham Pond cannot be more than a few thousand years old. Entrance to the Pond can be obtained only through the channels of underground caves that connect Walsincham to 1968 UNUSUAL BERMUDIAN POND 3 Castle Harbor; I suspect that the molluscan populations now liv- ing in the Pond owe their origin to the infrequent transport of larvae through these channels. If this isolation exerts a major control on composition, I would expect a disharmonious as- semblage of molluscan species with a predominance of forms hav- ing long and highly mobile larval stages. If ecology is the major control, then neither the young age of the Pond nor its isolation should affect the distribution of species. In this case, I would ex- pect a molluscan fauna composed of those species that characterize similar environments in open marine areas around Bermuda. This finding would carry the implication that most molluscan species had ample access to the Pond and that the composition is a func- tion of the normal environmental preferences of species rather than of chance introductions. I have endeavored to compile a complete listing of the molluscs in Walsingham Pond. Shells were obtained both by visual collect- ing and by bulk sampling with a Van Veen grab at various depths; composition does not seem to change with depth. With the excep- tion of Chama macerophylla, which occur infrequently, cemented to the limestone walls, all species inhabit the muddy bottom. Some burrow into the mud (Vermicularia spirata), others crawl along the surface (Nassarius ambiguus), and still others (Modulus modulus) commonly live among the algal colonies of Chaetomor- pha crassa. Class Bivalvia Family Mytilidae Brachidontes exustus (very few specimens) Condylocardiidae Carditopsis smithii (one specimen) Lucinidae Codakia orbiculata (most common bivalve in Pond) Chamidae Chama macerophylla (infrequently on limestone walls) Leptonidae Lasaea bermudensis (very common) Cardiidae Laevicardium laevigatum (one immature specimen) Veneridae Gouldia cerina (very common) Tellinidae Tellina mera (one specimen) Sanguinolariidae Tagelus divisus (rare) 4 bri-viora No. 308 Class Gastropoda Family Rissoidae Alvania platycephala ( very common) Rissoina cancellata (one specimen) Vitrinellidae Vitrinella helicoidea I moderately common ) Turritellidae Vermicularia spirata (most common mollusc in Pond) Caecide Caecum delicatulum (a lew specimens) Caecum termes ( very common) Modulidae Modulus modulus ( very common) Ccrithiidae Alahinu cerithioides < quite common ) Cerithium variabile (ver\ common) Nassariidac Nassarius ambiguus (quite common) Marginellidac Hyalina avena ( rare ) Bullidae Bulla striata ( rare) Atyidae Haminoea antillarum ( rare ) Retusidae Retusa candei < moderately common ) Pvramidellidae Odostomia didyma (moderateK common) I was able to obtain samples of the molluscan fauna from five additional localities of similar environment (mangrove roots and muddy substrates), but situated in areas having free access to the open sea. These "controls" should serve to 'test the hypothesis that composition of the Walsingham Pond fauna is unusual for a mangrove-mud area because of the Pond's isolation and young age. 1. Ely's Harbour, Sandys Parish (grab sample collected at a depth of 10 feet by Robert F. Schmalz). 2. Mangrove root area on north coast of Ferry Reach, 50 yards east of Bermuda Biological station (personally collected; three Van Veen grab samples at depths of 5-7 feet). Fortunately, three small samples of fossil shells were available to provide some temporal control on the stability of community composition during the past 10.000 years. Dried mud and man- grove root fragments were present in all samples. 968 UNUSUAL BERMUDIAN POND VI snjouii C r^ E a SHJDllJUDJ VUOJV0J3W o ^H • <-* ujpqwnjoj 1-H > - DlV\\d3UVO 00 \C »— 1 o OlUlUllll c 1— 1 vuD]\dwq ed ■~ *TJ 2 t3 VJL31U ON T3 3 O DUUPI s 13 o l-l snjsnxj 00 c- fl E xjjuopupv.ig SHSlAip c/3 =5 >n snjaSvx f- o 00 Ph X> ,_ o s T3 DJDUJS ON r- IXI o CG m»a —J l-i -3 OQ < snuSiquiD 1— «4-< O c o c SIlUDSSDtf snjnpoui «-> >* NO V) ■* (N r«") in m o c c 3 X> SlipipOJAJ niapioiqjo p DiyDpoj m -fr '"H ro "t X) > < 3[iqVUVA > lumqiud^ cs CS (S -. CO £ 5 ° a w o U T3 O £ w ca W °C/3 CO o Pm o o Ph 6 breviora No. 308 3. Pembroke Marsh, Pembroke Parish (collected by Charles Schuchert of Yale University in the 1920's). Of the five controls, only this sample cannot be identified as having been in open contact with the sea. 4. Longbird Bridge boring. 71.8 feet below present sea level. 5. Longbird Bridge boring. 85.5 feet below present sea level. Samples 4 and 5. kindly supplied by Walter S. Newman, were obtained from the test boring made prior to construction of Long- bird Bridge, St. Gcoriic's Parish. Table 1 lists all the mud-dwelling macromolluscs of Walsingham and these five localities in order of their abundance (only the rock- cementing Chama macerophylla is eliminated by this constraint on environment). I do not know how representative the small fossil samples are of actual diversity and abundances. The micromol- luscs are not included, because several oi the samples contained only the larger shells, with insufficient matrix for the screening of small species. Yet strong similarities almost surely exist between micromolluscs of Walsingham and the five controls. Several speci- mens oi Retusa candei, Caecum termes, and Odostomia didyma, for example, were obtained from dried mud caking the apertures of Vermicular ia spirata specimens in the Longbird 4 sample; all three of these micromolluscs are common in Walsingham Pond. The similarities between Walsingham and the controls are strik- ing. Five macromolluscs strongly dominate the Walsingham fauna — Vermicular ia spirant. Cerithium variabile, Codakia orbiculata, Modulus modulus, and Nassarius ambiguus, in that order. Except- ing the absence of Codakia in the Ely's Harbour sample, all of these species are present in all oi the five controls. Considering the small si/e of several samples (20 shells in Longbird 5. for ex- ample), this correspondence seems highly significant. Bulla .striata and Tagelus divisus, rare but consistently present in Walsingham. are found in three and two of the controls respectively. Of Brachi- dontes exustus and Tellina mera, the Walsingham "trace ele- ments." only the first is found at one other locality. A few unexplained discrepancies arise from species present in control samples but not in Walsingham Pond. The venerid clam Chione cancellata is present in three of the controls, dominant in one of these (Longbird 4) but absent from Ely's Harbour and Longbird 5 as well as from Walsingham. Batillaria minima, a homcomorph of Cerithium variabile, commonly occurs with the latter in rocky and sandy intertidal environments of the open sea (they are so associated in Castle Harbour, barely 100 yards from Walsingham Pond). Batillaria is absent from all but one of the 1968 UNUSUAL BERMUDIAN POND 7 mangrove-mud environments, but dominates among the mangrove roots east of the Biostation. This might argue for atypicality of the Biostation locality, but not for a lack of correspondence between Walsingham and the other controls. While it is not especially rele- vant to this study, the ecologic relationship of the homeomorphs Cerithium and Bat Maria is a fascinating problem.1 Thus, the association of Vermicularia spirata, Cerithium varia- bile, Codakia orbiculata, Modulus modulus, and Nassarius am- biguus in Walsingham Pond is no product of accidental access, but an expression of the normal molluscan community in mangrove- mud areas. Vermicularia is not often thought of as an inhabitant of muddy areas, for it usually lives cemented to hard objects in clear, open water. Olsson (1951), however, found abundant specimens on the mud flats in Tampa Bay, Florida. He described them as Vermicularia fargoi, but I have synonymized this species with V . spirata elsewhere (Gould, in press). Tagelus divisus prefers muddy environments (Fraser, 1967). In short, the unique isola- tion of Walsingham Pond is not reflected in the composition of its molluscan fauna. Characteristic species of the molluscan com- munity that inhabits mangrove-mud areas throughout Bermuda have all become established in Walsingham Pond despite the bar- riers to distribution. FORM OF THE MOLLUSCAN FAUNA Is the form of molluscan shells in Walsingham Pond different from that of the same species in open marine habitats? If such differences are found, we may again pose the question: Is the uniqueness of Walsingham molluscs attributable to the Pond's iso- lation per se or to some secondary ecologic result of that isolation? In the first instance, isolation of a small founding population main- tains the distinguishing gene frequencies present, by chance, in the founding individuals or attained, without regard to adaptive sig- nificance, by genetic drift. In the second case, an unusual condi- tion of water chemistry or bottom ecology works its influence upon the form of shells by direct phenotypic modification. A mixed case is also likely: unusual conditions establish strong selective pres- sures, while isolation favors the stabilization of new genotypes by 1 Where they occur together, distribution is not random. Patches of 20-30 individuals of one species are often found in large tracts dominated by the other. This phenomenon is particularly well seen in Coot Pond, St. George's Parish. 8 BREVIORA No. 308 removing the diluting effect o\' gene influx from normal popu- lations. When a single speeies differs from its 'control'" counterparts m a unique way, it may be difficult to distinguish among the three explanations. Walsingham specimens of Modulus modulus are often relatively higher than is normal for the species in Bermuda. The most spectacular modification, however, is found in Vermicu- laria spirata. This gastropod customaril) begins life as a normally coiled turritellid, but later cements to a hard object and uncoils its shell. In the absence of any hard substrate on the floor of Wal- singham Pond, more than 95 per cent o\' the Vermicular ia shells do not uncoil at all; those that do unwind slowly and regularly without a trace oi the normal attachment scar. We have here the curious case o\' an animal reverting, functionally, to the form and habitat oi its ancestor: Walsingham Vermicularia construct and inhabit burrows identical to those o\ its coiled ancestor Turritella. Since the Walsingham Vermicularia uncoil normally when pro- vided with hard coral substrate in a laboratory tank, this striking modification may have no genetic base other than the great pheno- ls pic plasticity permitted b\ the normal genotype. I have docu- mented this ease in some detail elsewhere (Could, in press). When many species differ from their "controls" in a similar way. some aspect o{ the Pond's unique ecolog) is probably involved in the explanation — either directly by phenotypic modification 01 indirect!) via selection pressures. There is one general modifica- tion that affects all the major Walsingham species: Pond shells appear to be thinner than those o\ the same species in open marine habitats. I tested this obser\ation by comparing plots oi combined linear dimensions vs. weight for the four most common Pond gastropods (Fig. 1) with similar plots for the same species in normal marine habitats (Haycock collection. Bermuda Museum ). Use of a combined linear dimension (length plus width) provides a better measure o\ overall size than length alone, fin Modulus modulus, for example. Pond shells of a given length are generally narrower — hence lighter — than are control specimens of the same length. The lower weight length ratios of Pond Modulus at a given length might be due not to shell thinness [the hypothesis to be tested] but to differences in shape between Walsingham and control shells. | In the ideal case of isometric growth, shell weight varies directh with shell volume, and shell volume increases as the cube of a single linear dimension or an added combination of such measures (using length plus width instead o\ length alone will modify the 1968 UNUSUAL BERMUDIAN POND mm Fig. 1. The four dominant gastropods of Walsingham Pond (natural size). Upper left: Cerithium variabile. Upper right: V ermicularia spirata. Lower left: Modulus modulus. Lower right: Nassarius ambiguus. y-intercept but not the slope of this ideal weight vs. linear dimen- sion curve ) : weight = b (length plus width)3 Such a curve will plot as a straight line on double logarithmic paper with a slope of 3 and a y-value of b at x = 1. This situation is closely approached in the eight plots of Figure 2: slopes vary from 2.57 to 3.53. In three of the four pairs of plots, slopes are similar for Pond and control samples (Table 2). The high slope of Wal- singham Modulus may be an artifact of a small and incomplete size range — see Bohlken, 1962:560. Pond b-values are lower than the control parameters in each case (Table 2), but this is not significant in itself, since x = 1 mm is too distant an extrapolation from the data size range to be biologically meaningful. The im- portant point (Fig. 2) is that for each of the four pairs of plots, the curve for Pond snails falls below that for control snails over the entire measured size range of the samples. (The intersection of the two Modulus curves at a size never reached by this species is a datum devoid of biological significance; the size range of actual data should always be specified in power function plots — see White and Gould, 1965.) At any given size for which data are available, 10 hreviora No. 308 Pond snails arc lighter than control snails in each of the four species. Similar modification of at least four species suggests that some common factor of Pond ecology is involved as direct or indirect cause. Due perhaps to the vast accumulation of decaying organic matter in Walsingham. Pond pH is a good deal lower than normal marine values. Six measures from various depths made in June 1967 ranged from 7.6 to 7.7. There is some suggestion in the literature (Okland, 1964: 190) that low pH might inhibit shell formation even in the presence oi sufficient dissolved calcium in the water; most Bermudian waters are supersaturated with respect to calcite (Schmalz and Chave, 1963). The lowered pH ma\ make it metabolicallv more difficult to deposit available calcium; a given expenditure of cnerg\ may therefore produce a thinner shell in the Pond than in open marine habitats of normal oceanic pll. (We are. indeed, dealing with a deposition phenomenon rather than subsequent shell solution in Pond snails; only uneroded snails were used in these analyses. | TABLE 2 Reduced Data ior Weight vs. Size Relationships in Four Gastropod Species: Comparison of Walsingham and Control Samples. Weight b (Height -\ Width )K /»( /;//// ) .0123 .0115 .0356 .0348 .0332 .0055 .0303 .0263 Nun ihcr of Specimens Slope(l I . Cerithium variabile Control 20 3.24 Walsingham 10 3.14 2 Vermicularia spirant Control 20 2.65 Walsingham 15 2.58 3. Modulus moil ul us Control 20 3.00 Walsingham 13 3.53 4. Xassarius ambiguus Control 20 2.90 Walsingham 8 2.78 1968 UNUSUAL BERMUDIAN POND 11 300 100- 30 X O 300 100 30 10 20 H+W CERITHIUM 30 10 20 H+W 30 VERMICULARIA 300 100 30 r o 300 10 20 H+W MODULUS 30 NASSARIUS Fig. 2. Curves of weight (mg) vs. length + height (mm) for the four snails of Figure 1, both in Walsingham Pond and in normal marine habitats. Regressions marked A are for control snails, B for Pond snails. Regression lines extend only over the range of size and weight covered by actual specimens. Walsingham curves all lie below control curves, demonstrating that Walsingham shells of all species are lighter at any given weight. See Table 2 for numerical data. 12 mrkviora No. 308 I cannot state whether the thinner shells of these four species are a direct phenotypic response to unusual conditions or a result of parallel evolution, but I favor the former explanation, which is based on both the \outh of Walsingham Pond and general notions of simplicity. Extreme phenotypic plasticity in shell thickness has long been recognized in land snails. H\ adding ground chalk to the diet of Helix aspersa, for example. Oldham ( 1934) found that shell weight increased fourfold over snails deprived oi calcium. Thus Walsingham Pond serves as a natural experiment in the form and composition of moliuscan communities. The unique iso- lation of the Pond has no apparent effect upon the presence or absence of species but influences, in similar ways, the shell form oi its dominant mollusean populations. ACKNOWLEDGMENTS I am deepl) grateful to a number oi people: to Mr. and Mrs. Rutherford, owners ol Walsingham Pond, for their interest, hos- pitality and permission to collect on their property; to R. Jensen, our leading student oi Bermudian molluscs, for help in identifying the micromolluscs (terra incognita for me); to L. Mowbrey, di- rector oi the Bermuda Aquarium and Museum, for placing the Haycock collection at my disposal; to R. Schmalz for collecting the Eh's Harbour sample: to W. Newman for supplying the Long- bird Bridge samples; to C. MacClintOCk for loaning the Pembroke Marsh sample; to A. C. Neumann for supplying the date on the Harrington Sound peat bed; to K. J. Boss for giving me free run of the library and collections oi the Mollusc Department at the Museum oi Comparative Zoology; and to R. Houston and C. Jones for drafting the illustrations. This paper is contribution — 437 from the Bermuda Biological Station. LITERATURE CITED Bon; m n. H. 1962. Problcme der Merkmalbewertung am Saugetierschiidel dar- gestellt am Bcispicl der Bos primigenius Bojanns 1827. Morph. Jb.. 103: 509-661. Kraser, T. H. 1967. Contribution to the biology of Tagelus divisus (Tellinacea: IMecypoda) in Biscay ne Bay, Florida. Bull. Mar. Sci., 17: 111-137. 1968 UNUSUAL BERMUDIAN POND 13 Gould, S. J. In press. Phenotypic reversion to ancestral form and habitus in a marine snail. Nature. Harris, M., and G. E. B. Morren 1966. The limitation of the principle of limited possibilities. Am. Anthrop., 68: 122-127. Okland, J. 1964. The eutrophic lake Borrevann (Norway) — an ecological study on shore and bottom fauna with special reference to gastropods, including a hydrographic survey. Folia Limn. Scan., 13: 1-337. Oldham, C. 1934. Further observations on the influence of lime on the shells of snails. Proc. Malacol. Soc. London, 21: 131-138. Olsson, A. A. 1951. New Floridan species of Ostrea and Vermicularia. Nautilus, 65: 6-8. SCHMALZ, R. F., AND K. E. CHAVE 1963. Calcium carbonate: factors affecting saturation in ocean waters off Bermuda. Science, 139: 1206-1207. White, J. F., and S. J. Gould 1965. Interpretation of the coefficient in the allometric equation. Am. Nat., 99: 5-18. (Received 4 June 1968.) BREVIORA Museumi of Comparative Zoology Cambridge, Mass. 31 December, 1968 Number 309 VARIABILITY OF RODENT INCISOR ENAMEL AS VIEWED IN THIN SECTION, AND THE MICROSTRUCTURE OF THE ENAMEL IN FOSSIL AND RECENT RODENT GROUPS John H. Wahlert Abstract. The microstructure in enamel of various fossil and Recent rodent incisors is described from thin sections. Measurements of enamel thickness and inclination of bands in the inner enamel layer were made from sagittal sections on large and small samples of Rattus norvegicus and on small samples of the Eocene forms Paramys copei and Knightomys depres- sus. Statistical analyses of these data show that a sample of size 10 yields an adequately close approximation to the population mean. Differences in enamel dimensions between upper and lower incisors and between incisors of different species are apparent. Thickness and inclination can be used in the identification of isolated incisors when data for other rodents have been compiled. Band width is of apparently similar utility, but small size pre- cludes anything more than rough measurement at 430 diameters magnifica- tion. Korvenkontio's "external index" and other measurements taken from sagittal sections are deemed unreliable. Rodent species whose enamel has been studied are listed according to the kind of enamel they possess. Pau- ciserial enamel, found only in Eocene and Oligocene forms, appears to be the structural predecessor of uniserial and multiserial enamels, which occur in all post-Oligocene rodents examined. At present, the microstructure of incisor enamel is useful as an additional character for determining the systematic position of rodent higher taxa, but it is no touchstone. INTRODUCTION That incisor enamel may be useful in the classification of ro- dents was first suggested by the work of Tomes (1850) and later by that of Korvenkontio ( 1934). Tomes examined the microstruc- ture of incisor enamel in a number of living rodents and found that within supposedly related groups the arrangement of enamel prisms is similar; his descriptions and figures showed that the enamel of modern rodents is of two basic kinds. Korvenkontio named these uniserial and multiserial. His extension of the research to other BKl \ l()k\ No. 3(W Figure 1. Sagittal sections of enamel in lower incisors: a. Rattus nor- vegicus; b. Metaphiomys schaubi; c. Paramys copei. Tips of incisors are to the right. Magnification is approximately 240 diameters. Abbreviations: o, outer enamel laser, i. inner enamel layer showing hands: d. dentine. 1968 RODENT INCISOR ENAMEL 3 modern rodents and to various fossil forms greatly enlarged Tomes1 work and led to the discovery of a third kind of incisor enamel structure, which he named pauciserial, among early fossil forms. In all rodents enamel covers the outer, labial side of the incisors; it may extend slightly on the mesial and distal sides. Korvenkontio published an extensive table (1934: 116-123) giving the dimensions of various parts of the enamel and indices derived from these measurements for the incisors of all the species he examined. These suggested the possibility that some dimen- sions and ratios, at least, might be characteristic of low level taxa. If this should prove to be true, it would be a relatively simple mat- ter to section, measure, and identify fossil rodent incisors, which are so frequently found separated from their jaws. Korvenkontio states (1934: 125, footnote) that in his study of 72 Recent and 33 fossil forms he used about 520 sections. He attempted to make sagittal, transverse, and frontal sections of each species wherever possible, and thus his sample size for measurement was very small. The primary purpose of the work here reported has been to de- termine the variability of enamel and the sample size needed to obtain meaningful measurements. In the course of the work I have had occasion to section incisors of a number of rodents, both Recent and fossil. These are included in Table 5, which lists all species whose incisor microstructure is known. THE KINDS OF RODENT INCISOR ENAMEL A sagittal section of uniserial enamel, for example that of Rat t us norvegicus (Fig. la), reveals two layers of enamel. The inner layer in this aspect appears to consist of bands one enamel prism wide that extend outward from the dentine and upward toward the tip of the tooth. On close examination each band is seen to be divided into small units, somewhat like a string of beads. Study of transverse and frontal sections reveals that each band is the cross- sectional view of a transverse lamella of enamel prisms. The width of bands and prisms is the same in sagittal section; Korvenkontio's data show a range of 2.2 to 5.0 microns for uniserial enamels. The prisms in a single lamella are parallel; they do not make a right angle with the dentine but are at some oblique angle to the sagittal plane of the tooth. A band is, therefore, the cross-section of prisms comprising a lamella. The prisms of every other lamella have the same orientation. There are two kinds of differences in orientation of prisms of adjacent lamellae. In transverse section the prisms of adjacent lamellae are seen to cross each other at a fairly constant angle. Frontal sections reveal that the prisms are 4 BREVIORA No. 309 tilted to one side o\' the sagittal plane in ever) other lamella and to the other in the alternating set. In the outer layer of the enamel the prisms of all lamellae are parallel, and lamination ceases; they are usually inclined more steepl) toward the tip o\' the tooth than in the inner layer. A sagittal section o\' multiserial enamel, lor example that o\ Metaphiomys schaubi (Fig. lb), also reveals two lasers of enamel. The hands o\ the inner layer are many prisms wide. Kor- venkontio found that band widths range from 12 to 30 microns for multiserial enamels. Prisms are directed obliquely with respect to the direction oi the bands; the obliquity differs in adjoining bands but corresponds in alternate ones. Prisms oi the outer layer are parallel and more steepl) inclined toward the tip of the tooth. as in uniserial enamels. The three-dimensional structure is more complicated than in uniserial enamels, and for m\ own work 1 have relied on Tomes' description ( 1850: 552-53), which 1 para- phrase closel\ here: In an oblique transverse section parallel with the course o\' the enamel lamellae, the inner enamel layer looks as though the prisms were thrown into waxes, the furrows oi which commence at the surface of the dentine and. proceeding obliquel) outwards, crop out where the prisms become parallel in the ex- ternal laser. The prisms pursue a serpentine course in the inner, lamelliform portion, where the) describe tolerablx uniform curves. B) altering the focus o\ the microscope it max be seen that the prisms of adjacent layers pursue a similar serpentine course, but are arranged so that the concavities and convexities point in oppo- site directions, thus producing a sort oi figure X. Pauciserial enamel, for example that oi Paramys copei < Fig. Ic). is also divided into two layers. The prisms oi the inner layer may or max not be organized into lamellae. In a sagittal section the bands of lamellar enamel are seen to be of somewhat variable width, depending on the number of prisms included, korxen- kontio found that band widths o\' pauciserial enamels fall between 5 and 16 microns, between the ranges for uniserial and multiserial enamels, with slight overlap. The structure appears similar to multiserial enamel but lacks the uniformity of organization. Bands are usually not inclined. Prisms in the external portion are slightly inclined toward the tip of the tooth, though this is not visible in the figure. Some pauciserial enamels have fewer prisms per band, and thus the bands are narroxver. sugges'ing uniserial enamel in appearance. These descriptions of the kinds oi enamel seen in sections of rodent incisors refer only to the xicinity of the midline of the 1968 RODENT INCISOR ENAMEL tooth, where the structure is clearly seen. The terms uniserial, multiserial, and pauciserial apply, of course, to the inner layer of the enamel only. MEASUREMENTS AND INDICES Sagittal sections are the best for determining what kind of enamel is present in a given rodent incisor, and they are the easiest to make; therefore, I have considered only measurements made from them. Total enamel thickness and thicknesses of the two enamel layers can be measured. Korvenkontio's external index, which is the per- centage of total enamel thickness occupied by the outer layer, compares these dimensions. Total enamel thickness is defined as the length of a line segment normal to the enamel surface at a point, in the vicinity of the midline of the tooth, at which the enamel is thinnest. If a sagittal section contains this segment, it occurs where the enamel is unworn and thinnest. The geometry of rodent incisor enamel distribution, as seen in transverse section (Fig. 2a and b), makes underestimation of the total enamel thick- ness impossible in a reasonably good section. I mm Figure 2. Transverse sections of upper incisors: a. Rattits norvegicus; b. Paramys copei. Mesial side is to the right. 6 BREVIORA No. 309 Band width and inclination oi bands arc also measured best in sagittal section. Korvenkontio defined inclination as the angle at which bands intersect a perpendicular to the dentine in the sagit- tal plane.1 When bands appear curved, a tangent to the midpoint o\ a band is used to determine inclination. Prisms in the external laser also have an inclination that can be measured; this is not visible in most sections, however, and I have not considered it. In this paper 1 use enamel thickness, external index, band width, and inclination in the internal enamel laser according to Korvenkon- tio's definitions for sagittal sections. MATERIALS [ncisors ol 48 individuals from a highl) inbred strain of albino Rattus norvegicus were available to me at the beginning of this project. Ihis is a large sample tor statistical analysis by paleon- tological standards. I he heads had been preserved in formalde- hyde, and sexes were not recorded. Incisors of Paramys (<>i>ci and Knightomys depressus were also available m lair quantity. I hese had been collected by Amherst College parties from the earl\ I ocene I ysite member of the Wind River Formation in the Wind River Basin. Wyoming. Individual incisors from a variety ol rodents were also sectioned; specimens were obtained from the col- lections of Amherst College, Albert E. Wood, the Museum of Comparative Zoology, the University ol lexas. ami the Yale Pea- bod\ Museum. PROCEDURE Standard sectioning techniques were used. All lour incisors from each rat were sectioned on one petrographic slide. Fossil incisors a\k\ individual modern incisors were sectioned singly. Preliminar) grinding was done on a water-bathed 600 grain dia- mond wheel that turned at approximate^ 100 rpm. Sections were hand held. Final polishing was done with wet 900 gram alundum on wood, and then sections were etched briefly with dilute hydro- chloric acid; this step makes the enamel structure visible. When sections of modern teeth become thin, water absorption causes them to buckle. This difficulty was solved by drying the section before it became thin and then impregnating it with mineral oil which acts as a waterproofing agent. ■Tomes (1850) measured inclination as the angle made b) bands and the dentine surface. 968 RODENT INCISOR ENAMEL 7 Thickness measurements were made by comparing the projection of a thin section with a micrometer slide. The number of bands within a standard unit of a micrometer eyepiece divided by the length of the unit in millimeters yields the average band width. Inclination was measured on the rotating stage of a petrographic microscope. In each case 430 diameters magnification was used. Tabulation and statistical analyses of the measurements are given in Tables 1-4. The formulae used in computation are those given by Simpson, Roe, and Lewontin (1960: 84, 90, 166). Ab- breviations are as follows: N, sample size; OR, observed range; % 3 0 .080 .090 .100 140 .150 .160 % 50 40 30 20 10 upper — lower Figure 3. Frequency distributions for incisors of Rattus norvegicus'. a. total enamel thickness; b. enamel prism inclination. 8 BREVIORA No. 30l) X, arithmetic mean: Conf. Int.. confidence interval of the mean; SD. one standard deviation unit; V, coefficient of variation. In all thickness measurements there is an error in estimation of ±.001 mm; this error is inherent in the mean. Frequency distributions for enamel thickness and inclination of bands in the large sample of albino laboratory rats appear in Figure 3a and b. Although all four incisors of forty-eight rats were sectioned, not all individual sections were usable, and thus the sample size varies slightly. Inclination was not measured in Paramys copei and Knightomys depressus because prism bands are not well defined and are nearly perpendicular to the dentine surface. The sample M/e in both cases was too small for frequenc\ diagrams to be constructed. Measurements of band widths were, at best, crude, because boundaries are not well defined and magnification was low for such small objects. Consequently, I present only average width and range under the appropriate headings. TABLE 1 Data for albino laboratory rat incisors. total enamel thickness (in mm) 959! Ineisoi N OK \ < onf. Int. SD V Upper 9ft .080-. 126 ii')-* ±.002 .009 X.7 Lower 93 .100-. 155 .129 ±.003 .012 9.1 Inclination of enamel prisms ( in decrees ) Uppei 95 23-42 36 ±.6 3 8.3 Lower 93 J2-51 46 ±.6 3 6.1 DESCRIPTION OF SAMPLES Rutins norvegicus (laboratory strain). The incisor enamel is uniserial (Fig. la), and the enamel thickness is greater in the lower than in the upper incisors o\ each individual rat. although the ranges of total enamel thickness for upper and lower incisors over- lap appreciably (Table 1 ). In only one specimen were the meas- urements within the error o\ estimation of each other, the enamel thickness of the upper right incisor being .126 mm and of the lower left. .127 mm. The overlap of ranges shows that it is unsafe to decide whether upper or lower incisors have the thicker enamel on the basis of only one or two specimens. The frequency distri- bution is graphed in Figure 3a. 1968 RODENT INCISOR ENAMEL 9 The 95 per cent confidence interval for the mean is quite small in relation to the total thickness. The standard deviations are greater for lower incisors than for uppers. The coefficients of varia- tion are high for a Recent species (Simpson, Roe, and Lewontin. 1960: 92). This may be accounted for by deviation of the section from the sagittal plane and also by natural variation, such as effects of age, sex differences, and inbreeding, none of which can be ex- cluded as a factor contributing to a high V. The distribution of enamel on the front of an upper incisor as seen in transverse sec- tion (Fig. 2a) is uneven; thus, deviation of sections from the sagittal plane could be the cause of considerable variation in thick- ness measurements. The enamel distribution is more uniform in lowers. Separate analysis of left and right incisors yielded the same mean in lowers and a difference of means in uppers that is within the estimated range of observational error. The number of incisors of a single extinct species collected from one locality is usually small, on the order of ten or so. To test the reliability of enamel thickness measurements for small samples, I considered the data for forty-eight rats as representing an entire population with a known mean and took from it ten random samples of ten upper and of ten lower incisors. The results of the analyses are presented in Table 2. Although the means of these samples may be as much as five thousandths of a millimeter from the mean of the population, the ninety-five per cent confidence in- tervals of the means for all but one sample (upper incisors, sample 8) include it. This same sample has both the highest mean thick- ness and the least variability. With any two samples representing upper and lower incisors, the relative difference in enamel thick- ness is apparent; the smallest difference in mean enamel thickness obtainable from these results amounts to .020 mm and the largest is .039 mm. Attempts to measure the thickness of either layer of the enamel separately were unsatisfactory, because there is no clear line of demarcation between the two. Figure 1 shows how bands of the inner layer project into the outer; it is impossible to establish a definite line that is consistently placed in each thin section. In the- sample the outer enamel layer occupies approximately 30 per cent of the mean total thickness in uppers and 19 per cent in lowers. The observed ranges are 20 to 39 per cent for uppers and 14 to 27 per cent for lowers. In two individuals the external index of an upper and a lower incisor is the same, but in no instance in an individual is that of the lower greater. A regression analysis of these measurements of outer layer thickness versus total enamel 10 HKI-.VIORA No. 31W TABLE 2 Variability of total enamel thickness (in mm) in ten samplings, each of ten incisors selected at random from data tor albino labora- tory rats. Upper incisors 95 S.uiip. OK \ ( onf. Int. SI) V 1 085-.120 101 ±.007 .010 : 080 126 .099 ±.009 01 i 13.1 ) 085 .110 .097 ±.006 008 8.2 4 OS5-.IK) 099 ±.005 .007 7.1 5 092-.109 .099 ±.004 ,006 6 I '• 094-.106 100 ±.003 .004 4.0 7 080-.1 12 095 ±.006 .009 9 s 8 [00 .107 .104 ±.001 .002 1.9 9 085-.126 .102 ±.009 .013 12.7 in 092-.1 10 101 ±.004 .006 5 9 1 owei incisors 1 1I0-. 155 .129 ±.01 1 015 1 1.6 : 1 I9-.153 .134 ±.008 .01 I 8.2 5 1 10-143 .127 ±.007 .010 7.9 4 [07- 143 .128 ±.007 .010 7.S 5 11 1-.138 .127 ±.007 Din 7.9 6 II 4-. 153 .129 ±.008 .01 1 8.5 7 I1I-.I45 .124 ±.007 .010 x.l 8 I07-.I5O .131 ±.01 1 .015 11.4 9 1 07-. 138 .126 ±.008 .01 1 8.7 10 1 10-.145 .129 ±.008 .012 9 ; thickness was carried out within upper and lower incisors of the entire sample. The correlation coefficient for upper incisors is .27; for lowers. ,18. This is extremely poor, but the gross difference between uppers and lowers remains. Band width in sagittal section appeared ncaii\ invariable at 450 diameters magnification. It is .0033 for both upper and lower in- cisors. The mean inclination of enamel prisms in the inner layer is distinctly greater in nearly all lower incisors (Table 1. Figure 3b). However, in one specimen the inclination of bands in ov^- lower incisor was less than that of the uppers; in another, an upper and a lower incisor yielded the same angle. Thus, two teeth, even I 95% OR X Conf. Int. SD V 088-.115 .104 ±.004 .008 7.7 065-.105 .088 ±.004 .009 10.2 1968 RODENT INCISOR ENAMEL 11 from the same individual, may not indicate the usual relationship. The standard deviation of inclination is the same for both upper and lower incisors. TABLE 3 Data for Paramys copei incisors. Total enamel thickness (in mm) Incisor N Upper 1 6 Lower 24 Paramys copei. The sample of incisors of Paramys copei from the Lysite represents a minimum of twelve individuals, all but two of the teeth being from the same locality. Enamel is pauciserial. In this species the distribution of enamel across the face of the tooth as seen in transverse section is fairly even (Fig. 2b). The error caused by the deviation of a section from the mid-plane is therefore not large when compared to the same situation in rat incisors. The data presented in Table 3 indicate that the mean enamel thickness in upper incisors is .016 greater than in lowers, the re- verse of the situation in rats. There is overlap in ranges, and examination of only a few teeth could show the opposite relation- ship. The standard deviations for upper and lower incisors are similar to the figure for upper incisors of rats. The sample is too small to construct a meaningful frequency distribution curve. The mean band width and observed range are .017 mm and .01 3-. 021 mm in upper incisors and .016 mm and .014-.023 mm in lowers. Inclination of bands is approximately zero. In any in- dividual tooth, bands may be slightly inclined toward or away from the tip. TABLE 4 Data for Knightomys depressus incisors. Total enamel thickness (in mm) 95% Incisor N OR x Conf. Int. SD V Upper 13 .103-.150 .120 ±.009 .016 13.3 Lower 6 .053-.095 .068 ±.005 .005 7.35 Knightomys depressus. The sample of Knightomys depressus from the Lysite represents a minimum of seven individuals. Enamel is pauciserial. The data presented in Table 4 show that the mean 12 BREVIORA No. 309 enamel thickness is much greater in upper incisors and that the ranges for upper and lower incisors o\o not overlap. The lack of overlap mav simplv mean that the sample size is too small. The confidence interval, standard deviation, and coefficient oi variation tor the lower incisors seem low with regard to these figures tor the other samples. 1 hey are in best agreement with those tor small samples of rat upper incisors I he distribution of enamel is similar to that in Paramys copei. 1 he mean hand width and observed range are .014 mm and .012- .017 mm in upper incisors and .012 mm and .009-. Old nun in lowers. Inclination oi bands is approximate!) zero, and the boun- dary between inner and outer layers of enamel is again not a distinct line. SAMPLE SIZE AND THE UTILITY OF MEASUREMENTS AND INDICES AS CRITERIA FOR IDENTIFICATION OF ISOLATED INCISORS A sagittal thin section of a rodent incisor is sufficient to show whether the enamel is pauciserial, uniserial. or multiscrial. A lew sections provide a rough measure of enamel thickness. A sample of ten or more upper and ten or more lower incisors yields a mean enamel thickness with a fairly small confidence interval and shows the relationship oi thicknesses. A sample o\ about a hundred individual upper and lower incisors is needed for graphing a frequency distribution oi thickness. The inclination oi bands in the inner enamel layer also vields a continuous curve when plotted lor a sample oi this size. I lie inclination oi prisms in the external layer is usually not visible. A rough measure of band width mav he made at a magnification oi 430 diameters, but variation is not apparent. The boundary between internal and external enamel layers is not a line, and thus the thickness oi either part cannot be measured precisely. This variability and the lack of correlation between thickness oi the outer layer and total enamel thickness within a sample suggests that a ratio of the two, Korvenkontio's external index, cannot be used. At present there are few numerical data available concerning the microstructure oi rodent incisor enamel. Identification of isolated incisors cannot be made on the basis of measurements alone. DISCUSSION OF INCISOR ENAMELS Pauciserial enamel was found by Korvenkontio to be present in ischyromyid. sciuravid. pseudosciurid. and some theridomyid 1968 RODENT INCISOR ENAMEL 13 rodents. I have sectioned incisors of some of the same Eocene species; these and thin sections of different ischyromyid and sciura- vid species reveal pauciserial enamel. Prosciurus relict us, a mid- Oligocene ischyromyid has uniserial enamel. Korvenkontio found a condition transitional between pauciserial and uniserial in the Oligocene theridomyid, Nesokerodon minor. Completely uniserial and multiserial enamels are first met with in incisors of early Oligocene rodents. Korvenkontio found uni- serial enamel in species of this age in the Theridomyidae and Ischyromyidae, and I have observed it in Cylindrodon fontis of the Cylindrodontidae. Other species having uniserial enamel are listed in Table 5. Multiserial enamel is characteristic of all caviomorph rodents so far examined. Wood and Patterson (1959: 292)- found it in a Deseadan (early Oligocene) incisor, very probably of Scotamys antiquus, and I have found it in Cephalomys arcidens from deposits of the same age. The early Oligocene African phiomyid rodents sectioned by me had acquired multiserial enamel; thin sections of incisors of Phiomys andrewsi, Metaphiomys schaubi, and a new species (Wood, in press) demonstrate this. The Recent Thryono- mys and Petromus, which Lavocat (1962) and Wood (in press) associate closely with the phiomyids, also prove to have multiserial enamel. Other species possessing multiserial enamel are listed in Table 5. Pauciserial enamel is a good structural predecessor for uniserial and multiserial enamels. It would appear to be the ancestral con- dition. As far as present evidence goes, it had probably been achieved by the time rodents made their first appearance in the record. It is known thus far in only a few post-Eocene rodents, and uniserial and multiserial enamels have not yet been recorded prior to the Oligocene. Presumably acquisition of two layers in which the organization of prisms is different served in some way to strengthen the enamel. The pauciserial structure, judging from the record, would appear to have been less efficient than the other two. There is no available evidence to suggest that one of the two surviving kinds is superior to the other or that one of them has ever evolved from the other. Seemingly, selection has favored increase in strengthening rather than any one method of accomplishing it. To the possible question: could the pauciserial condition be an artifact caused by diagenetic changes in materials of greater age? 2 A lapsus in this paper may be corrected. The authors state (p. 292 n.): ". . . the Theridomyidae are in the process of passing from a pauciserial to a multiserial type." For "multiserial" read "uniserial." 14 BREVIORA NO. 309 I would repl) in the negative. In none of the many slides of enamel of I ocene age that I have examined is there an\ indication that recrystallization has disrupted the tine structure. The enamels of all modern rodent incisors tall into one of the two major categories, uniserial and multiserial, but there are minor differences within each of the two as regards dimensions and orien- tation o\ lamellae. Stud) of the internal detail of lamellar structure is needed to understand the differences and to determine how the) ma) have evolved. ACKNOWLEDGMENTS I began this research at Amherst College and received support in 1964-65, through the Ceolog\ Department, from the Student Research Participation Program of the National Science Founda- tion. The work was continued in the summer o\ 1965 under Pub- lie Health Service (.rant 2 II (.\1 M)^-()h to the Biolog) Depart- ment of Amherst College, and other help was forthcoming from National Science Foundation Granl GB lc>77 to Professor Albert E. Wood. I am indebted to Professor Wood and to ProfeSSOl Bryan Patterson for their guidance, and to Miss Barbara Lawrence and Mr. Charles Mack of the Mammal Department. Museum of Comparative Zoology, for permitting me to section various speci- mens. TABLE 5 I name] structure in rodent incisors. ["he nun. ils aftei each species name .ne those of the authors who have described its incisoi enamel Species names are mainl) as in I Herman < 1441 ): familial and generic assignments are mainl) as in Simpson i 1945 I. Where a name given in the publications of those who have described rodent incisor enamels differs from the current one. I have included it in brackets. \ classification ol rodents above the familial level is given b) Woo. I i 1958) Abbreviations: B., Bohlin (1946); k. Korvenkontio (1934); I., lomes i I smi i ; \V . Wahlert; W. and P.. Wood and Patterson ( 1959). I OC., I ocene: Olig., Oligocene; Mioc, Miocene; Pleist., Pleistocene; I., early; \1 . mid; L., late. \f. Africa; Vs., Asia; In. Europe; N V., North America: S.A.. south America. (Geologic age and location given only for fossils.) I' \l ( isl RIAL ENAMELS lachyromyidae: Ischyrotomus petersoni W. L.Eoc. N.A. Knightomya de- pressusW., E.Eoc. N.A. Manitsha sp. W.. E.Olig. N.A. Microparamys lysitensis W., E.Eoc. N.A. Paramys c. copei K.. \\ I .Eoc. N.A. Paramys 1968 RODENT INCISOR ENAMEL 15 c. major K., W., E.Eoc. N.A. Paramys delicatior K., M.Eoc. N.A. Paramys excavatus K., W., E.Eoc. N.A. Thisbemys perditus W., E.Eoc. N.A. Sciuravidae: Mysops parvus W., M.Eoc. N.A. Sciuravus nitidus K., M.Eoc. N.A. Theridomyidae : Archaeomys gracilis K., L.Eoc.-M.Olig. En. Archaeomys major K., L.Eoc.-M.Olig. Eu. Theridomys gregarius K., L.Eoc.-M.Olig. Eu. Theridomys vaillanti K., Eoc. Eu. Pseudoseiuridae: Pseudosciurus suevicus K., E.Olig. Eu. Sciuroides quercyi K., L.Eoc.-M.Olig. Eu. Sciuroides sp. K., L.Eoc.-M.Olig. Eu. UNISERIAL ENAMELS Ischyromyidae: Ischyromys typus K., M.Olig. N.A. Prosciurus relictus W., M.Olig. N.A. Titanotheriomys veterior K., E.Olig. N.A. Cylindrodontidae: Cylindrodon j otitis W., E.Olig. N.A. Aploduntidae : Allomxs nitens K., L.Olig. N.A. Aplodontia rufa K. Myiagaulidae : Mesogaitlus novellus W., M.Mioc. N.A. Mylagaulus sp. W., L.Mioc. N.A. Muridae: Acomys sp. K. Arvicanthis sp. K. Conilurus [Hapalotis] albipes T. Hydromys chrysogaster T. M//.v. musculus K. Notomys [Hapalotis] longicaudatus T. Otomys sp. K. Rattus norvegicus [Mus decumanus T.. KJ T., K., W. /?a«HS /"M/w) ra««J K. Cricetidae: Cricetodon minor K., L.Mioc. Eu. Eumys elegans K., W., M. -L.Olig. N.A. Eumys gracilis K., Olig. N.A. Sigmodon sp. W., L.Pleist. N.A. Akodon arenicola K. Arvicola amphibius T. Arvicola [Microtus] terrestris K. Clethrionomys [Arvicola T., Evotomys K.] glareolus T., K. Cricetulus migratorius [phaeus] K. Cricetus cricetus [fumentarius T.] T., K. Ellobius talpinus K. Gerbillus sp. K. Lemmus lemmus [norvegicus] T., K. Meriones [Gerbillus] shawi T. Microtus [Arvicola] nivalis T. Myopus schisticolor K. Ondatra [Fiber] zibetluca T., K. O/'v^o'H}'^ flavescens [longicaudatus] K. Oxymycterus rufus K. Phyllotis [Hesperomys] darwinii T. Tatera sp. K. Spalacidae: Spalax micro phthalmus [typhlus] T., K. Tachyoryctes splen- dens K. Geomyidae: Entoptychus cavifrons K., E.Mioc. N.A. Entoptychus sp. K., L.Olig. N.A. Entoptychine geomyid W. M.Mioc. N.A. Heterogeomys hispidus K. Thomomys [Geomys] umbrinus T. Heteromyidae : Heteromyid W.. M.Mioc. N.A. Dipodomys sp. K. Pero- gnathus fasciatus K. Perognathus sp. K. Perognathus [Cricetodipus] sp. K. 16 BREVIORA NO. 309 Eomyidae: Adjidaumo sp. \\ '.. M.Olig. N.A. Paradjidaumo trilophus W.. M.Olig. N.A. Dipodidae: Alactaga sibirica [saliens] k. Jaculus jaculus [Dipits hirtipes] k. Jaculus [Jerboa T., />//>/n K.J orien talis [aegyptius] T. K. Zapodidae: Napaeozapus insignis k. Sicista [Sminthus] subtilis K Zapus hudsonius k. Cliridae: C/h [Myoxus] wetzleri k. L.Olig Eu. Dryomys nitedula k Eliomys quercinus k. G/u [Myoxus k./ g/1'5 k . W. Graphiurus ocularis [capensis] k Muscardinus [Myoxus 1/ avellanarius I.. k Seiaridae: Sciurus feignouxi k.. E.Mioc, Eu. Callosciurus [Sciurus] pre- vosti k. Citellus [Spermophilus] eversmanni k. Citellus parryii [Spermo- philus empetra] k. Citellus [Spermophilus] sp. I. ( ynomys ludovicianus k. Harmota caligata [Arctomys pruinosus] I Marmoia [Arctomys] monax [empetra I / I., k. Marmota I Arctomys k./ sp. k.. W. Petaurista volans [Pteromys russicus] k. Ratufa [Sciurus] macroura k. Sciurus niger V. Sciurus vulgaris k. lamias sibiricus [Eutamias asiaticus] k. Tamias striatus [lysteri 1/ I . k Castoridae: Steneofiber eseri K., 1 Mi. \ I . \m» B. Patti rson 1959 I he rodents of the Deseadan Oligocene of Patagonia and the beginnings of South American rodent evolution. Bull. Mus. ( omp Zool . I2<> 281-428. I Received 1 5 August 1968 I BREVIORA MuseiMin of Comparative Zoology Cambridge. Mass. 31 March, 1969 Number 310 A SHOVEL-TUSKED GOMPHOTHERE FROM THE MIOCENE OF KENYA Vincent J. Maglio Abstract. A tusk fragment of the first recorded occurrence in Africa of Platybelodon, a shovel-tusked gomphothere, is described, and its relation- ships to the Asiatic and North American representatives of the Amebelo- dontinae are discussed. It is suggested that the African form represents a very early member of the group, and that Africa may have been the place of origin of the subfamily. INTRODUCTION In 1927, E. H. Barbour described as Amebelodon fricki the first specimen of a rather bizarrely specialized group of probosci- deans, the shovel-tusked gomphotheres — Amebelodontinae. The geological and geographical range of this subfamily has hitherto been believed to extend from the late middle Miocene (Tortonian)1 of Asia to the late Pliocene of North America. Osborn (1936) proposed a subfamilial separation of the North American Amebelo- don from Platybelodon, which occurs in both North America and Asia, suggesting an independent origin of the two genera — Amebelodon in North America from a stock of the "Bunomasto- dontidae" and Platybelodon from the "Serridentidae" in Asia. Gregory (1945) considered the two genera to represent a more closely related group than did Osborn and placed them both in a single subfamily, the Amebelodontinae. This is the view sub- scribed to here. Until now, it has seemed likely that the shovel- tusked gomphotheres originated in Asia from a stock of the Gomphotheriidae sometime during the early to middle Miocene, 1 Platybelodon danovi Borissiak, the earliest known species, occurs in the Chokrak beds of the Kuban region in the North Caucasus, considered to be of late middle Miocene age (Tortonian of western Europe) by Nalivkin (1960). 2 BREVIORA No. 3 10 migrating to North America during the early Pliocene. The speci- men described in the present note, however, demonstrates the presence in east Africa during middle Miocene time of shovel- tusked gomphotheres very similar to the Asiatic amebelodontines. The fragment (MCZ field number 3S-64K) was collected by Mr. James (i. Mead during the 1964 field season of the Museum of Comparative Zoology expeditions to east .Africa- directed by Professor Bryan Patterson. I he locality is 35 50' E, 02 20' N. approximated 4 mi. NM of the site of the abandoned town of Loperot and 50 mi. SSI- of I.odwar. in the Loperot area of southern Turkana District, northwestern Kenya (map in Joubert, 1966). Although none of the associated fauna has yet been studied in detail, the following elements have been identified: Deinotherium hobleyi, Dorcatherium sp. cf. D. pigotti, Chilotherium sp., ?Brachyodm sp.. and hyracoid aff. Prohyrax. The specimen, weathered from the Turkana Grit, was found on the surface near the top of the formation, about 20 feet below a fine-grained olivine basalt sequence. Joubert recognized three successive basaltic Hows overlying the Turkana Grit in this region and on geological evidence placed the onset of the basal flow in the lower Miocene (1966: 35). I he associated fauna collected from the Turkana Grit is not wholl) incompatible with this view, in that it appears to be close to those in the Rusinga scries and. although less so. to that of l.otludok. both usually considered to be of earl) Miocene age. Although detailed faunal comparison is not possible at present. the Loperot fauna probabl) correlates with part o\' the Rusinga complex'; it docs not appear to be equivalent to that of Lothidok. Dorcatherium pigotti and Chilotherium s/>. occur on Rusinga Island. -The field work was supported in National Science Foundation Grant GP-l I ss :! The name Losodok as used b\ Arambourg (1933), Whitworth (1954). and Hooijer (1966), is replaced b) the name Lothidok on most current geo- logic maps of the area. as. for example. SK. 57 Kenya. I:IM (Special). Survej of Kens a. 1965. 4 As Simpson has recently emphasized (I9(S7: 43). the stratigraphic and faunal successions at Rusinga are at present inadequate!) knovvn. as are the facies and age relationships of the several other earlier Miocene faunas known from the Kavirondo Gulf region. Evidence in support of the often quoted "earl) Miocene" age. or indeed of any other age. for these deposits is inconclusive, although the) are almost certainly older than those at Fort Ternan. for uhich an acceptable age (12-14 million years) is available. The 1969 SHOVEL-TUSKED GOMPHOTHERE FROM KENYA 3 The hyracoid is close to, but lower crowned than, Prohyrax from Langental, South Africa (Stromer, 1926), and may be the same as Meroehyrax from Rusinga (Whitworth, 1954). The Lothidok fauna includes Dorcatheriwn chappuisi, Megalohyrax championi (see Whitworth, 1954), and Aceratheriwn acutirostratum. The lowest basalt in the Loperot sequence has been dated at 17.5 ± 0.9 million years by Geochron Laboratories, Inc., using the Potassium/ Argon method (Patterson, personal communica- tion). This date indicates a Vindobonian age for the basalt im- mediately overlying the fossil-bearing sediments. There does not appear to be any significant time interval between deposition of the upper part of the Turkana Grit and the basalt flow, and this age is here considered approximately correct. The specimen under discussion is too fragmentary for positive identification, but, as far as can be determined from what remains, the form it represents was closely related to Platybelodon. It is therefore tentatively placed in that genus, pending further knowl- edge. PLATYBELODON SP. The specimen (Fig. 1 ) is a fragment from the middle of a left lower incisor, and measures 56.8 mm in maximum length. The upper surface is broadly concave and the lower convex. The two surfaces are not parallel but diverge somewhat towards the sides, so that the tooth is thicker along the medial and lateral edges than at the center. The medial edge is flattened, with a broad, shallow, concave groove along the entire length of the fragment; the two lower incisors may have been closely appressed during life, at least in this region of the tusk. The specimen tapers in all dimen- sions toward one end, which is probably anterior. A similar taper- ing toward the tip of the tusk is seen in more complete specimens of other amebelodontines, e.g., the specimen of Amebelodon fricki age of 15.3 million years suggested by Evernden ct al. (1964) as the maxi- mum for the Rusinga series seems unlikely. The sample used for this de- termination was taken from a locality at the base of Kiahera Hill and, as suggested by Savage (1965) and Leakey (1967), these deposits may not be part of the Kiahera Series, supposedly the oldest formation on Rusinga Island. Until the deposits of Rusinga are better understood, correlation within the east African Miocene will remain uncertain. The relative position of the various deposits within the Miocene depends in part on the inclusion in or exclusion from this epoch of the Aquitanian and Pontian stages. BRt A IOR\ NO. 3 1 0 described b\ Gregor) ( L945). The medial thickness tapers from 14.6 mm to 11.6 mm anteriorly. The lateral edge is thinner than the medial and measures S.5 mm to 7.0 mm. In overall width, the fragment decreases from 78.5 mm to 74.5 mm. re 1. Platybelodon sp. Tusk fragment, NICY field number 3S-f>4K. from the Loperoi area, Kenya, a, Dors.il view; horizontal line indicates plane of section, ft, Nnterior view of cut surface. Shaded area, investment of laminated dentine. • 1. The broken ends reveal a complex internal structure of dentine organized into dentinal system^ or rodlike masses as in most other amebelodonts. although they have not weathered out as discrete and isolated rods, such as those described in Platybelodon grangeri (Osborn and Granger. 1931) and Amebelodon jricki (Barbour. i941 ). In thin section (Fig. 2). the dentine has a fibrous texture, consisting of numerous microscopic dentinal tubules. These tubules 5 The term dentinal system was restricted by Tomes ( 1850) to that portion of a tooth consisting of a canal from which dentinal tubules radiate and around which there is no investment of enamel. 1969 SHOVEL-TUSKED GOMPHOTHERE FROM KENYA CJ ■, g CB CJ '72 U co -5 i cj CO CO O - o ~ 1-^ CJ « C f c r3 CJ -o "S cj T3 6 CJ ao C c s s CO 0 -J X! CO " C u E OX) C ca CJ u 4— CO >, M CO co *- 03 C <4-H ■rj o c CJ c o Q u "3 CO T3 C x X H CtfJ 1 — ' D.T3 CO CJ <2 E CO ; C ^ 03 -c L* ^— - , *-» oc ^ e ft. Lo — ' CN X CJ 3 O/J O O d BRE\ [ORA No. 3 10 appear to radiate from the center ol each dentinal system and ma\ curve partially around its outer edge. Near the center o! each dentinal system is a round to irreguiari) shaped canal, secondaril) filled with calcite, as are the numerous small cracks which cut across the tubule boundaries. I he canals probabl) correspond to the "small lumen" noted in the dentinal s\stems ("rod-cones") of Platybelodon by Osborn ( 1936: 465). The tooth is thus com- posed of a series of dentinal systems, each with a medullar) canal as m Orycteropus (Tomes, 1850). Some of the dentinal systems are closel) aggregated into bundles of two or three. A laminar substance is seen to swirl around and between the systems and bundles, completely tilling the spaces between them. Optically, this material resembles dentine; its structure s that of dentine and in no wa\ resembles that of cement. Such a condition has been described b\ I nines ( 1850) as confluent dentinal systems, m which "parallel dentinal systems are united to each other, throughout their length. b\ a thin longitudinal lamina of dentine.*' I he individual dentinal sWcnis are large, about one and one-hall mm in diameter, and loosel) packed: there are approximately 70 visible on the complete cross-section of the tusk in the region represented b\ the fragment. I his is m strong contrast to the \er\ small, discrete dentinal systems found in Platybelodon gran- gers, in which there are about 250 in the cross-section (Osborn and Granger, 1931 ). In a tusk referred to Amebelodon fricki (Colo- rado Museum no. 1319), Osborn and Granger noted the presence of dentinal systems ("rod-cones") "larger and much less numerous" than in /'. granger i, the total number being estimated at 100 to 150. Dentinal s\stems have not been dcfimtcK recognized in Platybelo- don danovi, although Osborn (1936: 4(>2 ) interpreted the "ir- regular longitudinal grooves," described b\ Borissiak ( lc->28) on the upper surface of the tusk, as indicating their presence. If, however, P. danovi was ancestral to /'. grangeri, or at least repre- sented a more primitive type, we might expect the internal structure of the tusk to be less advanced and closer to that o\ the African form. The outer surface of the tooth was sheathed in life b\ a thick (1.5 mm) layer o\ what appears to be strongl) laminated dentine with dentinal tubules oriented perpendicular to the surface. This superficially resembles enamel, but its internal edge is seen to merge into the underlying dentine, with which the tubules are continuous, without the sharp contact and contrasting structure characteristic of dentine-enamel junctions. Its microscopic structure is also not 1969 SHOVEL-TUSKED GOMPHOTHERE FROM KENYA 7 at all like that of cement. This sheath has been broken away from much of the tooth surface, but enough remains to indicate its former presence over the entire fragment; there is no indication of enamel or cement. In the degree of flattening, the present specimen exceeds Platy- belodon danovi and approaches P. grangeri, which shows the most extreme flattening of any species of the subfamily yet described. In size, the African form, as determined from available measure- ments, is the smallest amebelodont, with a tusk width about 28 per cent smaller than in P. danovi. DISCUSSION The present specimen demonstrates that by Vindobonian times a shovel-tusked gomphothere was present in east Africa. The internal modifications of the dentine into discrete dentinal systems, presumably for strengthening the tusk, were in a relatively rudi- mentary stage as compared with those of P. grangeri, being large, loosely compacted, and generally poorly defined. Yet if the degree of flattening can be used to infer the degree of adaptation to a rela- tively specialized ecological role, we must conclude that the present form had already reached a degree of specialization with respect to tusk shape equal to that of P. grangeri, despite the relatively primi- tive organization of the dentinal systems. It appears reasonable to suggest that the present form represents (with P. danovi) an early specialization toward the adaptations that were to characterize the later amebelodonts. Its exact relation- ship to the Asiatic representatives of this genus is uncertain, but in view of its similarity to those species, as far as can be determined from this specimen, a close relationship to P. danovi may be sug- gested, both forms being among the earliest members of the sub- family. The question arises as to the place of origin of the Amebelo- dontinae. The mere presence of this group in east Africa during the Miocene does not, of course, positively answer this question. However, unless the African form proves, on further evidence, to be an independent adaptation to this particular ecological situation, the existence of this group in Africa and Asia at roughly the same time suggests a possible African origin for the subfamily. This view becomes more probable when one considers the long prior evolu- tionary history of the Gomphotheriidae in Africa and their relatively late appearance in Eurasia during the early Miocene. It is to be hoped that further specimens will be unearthed and also that among 8 BREVIORA No. 310 the isolated gomphothere teeth and bones from the African Miocene in museum collections additional remains of this group will be recognized. ACKNOWLEDGMENTS I would like to express thanks to Professor Bryan Patterson for permission to Stud) and section the specimen, and for his comments on the manuscript. TABLE 1 Comparative measurements of Platybelodon species from Asia with Platybelodon from Africa, in mm WIMI 26200 P. danovi1 P grangeri1 M< / M< Z Pype Type J8-64K2 ^ '>4k:; width HO 166 79.5 74.5-78.5 medial thickness ;" ;; '6.1 11.6-14.6 lateral thickness 15 25 10.8 07.0-08.5 medial thickness width (i :?2 0.199 0.224 average thickness Q2Q] () ,75 Q ,6g width 1 Measurements foi P. danovi and P. grangeri taken from Osborn and Granger 1 1931, p. 2). I stimated mean measurements corrected for missing outer layer ol laminated dentine. Vctual measurements of specimen. LITERATURE CITED Aramboi rg, ( 1933. Mammiferes Miocenes du rurkana (Afrique Orientale). \nn. Paleoni.. Paris. 18: 123-146. Barboi r, E. H. 1927. Preliminary notice of a new proboscidean, Amebelodon fricki, gen. et sp. nov. Bull. Nebraska State Mus. I: 131-134. 1941. A shovel-tusked mastodon, Amebelodon fricki, from Kansas. Bull. Univ. Nebraska State Mus.. 2: 37-46. 1969 SHOVEL-TUSKED GOMPHOTHERE FROM KENYA 9 Borissiak, A. 1928. On a new mastodon from the Chokrak beds (middle Miocene) of the Kuban region, Platybelodon danovi, n. gen. n. sp., Ann. Soc. Paleont. Russ., 7: 105-120. Evernden, J. F., D. E. Savage, G. H. Curtis and G. T. James 1964. Potassium-Argon dates and the Cenozoic mammalian chronol- ogy of North America. Amer. Jour. Sci., 262: 145-198. Gregory, J. T. 1945. An Amebelodon jaw from the Texas Panhandle. Univ. Texas Pub., No. 4401: 477-484. Hooijer, D. A. 1966. Miocene rhinoceroses of East Africa. Fossil mammals of Africa, No. 21. Bull. Brit. Mus. (Nat. Hist.), Geol., 13 (2): 119-190. JOUBERT, P. 1966. Geology of the Loperot area. Geol. Surv. Kenya, Rep. No. 74: 1-52. Leakey, L. S. B. 1967. Notes on the mammalian faunas from the Miocene and Pleistocene of East Africa. In: W. W. Bishop and J. D. Clark, eds.. Background to Evolution in Africa. Chicago and London, Univ. Chicago Press, pp. 7-29. Nalivkin, D. V. 1960. The Geology of the U.S.S.R. — A Short Outline. London, Pergamon Press, 70 pp. [Translation.] Osborn, H. F. 1936. Proboscidea, a Monograph of the Discovery, Evolution, Mi- gration and Extinction of the Mastodonts and Elephants of the World. Vol. 1. New York, Amer. Mus. Nat. Hist., 802 pp. Osborn, H. F., and W. Granger 1931. The shovel-tuskers, Amebelodontinae, of Central Asia. Amer. Mus. Novit., No. 470: 1-12. Savage, R. J. G. 1965. The Miocene Carnivora of East Africa. Fossil mammals of Africa, No. 19. Bull. Brit. Mus. (Nat. Hist.), Geol., 10 (8): 241-316. Simpson, G. G. 1967. The Tertiary lorisiform primates of Africa. Bull. Mus. Comp. Zool., 136 (3):39-61. Stromer, E. 1926. Reste land- und susswasser-bewohnender Wirbeltiere aus den Diamantfeldern Deutsch-Slidwestafrikas. In: Kaiser, E., Die Diamantenwiiste Siidwestafrikas, 2: 107-153, Berlin. 10 BRI MORA No. 3 10 I OM1 s. I. 1850. On the structure of the dental tissues of the order Rodentia. Phil. I rans Roy. Soc. London, Pari II: 529-567. Will I WOH I M. T. Il>54. I he hyracoids ol I ast Africa. I ossii mammals of Africa, No. 7. London, Brit. Mus. (Nat. Hist.), 58 pp. i Received 25 April 1968.) BREVIORA Museum! of Contiparatlve Zoology Cambridge, Mass. 31 March, 1969 Number 311 THE GENERA APENESIA AND DISSOMPHALUS IN ARGENTINA AND CHILE (HYMENOPTERA, BETHYLIDAE) Howard E. Evans Abstract. Agenesia and Dissomphalus, two of the three genera of Pristocerinae occurring in Argentina and Chile, have been known from those countries from only a few specimens representing 5 species. Recent use of Malaise traps has greatly increased the available material and has made a further study of these genera desirable. In Apenesia, 12 species are here reported from Argentina, 6 of them new; 2 species are reported from Chile, 1 of them new. In Dissomphalus, 9 species are reported from Argen- tina, 8 of them new; this genus has not been collected in Chile. Keys to species are presented and important taxonomic features are illustrated. This is a supplement to two papers published in the Bulletin of the Museum of Comparative Zoology.1 It covers recently col- lected material of two of the three genera of the subfamily Pristo- cerinae occurring in southern South America. The remaining genus, Pseudisobrachium, has been treated elsewhere.2 Most of the material reported on here was taken by Dr. Lionel Stange in Malaise traps in the province of Tucuman, Argentina. The value of these traps as faunal samplers is shown by the fact that Apenesia was previously known from Argentina from two species represented by 7 specimens; in the present paper 12 species are reported, 6 of them new, these species known from over 150 speci- mens. Dissomphalus was previously known from Argentina from 3 species; 9 additional species are here reported from that country, 8 of them new. The figures for Pseudisobrachium are even more impressive: previously known from Argentina from only 4 species, each represented by only one or a few specimens, this genus is now i Bull. Mus. Comp. Zool., 130: 249-359 (1963). Bull. Mus. Comp. Zool., 132: 1-222 (1964). 2 Acta Zoologica Lilloana (in press). 2 BRI- VIORA No. 3 1 1 known from 22 species, represented in all by several hundred specimens. Altogether, the number of known species of Pristo- cerinae in Argentina has been increased from 9 to 48, the number in Chile from 0 to 3. Material quite recently collected by Dr. Stange near Tucuman and by Charles C. Porter in several localities has consisted wholly of these same species, suggesting that a fairly high percentage of the species occurring in the areas sampled is now known. A discussion o\' the terminology employed will be found in the papers cited earlier. The abbreviations used are listed here for the sake oi ready reference: DAO: diameter of anterior ocellus (transversely ) HE: height of eve ( maximum, lateral view) LFW : length o( fore W ing 111: length o\' head (frontal view, including clvpeus but not mouthparts ) OOL: ocello-ocular line (minimum distance between lateral ocelli and nearest eye margin) WF: width of front (at minimum point, between eyes) Wll: width of head (full frontal view, including eyes) WOT: width of oeellar triangle ( including lateral ocelli). All the material discussed in this paper is in the collections of the [nstituto Miguel Lillo, Tucuman. Argentina, and oi the Museum of Comparative Zoology, Cambridge, U.S.A. All holotypes have been deposited in the Institute Miguel Lillo. GENUS A PEN ESI A WESTWOOD This genus has been known from 35 species in South America, to which 7 more species are added here. All South American species are presentlv known from only one sex. and this unfortunate situation will prevail until these insects arc reared in scries from their hosts or taken in copula. No females of the genus Agenesia are as yet recorded from Argentina or Chile. In the treatment that follows, arrangement of the species-groups follows my 1963 revision. Key to male Apenesiu known from Argentina and Chile 1. Pronotal disc without a transverse carina anteriorly 2 Pronotal disc with a transverse carina anteriorly 10 2. Eyes glabrous: clvpeus with a median lobe, not prominent on the sides "(Figs. 10-13) 3 1969 APENESIA AND DISSOMPHALUS 3 Eyes hairy; clypeus large, well developed on sides, and without a distinct median lobe (Figs. 7-9) {dissomphaloides group) 8 3. Abdomen with a distinct, moderately long petiole; LFW 2.1-2.4 mm (laevigata group) 9. cremdata (Kieffer) Abdomen sessile; LFW exceeding 3 mm 4 4. Propodeum with no evidence of a transverse carina margining the disc behind; pronotum very short, strongly sloping (exilis group) 8. chilena Evans Propodeum with a transverse carina margining the disc behind; pro- notum longer and with a more or less flat dorsal surface 5 5. Antennae with erect setae on the under surface, which stand out strongly above the pubescence (most noticeable on segments 4-7) (pilicornis group) 6 Antennae with erect setae sparse and standing out slightly if at all above the prominent, bristling pubescence {columbana group) 7 6. Transverse carina of propodeum complete, median carina also com- plete and reaching transverse carina; LFW 3.3-3.5 mm 1 . angusticeps Evans Transverse carina of propodeum obsolescent in middle, median carina not nearly reaching posterior margin of disc; LFW 4.0-5.6 mm .... 2. reducta Evans 7. Ocelli of moderate size; OOL at least as great as WOT; head about as wide as high 3. flammicornis Evans Ocelli greatly enlarged; OOL about half WOT; head considerably wider than high 4. photoplula (Ogloblin) 8. Mandibles broad, with four teeth (Fig. 23); clypeus with a sharp median tooth but no additional processes (Fig. 9); notauli present only on an- terior half of mesoscutum 7. pygmaea n. sp. Mandibles slender, with two or three teeth; clypeus with a small median tooth flanked by two rounded processes (Figs. 7, 8); notauli com- plete 9 9. Mandibles bidentate (Fig. 21); punctures of front small but quite dis- tinct; legs testaceous 5. lilloana n. sp. Mandibles tridentate (Fig. 22); front obscurely punctate; coxae and femora medium brown 6. simulata n. sp. 10. Abdomen sessile, first tergite broadly reaching articulation with pro- podeum; mandibles with three large teeth (Fig. 24); inner margin of volsella with only a few setae just below cuspis {mexicana group) ..11 Abdomen short-petiolate, first tergite not reaching extreme base; man- dibles with 4 or 5 teeth (Figs. 25, 26); inner margin of volsella setose for a considerable distance (brasdiensis group) 12 11. Clypeus broadly subangulate (as in Fig. 12); antennae elongate, the pubescence short and subappressed; head and thorax polished, non- alutaceous 10. laevicornis n. sp. Clypeus truncate or bidentate apically (Fig. 10); antennae shorter, with suberect pubescence; head and thorax somewhat alutaceous 11. inca Evans 4 HRI-VIORA No. 31 1 12 Clypeus short, broadly truncate (Fig. 13); mandibles with Eour teeth ( Fit;. 26); front polished, non-alutaceous, strongly punctate 14. spinipes n. sp. C lypeus more prominent, subangulate (1 igv 1 I. 12): mandibles with five teeth (Fig. 25); front somewhat allltaceoUS, punctures small and u idel) spaced 13 13. Ocelli enlarged, DAO .20-.22 - WF, OOL .80-.9S x WOT; mesoscutum evenl) alutaceous; parameres noi bighl) modified (Fig. 4> 12. spatulata n. sp. Ocelli not or but slightly enlarged, DAO .16 to .20 x WF. OOL at least slightly exceedini; WOT; mesoscutum not or obscurely alu- taceous. at least medially; parameres each divided into two separate lobes (Fig. 5) 13. hut rata n. sp. 1 . Apcnesm angusticcps Evans This species lias been known from one male from Santa Cruz, Bolivia. Five Argentinian males are similar to the type in color and sculpture. However, the antennae are somewhat shorter (seg- ment three 1.6-1.9 ■ as long as wale, segment eleven about twice as long as wide) and the front is somewhat broader (WF 1.1-1.3 X HE"). 1 I \\ varies from 3.2 to 3.8 mm. Specimens examined. — ARGENTINA: 1 '. Horco Molle, Tucuman. 3-10 April 1966 (L. Stange); 2 ■ ;. Horco Molle, Oct.. Dec. 1967 (C C. Porter ); I t, 11 km W Las Cejas, Tucu- man. 1-21 Feb. 1968 (I.. Stange); 1 . San Pedro Colalao. Tucu- man, Nov. -Dec. 1967 ( L. Stange). 2. Apenesia reducta Evans I his species has been known from a single male from Chapada, Brazil. Twenty-three males now before me Ice) well to this species if allowance is made for the I act that in about half the specimens the ver\ small fourth mandibular tooth is altogether absent. In some specimens the tip of the abdomen is weakly suffused with brown, while in others the last two segments are rufous as in the type. All these specimens are larger than the type (LFW 4.2-5.6 mm) except lor the two from Tucuman. which are slightly smaller (LFW 3.6-3.9 mm). Throughout this series, the propodeum is broader than in the type (disc 1.2-1.4 X as wide as long), al- though similar in sculpturing. Specimens examined. — ARGENTINA: 2 6 6 , 1 1 km W Las Cejas, Tucuman. Jan. -Feb. 1968 (L. Stange): 5 ' . Oran, Abra Grande. Salta. 29 Jan.-7 Feb. 1967 (R.^Golbach); 4 $ S, Rio Pescado. Salta, 19-25 Nov. 1967 (C. Porter. E. Willink). PARA- GUAY: 12 I i . Carumbe, 1 Feb.-8 March 1966 (R. Golbach). 1969 APENESIA AND DISSOMPHALUS 5 3 . A penesia flammicornis Evans This species was described from Santa Cruz, Bolivia, and later reported from Jujuy, Argentina."' Nine additional males from northern Argentina compare favorably with the type, but some are smaller (LFW 4.0-5.5 mm), and there is considerable variation in some standard measurements (OOL 1.05-1.40 X WOT; WF 1.05-1.20 X HE). Specimens examined. — ARGENTINA: 7 $ $, Oran, Abra Grande, Salta, 29 Jan.-7 Feb. 1967 (R. Golbach); 2 s $ , Rio Pescado, Salta, 19-25 Nov. 1967 (C. Porter, E. Willink). 4. Apenesia photophila (Ogloblin) This species is known from a series from Loreto, Misiones, Argentina. It is closely related to the preceding, but has much larger ocelli and a reddish abdomen. I redescribed the species in mv 1963 revision of this genus. 5. Apenesia lilloana new species Holotype. — $ , ARGENTINA: Horco Molle, San Xavier Mts., Tucuman, 1-7 May 1966 (L. Stange). Description of type. — Length 2.6 mm; LFW 2.4 mm. Head black; thorax and propodeum fusco-castaneous; abdomen castane- ous, irregularly banded with light brown, base of first segment piceous; mandibles in large part testaceous; first two antennal seg- ments testaceous, remainder of antenna dark brown; legs testace- ous, coxae and femora weakly suffused with darker brown; wings subhyaline, with dark setulae, veins and stigma brown. Head, thoracic dorsum, venter, and basal parts of legs clothed with dense, short hair. Mandibles slender, bidentate (Fig. 21). Clypeus large, well developed in front of antennal sockets, below which there are large, semicircular depressions; median carina sharply defined, nearly straight in profile, forming a small median tooth at its terminus, this tooth flanked by two small, rounded processes, so that the clypeus is weakly tridentate (Fig. 7). Antennae very slender, scape elongate, first four segments in a ratio of about 3:1:1:1, segments three and eleven each nearly 3 X as long as wide; flagellar pubes- cence erect, bristling, setulae nearly as long as width of flagellum. Eyes hairy, somewhat protuberant; WH .97 X LH; vertex broadly rounded, nearlv straight in middle; distance from eye tops to vertex crest somewhat less than eye height. WF 1.25 X HE; ocelli not notably enlarged, in a compact triangle, the front angle less than 3 Evans, H. E., Acta Hymenopterologica, 2: 103 (1966). 6 BREVIORA No. 31 1 a right angle; OOL 1.3 \ WOT. Front strongly alutaceous, weakly shining, with an abundance of shallow but well-defined punctures. Thoracic dorsum wholly alutaceous. obscurely punctate; pronotum short, without a transverse carina: notauli linear, complete, curved somewhat mesad posteriorly. Propodeum of moderate length, with a somewhat irregular transverse carina along the top of the de- clivity, the median carina not quite reaching this transverse carina: surface strongly reticulate basally, polished apically; disc measur- ing 1.1 as long as wide. Mcsopleura alutaceous. without a well-defined callus. Pore wing with the discoidal vein strongly pigmented lor a distance greater than length of basal vein. Abdo- men verj slender at base, subpetiolate; subgenital plate broadly truncate; genitalia with the aedoeagUS terminating in twit simple lobes, the parameres with a lateral, digitiform process (Fig. 1). Paratypes. — 4 ■ ■ . same data as type except various dates March-July 1966. 1967 (L. Stange) Variation. — The paratypes varv slightly in si/e (II \V 2.3-3.2 mm): \\ 1 1 111 \anes from .92 to >>X. \\ I III' from 1.15 to 1.25. OOL/WOT from 1.15 to 1.25. There is no noteworthy variation in color or sculpture. Remarks. I would assign this species to the dtssomphaloides species-group, previously known from Mexico and the southwestern United States, although it will not run to that group in my revision of Apenesia ( 1963 I because of the presence of a transverse carina on the propodeum. However, the hair\ e\es. tridentate clvpeus. and male genitalia all suggest that it belongs in that group. The two species that follow are closely related to lilloana and these remarks applv equall) well to them. 6. Apenesia simuUzta ni \\ spi (ins Holotype. \ . ARGENTINA: Horco Molle. San Xavier Mts.. Tucuman. 4-12 Jan. 1966 (L. Stange). Description of type. — Length 3.0 mm: LFW 2.6 mm. Head, thorax, and base of first abdominal segment black: remainder of abdomen dark castaneous. irregularly banded with light brown: mandibles in large part testaceous; clvpeus dull ferruginous; first two antennal segments testaceous, remainder of antenna dark brown; legs medium brown except trochanters, tarsi, and femoro- tibial joints paler; wings subhyaline. with dark setulae. veins and stigma brown. Head, thoracic dorsum, venter, and basal parts of legs clothed with short, brownish hair. Mandibles slender, tridentate (Fig. 22). Clvpeus much as in lilloana, but the median tooth very small and flanked by two fairly large, rounded processes (Fig. 8). 1969 APENESIA AND DISSOMPHALUS 7 First four antennal segments in a ratio of about 16:5:6:5, seg- ments three and eleven each about 2.5 X as long as wide; flagellar pubescence erect, bristling, setulae nearly as long as width of flagellum. Eyes hairy, strongly protuberant; WH .96 X LH; sides of head roundly convergent behind eyes; occipital carina unusually strong, readily visible for a considerable length when head is viewed from front. Front broad, WF 1.4 X HE; ocelli small, in a compact triangle far above eye tops, OOL 1.55 X WOT. Front moderately alutaceous, somewhat shining, punctures small and shallow, sepa- rated by 2-4 X their own diameters. Thoracic dorsum alutaceous, obscurely punctate; notauli strong, complete. Propodeal disc slightly longer than wide, with a well-defined transverse carina behind; sculpturing similar to that of preceding species. Features of mesopleura and wings as in lilloana, and abdomen subpetiolate as in that species. Subgenital plate broadly emarginate apically; genitalia similar to those of lilloana but the parameres simple, the aedoeagus with prominent ventral rami and with elongate apical lobes which are serrate on their inner margins (Fig. 2). Paratypes. — 6 $ $ , same data as type except various dates Jan.-May 1966 (L. Stange). Variation. — LFW varies from 2.2 to 2.9 mm, WH/LH from .95 to .97, WF/HE from 1.30 to 1.45, OOL/WOT from 1.3 to 1.6. In the majority of specimens the propodeal disc is about as long as wide. 7. Apenesia pygmaea new species Holotype. — <5 , CHILE: Queb. de la Plata, Rinconada Maipu, Santiago Prov., 510 meters, 26 Dec. 1966 (L. Stange). Description of type. — Length 1.8 mm; LFW 1.6 mm. Head and thorax dark brown, abdomen somewhat lighter brown except base of first tergite black; apical half of mandibles light brown; antennae dark brown except second segment testaceous; legs brown, tarsi and apices of tibiae somewhat lighter than remainder; wings hyaline, with dark setulae, veins and stigma brown. Head and thorax dorsum with rather dense short, brownish setae. Mandibles with a large apical tooth and three small, sharp teeth above it (Fig. 23). Clypeus large, well developed in front of antennal in- sertions, with a median tooth but no additional irregularities; median carina very strong basally, then abruptly declivous to apex (Fig. 9). Antennae very slender, first four segments in a ratio of about 7:4:3:4, segments three and eleven each about 3 X as long as wide; flagellar pubescence suberect. bristling, setulae about as long as width of flagellum. Eyes small but strongly convex, hairy; 8 BREVIORA NO. 31 1 WH .97 X LH; vertex broadly rounded, distance from eye tops to vertex crest subequal to eye height. WF 1.25 X HE; ocelli small, widely spaced, front angle of ocellar triangle less than a right angle; OOL 1.3 X WOT. Front alutaceous although rather Strongly shining, obscurely punctate. Thoracic dorsum also alutace- ous, somewhat shining; pronotum unusually short, without a trans- verse carina; mesoscutum with a few small punctures medially, notauli strong on anterior half, absent behind; scutum longitudinally depressed on each side. Propodeum elongate, with a weak, in- complete carina margining the disc behind; disc 1.3 X as long as wide, with a long median carina and several short, radiating carinae basallv. Features of mesopleura and wings as in preceding species; subgenital plate truncate as in that species; genitalia not studied. Remarks. — I have seen only one specimen of this minute species, but since it is the second species of the genus known from Chile, it seems worth describing at this time. The other Chilean species, chilena, is much larger and has a very different clypeus, glabrous exes, larger ocelli, and several other prominent differences. 8. Apenesia chilena Evans I described this species from one specimen from Pichinahuel, ArauCO, Chile.' I have seen do additional specimens, nor have T seen other South American species closely related to it. I assigned the species tentatively to the exilis group, known otherwise from the southern United States. 9. Apenesia crenulata (Kieffer) This very distinctive species has been known only from the type. from Para. Brazil. Two specimens before me agree well with the type but are both slightly smaller (LFW 2.1-2.2 mm) and have the ocelli less far removed from the eyes (OOL 1.1-1.2 y WOT); both have propodeal sculpturing generally similar to that of the tvpe. but the median carina is well defined. Specimens examined. — ARGENTINA: 1 ". Oran. Abra Grande, Salta, 29 Jan.-7 Feb. 1967 (R. Golbach). BRAZIL: 1 i . Nova Teutonia, Santa Catarina, Feb. 1964 (F. Plaumann). 1 0. Apenesia laevicornis new species Holotype.— - . ARGENTINA: 11 km W Las Cejas. Tucu- man, 13 ' July-2 Aug. 1967 (L. Stange). Description of type. — Length 3.8 mm; LFW 3.2 mm. Head and thorax black; abdomen dark castaneous. irregularly mottled 4 Proc. Ent. Soc. Washington, 69: 271 (1967). 1969 APENESIA AND DISSOMPHALUS 9 with lighter brown; mandibles black except suffused with dull fer- ruginous on apical third; antennae very dark brown; legs dark brown except tarsi somewhat paler; wings hyaline, with pale setulae, veins and stigma brown. Body setae relatively sparse and long. Mandibles with three large apical teeth (Fig. 24). Clypeus broadly subangulate, median part somewhat tectiform but not carinate (as in Fig. 12). Antennae slender, first four segments in a ratio of about 14:4:9:8, segment three 3 X as long as wide, segment eleven 3.5 X as long as wide; flagellar pubescence short, sub- appressed, a few slightly longer, erect setulae standing above it. Eyes with scattered short setae, only slightly protuberant; vertex broadly rounded off well above eye tops; WH .97 X LH. Front broad, WF 1.47 X HE; ocelli small, widely spaced, in a right triangle; OOL 1.25 X WOT. Front strongly polished, non-alutace- ous, punctures strong, rather irregularly spaced, mostly separated by 5-8 X their own diameters. Thoracic dorsum also wholly non- alutaceous, strongly polished, sparsely punctate; pronotum with a transverse carina on the anterior slope; notauli and parapsidal fur- rows strong, nearly complete. Propodeal disc as long as wide, with a complete median carina and a rather irregular transverse carina; basal part of disc alutaceous and with strong reticulations, apical part smooth and shining. Mesopleura wholly polished, non- alutaceous, very sparsely punctate. Discoidal vein of fore wing very weakly developed, arising a short distance down on transverse median vein. Abdomen sessile; subgenital plate arcuately emar- ginate; aedoeagus complex although nearly parallel-sided; ventral arms of cuspides narrowly rounded; parameres with a rounded apical enlargement (Fig. 3). Remarks. — This unusual species is known from a single speci- men. It is a member of the mexicana group, similar to neotropica Evans in many details, but with longer and smoother antennae, a much broader and more polished front, and smaller ocelli. 1 1. Apenesia inca Evans This distinctive species was described from Peru and Ecuador in 1963; in 1966 I recorded it from two localities in eastern Brazil. Nineteen Argentinian males have the usual tridentate mandibles and bidentate clypeus (Fig. 10), as well as genitalia resembling those of the type closely. LFW varies from 2.7 to 3.8 mm, WF from 1.0 to 1.2 X HE; in the majority of specimens OOL is subequal to or even slightly less than WOT. Specimens examined. — ARGENTINA: 19 S $ , 1 1 km W of 10 KREVIORA No. 3 I I Las Cejas, Tucuman, Sept.. Nov., Dec. Jan.. Feb.. 1967-68 (L. Stange). 1 2. Apenesia spatulata new species Holotype.- - . ARGENTINA: Horco Molle. Tucuman. 7-26 March 1967 (L. Stange). Description of type. — Length 4.S mm; LFW 4.0 mm. Head, thorax, and abdominal petiole black; remainder of abdomen dark castaneous with irregular banding with lighter brown; apical hall' o( mandibles testaceous, teeth rufous; antennae medium brown except second segment light brown; legs entirely testaceous; wings subhvaline. with dark setulae. vein and stigma dark brown. Mandi- bles broad, with five strong teeth (big. 25). Clypeus broadly subangulate. its median line tcetiform but not carinate (Fig. 1 1 ). First four antenna] segments in a ratio of about 20:7:10:10, seg- ment three 2.5 ■ as long as wide, segment eleven 3 X as long as wide; flagellar pubescence pale, suberect. majority of setulae about .7 as long as width of llagellum. Fycs large, somewhat protuberant, glabrous; w II 1.03 1. 1 1; vertex forming a broad, even arc above eye lops. \\ I 1.1 < HE; ocelli large, less than their diameters apart, front ocellus located on an imaginary line drawn between eye tops; DAO .22 WF; OOL .85 WOT. Front wholly alutaceous, weak!) shining, punctures small but sharply defined, separated for the most part by 3-5 their own diameters. Pronotal disc margined m front by a transverse carina, weakly alutaceous and with scattered, minute punctures: mesoscutum also weakly alutaceous. shining, with punctures slightly stronger than on pronotum; QOtauli nearly complete. Propodeum elongate, the disc margined behind by a series of line transverse ridges, disc measuring slightly longer than wide: median carina complete, disc also reticulate medio-basally. Mesopleurum wholly weakly alutace- ous. obscurely punctate anteriorly. Middle tibiae strongly spinose above. Discoidal cell of fore wing weakly outlined, the discoidal vein itself Strongl) pigmented for a distance about equal to basal vein. Abdomen petiolate. but the petiole very short; subgenital plate truncate: aedoeagus complex; ventral lobes of cuspides spatu- late. narrowly rounded apicallv. paramercs with a broad, sub- quadrate lobe which is directed mesad (Fig. 4). Paratxpcs. — 20 $ $ , same data as type but various dates Jan.-July 1966 (L. Stange). Variation. — In the paratvpes. LFW varies from 3.2 to 4.1 mm. WH from 1.03 to 1.06 LH. WF from 1.05 to 1.20 X HE. OOL 1969 APENESIA AND DISSOMPHALUS 11 from .80 to .95 X WOT, DAO from .20 to .22 X WF. There is no noteworthy variation in other characters. Remarks. — This species is a member of the brasiliensis group, apparently related to angustata (Evans). The species name refers to the large, spatulate lobes of the cuspis. 13. Apenesia lacerata new species Holotype.— $ , ARGENTINA: 11 km. W of Las Cejas, Tucuman, 16-29 April 1967 (L. Stange). Description of type. — Length 3.3 mm; LFW 3.0 mm. Head, thorax, and abdominal petiole piceous; remainder of abdomen dark castaneous, irregularly banded with lighter brown; mandibles testaceous, darker at extreme base and apex; antennae testaceous basally, grading into medium brown beyond segment three; legs wholly testaceous; wings hyaline. Mandibles with five teeth (as in Fig. 25); clypeus broadly subangulate, tectiform medially (Fig. 12). First four antennal segments in a ratio of about 18:5:10:9, segment three 3 X as long as wide, segment eleven 3.5 X as long as wide; flagellar pubescence suberect, nearly as long as width of flagellum. Eyes weakly hairy, slightly protuber- ant; WH 1.03 X LH; vertex forming a broad, even arc above eye tops. WF 1.22 X HE; ocelli slightly enlarged, DAO .17 X WF; front ocellus slightly above a line connecting eye tops; OOL 1.1 X WOT. Front wholly alutaceous, somewhat shining; punctures small, separated by 3-5 X their own diameters. Pronotum strongly polished, non-alutaceous, the disc margined in front by a transverse carina; mesoscutum strongly polished, very weakly alutaceous laterad of the very strong, complete notauli. Propodeal disc as wide as long, sculptured exactly as in the preceding species. Mesopleurum shining, barely alutaceous, strongly punctate in front. Discoidal vein of fore wing very weakly outlined by pig- mented lines. Abdominal petiole very short; subgenital plate subtruncate; genitalia of unusual form in that the ventral arms of the cuspides are very long and pointed, the parameres deeply divided into a slender process and an enlarged mesal lobe (Fig. 5). Paratypes. — 17 $ $ , same data as type but various dates July-Nov. 1967 and Jan., Feb., May 1968 (L. Stange). Variation. — LFW varies from 3.0 to 3.3 mm, WH from .98 to 1.03 X LH, WF from 1.12 to 1.30 X HE, DAO from .16 to .20 X WF, OOL from 1.1 to 1.2 X WOT. In the majority of paratypes the propodeal disc is longer than wide. In a few speci- mens the front coxae and femora are suffused with brown. In two specimens the mesoscutum is somewhat alutaceous, even medially. 12 KREVIORA No. 31 1 Remarks. — This species resembles the preceding closely, and I would be inclined to consider the two conspecific if it were not for the distinctive genitalia. The name laceruta refers to the greatly dissected parameres of the genitalia of this species. 14. Apenesia spinipes new species Holotype. — 6 , ARGENTINA: 1 1 km W of Las Cejas, Tucu- iiuiii. 3-19 June 1966 (L. Stange). Description of type. — Length 4.3 mm; LFW 3.5 mm. Head and thorax black, abdomen dark castaneous with lighter markings along margin of first tcrgite as well as apically; mandibles testaceous except teeth rufous; antennae wholly testaceous, legs also of this color except coxae and femora suffused with brown; wings hyaline. veins and stigma dark brown. Mandibles with four large apical teeth (Fig. 26). Ctypeus very short, broadly truncate (Fig. 13). 1 irst four antenna] segments in a ratio of about 4:1:2:2, seg- ments three and eleven each about 3 X as long as wide; flagellar pubescence erect, pale, setulae about one third as long as width of tlagellum. Lyes ver\ weakl) hairy; vertex forming a broad, even arc above eye tops, distance from eve tops to vertex crest nearl) equal to HE. WH 1.02 X LH; WF 1.2 X HE; ocelli small, in a compact triangle. OOL 1.4 X WOT. Front polished, non- alutaceous, with strong punctures which are separated by only 1-2 X their own diameters. Thoracic dorsum polished, non- alutaceous; pronotal disc with a strong transverse carina in front, its surface covered with small punctures except along a median band. Mesoscutum with scattered, sharply defined punctures; notauli strong, complete; scutellar disc impunctate medially. Pro- podeal disc 1.2 X as wide as long, the transverse carina weak although complete, median carina also complete; disc reticulate medio-basallv. elsewhere strongly polished and smooth; posterior face shining, with a lew punctures and some medial roughening, but no median carina. Mesoplcurum in large part smooth and polished, the callus convex. Middle tibiae very strongly spinose. Discoidal cell of fore wing weakly outlined by pigmented lines. the subdiscoidal vein more heavily pigmented than the discoidal vein. Abdomen short-petiolate; subgenital plate truncate; genitalia with the aedoeagus narrowed subapically, with a pair of curved apical processes; ventral arms of cuspides large, rounded; para- meres with broad mesal lobes (Fig. 6). Paratypes. — 41 6 Z , same data as type except various dates Mav. Aug.. Oct.. Dec. 1967; Jan., Feb., 1968 (L. Stange). 1969 APENESIA AND DISSOMPHALUS 13 Variation. — Several specimens have the coxae and femora more heavily infuscated than in the type, and the apical third of the antennae may also be somewhat infuscated. LFW varies from 2.7 to 4.2 mm, WH/LH from 1.01 to 1.05, WF/HE from 1.10 to 1.25, OOL/WOT from 1.20 to 1.45. Remarks. — This species runs to brasiliensis (Kieffer) in my 1963 key, although the transverse carina of the propodeum is not as strong. The genitalia are very different from those of brasiliensis and the sculpturing of the basal triangle of the propodeum quite different. GENUS DISSOMPHALUS ASHMEAD Seventeen species of this genus have so far been described from South America, and 8 more are added here. I would estimate that there are at least 2 to 4 times that many species on that continent, even though some synonymy will be discovered when the sexes are properly associated. The following three species are known from Argentina from the female sex and are not treated here: attaphila (Bruch), azarai (Ogloblin), and platensis (Bruch). This genus has not yet been found to occur in Chile Key to male Dissomphalus known from Argentina 1. Second abdominal tergite with two large, pubescent pits close together medially and sharing a common depression (Figs. 16, 17) 2 Second abdominal tergite not as above, containing two medium-sized to minute pits, marks, or processes which are separated by much more than their own length (Figs. 18-20) 3 2. Pronotum with a strong transverse ridge; second tergite with four pits, the two middle ones very large, the tergite swollen and distorted be- hind them (Fig. 16) 1 . deformis n. sp. Pronotum without a transverse ridge; second tergite with two somewhat smaller pits medially, the tergite not swollen and distorted behind them (Fig. 17) 2. ulceratus n. sp. 3. Notauli absent on posterior two-thirds of mesoscutum; mandibles with four teeth (Fig. 31) 6. teren n. sp. Notauli complete; mandibles with 2 or 3 teeth 4 4. Second tergite with fairly large, round pits with raised rims (Fig. 18); subgenital plate with a large, V-shaped apical emargination; WF not exceeding HE 3. puteolus n. sp. Second tergite with minute pits, slits, or processes; subgenital plate with- out a V-shaped emargination: WF exceeding HE 5 5. Clypeus tapering to a long, sharp median point (Fig. 15); WH only .9 X LH; second tergite with a pair of small, spatulate processes antero- medially 8. bispinulatus n. sp. 14 BREVIORA No. 31 1 Clypeus not so strongly pointed apically and WH subequal to HE; second tergite not modified as above 6 6. Second tergite smooth, with a pair of pale, roughened spots at extreme anterior margin laterally, but DO Other modifications; clvpeus with a median process, not or very indistinctly tridentate. .9. incomptus Evans Second tergite with small pits, slits, or processes not nearly so close to anterior margin (I igs. 19, 20); clypeus Strongly tridentate (as in I ig. I4i 7 7. Modifications of second tergite in the form of oblique slits (Fig, 20); propodeal disc short. 1.7 x as wide as long J.infissus n. sp. Modifications Of second tergite in the form of minute pits or hair pencils ( 1 ig. Il>); propodeal disc 1.0-1.3 X as wide as long 8 8. LFW l.S mm; eyes and ocelli small. WF 1.2 X HE, OOL 1.05 X WOT; wings pale, discoidal \cin unpigmented, basal vein subered 4. mendicus n. sp. I I w :.(> mm; eyes and ocelli rathei large, WF 1.1 x HE. OOL only .8 > WOT; wings with dark veins and stigma, discoidal vein a short. hi own streak, basal vein strongly oblique 5. microstictus n. sp. 1 . Dissomphalus deformis new species Holotype — 6 , ARGENTINA: Oran. Abra Grande, Salta, 8-15 Feb. 1967 (R. Golbach). Description <>/ type. — -Length 2.7 mm; LFW 2.4 mm. Head and thorax black; abdomen dark brown, still used with lighter brown on parts oi basal and apical segments; mandibles largely testaceous; antennae testaceous, sufTused with brown on apical half; coxae and femora dark brown, hind tibiae medium brown, legs Otherwise testaceous; wings hyaline. Mandibles tridentate, the middle tooth the smallest (Fig. 27). Clypeus tridentate, median carina strong, straight in profile (Fig. 14). First four antenna] segments in a ratio of about 3:1:1:1, segment three 1.5 ■ as long as wide, segment eleven twiee as long as wide. Head as wide as high, the eyes large, convergent below; WF 1.1 X HE; ocelli widely spaced, front angle of triangle less than a right angle; OOL 0.9 )< WOT. Front alutaceous although somewhat shining, covered with shallow punctures. Thoracic dorsum strongly alutaceous, obscurely punctate; pronotal disc crossed anteriorly by a very strong carina; notauli strong, complete. Propodeal disc 1.2 X as wide as long, disc and declivity wholly coarsely reticulate. Discoidal vein of fore wing barely indicated. First tergite short, with a strong median groove for much of its length; second tergite with two very large, pubescent, elliptical pits close beside the mid- line, the tergite strongly gibbous behind and slightly laterad of these pits; second tergite also with a pair of much smaller pits in 1969 APENESIA AND DISSOMPHALUS 15 addition, so that there are four pits in a transverse series; tergite with a median band of small setae behind the pits (Fig. 16). Subgenital plate broadly concave apically; parameres large, tri- angular in lateral view, subacute apically. Paratypes. — 3 6 £ , same data as type. Variation. — The paratypes are all slightly larger than the type (LFW 2.5-2.7 mm) and all have the propodeal disc slightly shorter and wider (1.25-1.35 X as wide as long). There is no noteworthy variation in standard measurements or in the re- markable configuration of the second tergite. 2. Dissomphalus ulceratus new species Holotype. — s , ARGENTINA: Horco Molle, San Xavier Mts., Tucuman, 1-7 May 1966 (L. Stange). Description of type. — Length 2.0 mm; LFW 1.9 mm. Dark brownish-fuscous, sides of basal abdominal segments suffused with light brown; mandibles testaceous on apical half; mandibles dark brown except second segment somewhat lighter than remainder; coxae and femora dark brown, legs otherwise testaceous; wings hyaline. Mandibles tridentate, the uppermost tooth the smallest (Fig. 28). Clypeus with a strong median angulation and two small, rounded teeth beside it (much as in Fig. 7); median carina straight in profile. Third antennal segment 1.5 X as long as wide. Head as wide as high, eyes not convergent below, front broad, WF 1.3 X HE; ocellar triangle compact, OOL 1.2 X WOT. Front alutaceous, somewhat shining, covered with shallow punctures. Thoracic dorsum alutaceous although somewhat shining; pronotum short, without a transverse carina; notauli complete. Propodeum short, disc measuring 1.35 X as wide as long; disc mostly reticulate, but with a smooth area behind. Discoidal vein of fore wing present as a long but weakly pigmented streak. First tergite more rounded behind and with a shorter median groove than in deformis, and the second tergite not gibbous behind the pits as in that species; pits large, narrowly separated medially, sharing a common bowl-shaped depression, the tergite without other pits and with only a few scattered setae (Fig. 17). Subgenital plate broadly truncate. Paratype. — 1 $, same data as type except 3-10 April 1966 (L. Stange). Variation. — In the paratype, LFW is 2.0 mm, WF 1.4 X HE. This specimen is closely similar to the type in every respect. 16 BREVIORA No. 31 1 3. Dissomphalus puteolus new species Holotype. — 6 , ARGENTINA: 1 1 km W of Las Cejas, Tucu- nuin. 13-27 May 1967 (L. Stange). Description of type. — Length 2.5 mm; LFVV 2.1 mm. Head and thorax black; abdomen dark brown, with paler markings toward base and apex; mandibles in large part testaceous; scape medium brown, tlagellum light brown; coxae and femora dark brown, remainder of legs light brown; wings hyaline. Mandibles with three strong apical teeth, the most basal tooth somewhat rounded (Jig. 29). Clypeus with a projecting median lobe which is tridentate (much as in Fig. 14). First four antenna] segments in a ratio of about 10:4:3:3, segment three only slightly longer than wide, segment eleven 1.5 X as long as wide. \VH 1.02 X LH; eyes large, strongly convergent below, WF only .93 X HE; ocelli widel) spaced. OOL onl\ .8 WOT. Front alutaeeous and Strongl) roughened b\ punctures which are separated by less than their own diameters. Thoracic dorsum alutaeeous; pronotum short, without a transverse carina: mesoscutum somewhat shining, with numerous small punctures, ootauli complete. Propodeal disc unusually short and broad. 1.7 • as wide as long, surface reticu- late except smooth over a small area behind. Discoidal vein of fore wing absent. First tergite relatively longer and second tergite shorter and less convex than in preceding two species; second tergite with a pair of fairly large, circular pits dorso-laterally, each pit in a depression, with a raised rim and giving rise to a small tuft of setulae; second tergite with numerous setae, especially laterad of the pits (Fig. 18). Subgenital plate with a strong. V-shaped apical emargination; parameres broad, tapering to an acute apex. Paratypes. — 5 6 & . same data as type except various dates Jan.. March, April. June, 1966-1968 (L. Stange). Variation, — The paratypes are very similar to the type; LFW varies from 1.9 to 2.1 mm; all have the basal flagellar segments testaceous, contrasting with the darker apical segments; in two the propodeum is not quite as short as in the type, the disc measuring 1 .6 X as wide as long. 4. Dissomphalus mendicus new species Holotype. — 6 , ARGENTINA: 11 km W of Las Cejas. Tucu- man, 17 June- 12 July 1967 (L. Stange). Description of txpe. — Length 2.1 mm; LFW 1.8 mm. Head and thorax dark brownish-fuscous except pronotal collar medium 1969 APENESIA AND DISSOMPHALUS 17 brown; abdomen dark castaneous except first tergite bordered with light brown, second tergite light brown on extreme sides; mandi- bles in large part testaceous; antennae dull brown except second segment and adjacent parts of first and third segments testaceous; legs brown, tarsi and femoro-tibial joints straw-colored; wings hyaline, with pale setulae and pale veins and stigma. Mandibles tridentate, the basal two teeth small (much as in Fig. 28). Clypeus as described for the preceding species. First four antennal seg- ments in a ratio of about 5:2:2:2, segments three and eleven both about 1.5 X as long as wide. WH and LH subequal; eyes weakly converging below, WF 1.2 X HE; ocelli small, front angle of ocellar triangle less than a right angle, OOL 1.05 X WOT. Front moderately alutaceous, shining, the punctures small and shallow, separated by 2-4 X their own diameters. Thoracic dorsum alutaceous although somewhat shining; pronotum short, without a transverse carina; mesoscutum weakly punctate, notauli complete. Propodeal disc 1.25 X as wide as long, surface reticu- late. Discoidal vein of fore wing distinct for a short distance although barely pigmented. First tergite rounded behind, with a strong median groove; second tergite with a pair of minute lateral pits with raised rims, the pits occupying very shallow depressions and flanked by several setae (much as in Fig. 19). Subgenital plate broadly truncate apically. Remarks. — This small but distinctive species is known from a single specimen. 5. Dissomphalus microstictus new species Holotype. — $ , ARGENTINA: 11 km W of Las Cejas, Tucu- man, 3-19 June 1966 (L. Stange). Description of type. — Length 3.0 mm; LFW 2.6 mm. Head black; thorax dark brownish-fuscous; abdomen dark castaneous, with paler markings basally and apically; mandibles in large part testaceous; antennae medium brown except segments two and three and apex of one testaceous; legs straw-colored except front and hind femora weakly suffused with brown; wings hyaline, with dark veins and stigma. Mandibles with two strong apical teeth (Fig. 30). Clypeus with a tridentate median lobe (much as in Fig. 14). Third and eleventh antennal segments both about twice as long as wide. WH and LH subequal; eyes prominent, short-haired, inner orbits subparallel on lower half; WF 1.1 X HE; ocelli slightly en- larged, in about a right triangle, OOL only .8 X WOT. Front wholly strongly alutaceous, weakly shining, punctures very shallow, 18 BREVIORA No. 31 1 separated by 1-3 X their own diameters. Thoracic dorsum alu- taceous although less strongly so than front; pronotum short, with- out a transverse carina; mcsoscutum with scattered small punctures, notauli complete. Propodeal disc elongate, barely wider than long, disc smooth and shining behind, the median carina not reaching the transverse carina. Discoida] vein of fore wing lightly pigmented for a considerable distance. First tergite elongate, median groove con- fined to basal half; second tergite with a pair of widely spaced very small pits with raised rims, these pits in very shallow, broad depres- sions and flanked by a few setae (Fig. 19). Subgenital plate broadly, shallowly emarginate; parameres twisted mesad apically. Paratypes. — ARGENTINA: 53 <5 c5 , same data as type except various dates Nov. -Dec. 1967. Jan. -May lc)68; 1 • . San Pedro Colalao, Tucuman. Jan. 1968 (L. Stange). Variation. — LFW varies from 2.1 to 3.6 mm. The legs vary from wholly straw-colored to almost wholly brown. WF varies from 1 . 1 to 1 .3 X HE. The ocelli of some of the smaller specimens are only slightlv enlarged, and in these specimens OOL is only slightly less than WOT. There is little variation in the form of the second tergite in this long series. (•>. Dissomphalus teren new species Holotxpc. — | . ARGENTINA: Horco Molle. San Xavier Mis., Tucuman. 3-1 1 June lcK>h (L. Stange). Description of type. -- Length 1.8 mm; LFW 1.6 mm. Dark brownish-fuscous, including mandibles and antennae; legs dark brown except tarsi testaceous; wings h valine, \eins and stigma dark. Mandibles with four teeth, basal two teeth connate (Fig. 31 ). Cly- peus tridentate. with a strong median carina (much as in Fig. 14). First four antenna! segments in a ratio of about 10:4:3:3, segments three and eleven each about 1 .5 X as long as wide. Mead elongate, vertex broadly rounded off far above eye tops; WH .93 X LH; WF 1.25 HE; front angle of ocellar triangle less than a right angle, OOL 1.3 X WOT. Front weakly alutaceous. moderately shining, punctures small and shallow. Thoracic dorsum smooth, somewhat shining, obscurely punctate; pronotum without a transverse carina; notauli present on anterior third of mcsoscutum, absent behind. Propodeal disc 1.3 X as wide as long, with a transverse polished band posteriorly. Discoidal vein of fore wing weakly pigmented. First tergite rounded apically. with a median groove on the basal .6; second tergite smooth except for scattered small setae and a pair of curved hair-pencils which arise from slightly elevated bases, these located dorsally but separated by about twice the length of one of 1969 APENESIA AND DISSOMPHALUS 19 the hair-pencils. Subgenital plate unusually broad, arcuately emar- ginate; parameres very slender apically, their tips directed mesad. Remarks. — This species possesses a number of unique features: the 4-tooth mandibles, reduced notauli, and broadly emarginate subgenital plate. It is known only from the type specimen. 7. Dissomphalus infissus new species Holotype. — <$ , ARGENTINA: Oran, Abra Grande, Salta, 16- 23 Feb. 1967 (R. Golbach). Description of type. — Length 3.2 mm; LFW 2.6 mm. Dark brownish-fuscous; mandibles in large part testaceous; antennae medium brown, dark brown beyond basal .4; front coxae, all femora, and hind tibiae dark brown, remaining coxae, trochanters, femora-tibial joints, middle and front tibiae, and all tarsi testaceous; wings subhyaline, with dark setulae and dark veins and stigma. Mandibles tridentate (much as in Fig. 28). Clypeus with a triden- tate median lobe (much as in Fig. 14). First four antennal seg- ments in a ratio of about 3:1:1:1, segments three and eleven each about 1.5 X as long as wide. Head about as wide as high; front broad, WF 1.2 X HE; vertex slightly emarginate behind ocellar tri- angle, the latter about a right triangle, OOL 1.15 X WOT. Front strongly alutaceous, covered with shallow punctures which are separated by only 1-2 X their own diameters. Pronotum short, roughened but not carinate along anterior margin of disc; meso- scutum somewhat shining, with scattered small punctures; notauli strong, complete. Propodeal disc short, 1.7 X as wide as long, covered with coarse reticulations. Discoidal vein of fore wing inter- stitial with media, pigmented for a distance exceeding basal vein. First tergite rounded apically, with a strong median groove on basal half; second tergite with a pair of dorso-lateral slits with raised rims, each slit flanked by a group of setae (Fig. 20). Subgenital plate broadly truncate. 8. Dissomphalus bispinulatus new species Holotype. — $ , ARGENTINA: Oran, Abra Grande, Salta, 16- 23 Feb. 1967 (R. Golbach). Description of type. — Length 1.7 mm; LFW 1.5 mm. Dark brownish-fuscous; mandibles testaceous apically; antennae medium brown except basal three segments light brown; legs straw-colored except front coxae, all femora, and hind tibiae medium brown; wings hyaline, with dark setulae and dark veins and stigma. Man- dibles with a large apical tooth and two minute teeth above it. Clypeus tapering medially to a long, acute process which is at the 20 BREVIORA No. 31 1 terminus of a strong median ridge (Fig. 15). First four antennal seg- ments in a ratio of about 9:4:3:3, segments three and eleven each about 1.5 X as long as wide. Head elongate, WH only .90 X LH; eyes glabrous, inner orbits slightly diverging above and below middle; WF 1.17 X HE; ocelli small, in an acute triangle, OOL 1.3 X WOT. Front shining, weakly alutaceous, obscurely punc- tate. Pronotum much expanded from front to rear, sides concave as seen from above, without a transverse carina; mesoscutum weakly alutaceous, obscurely punctate, notauli complete. Propodeal disc 1 .2 X as wide as long, most of its posterior half smooth and shining, median carina not nearly reaching transverse carina. Discoidal vein of fore wing strongly pigmented for a considerable distance. First tergite rounded apicallv. with a median groove on basal half; second tergite with a pair oi small, spatulate processes close behind the margin of the first tergite. these separated medially by about twice their own length; second tergite also with a pair of roughened spots anterolaterally. Subgenital plate very broadly truncate apically. 9, Dissomphalus incomptus Evans I described this species from eight males from Santa Catarina, Brazil, in im 1964 synopsis. Its presence in Paraguay and in two provinces of Argentina suggest that it is widely distributed and not uncommon in southern South America. The specimens before me key readily to this species in my 1966 paper1 and agree with the tvpe series in most details, including the genitalia. They average somewhat larger I I I \\ 1.9-2.6 mm) and show minor variation in head measurements (WF 1.1-1.3 X HE, OOL 1.15-1.30 WOT). The sinuations beside the median process of the clypeus vary in development, such that in some specimens the clypeus may be said to be indistinctly tridentate. Specimens examined. — PARAGUAY: 1 & , Carumbe, 1 Feb. -8 March 1966 ( R. Golbach). ARGENTINA: 2 6 6, Oram Abra Grande. Salta. 1 Jan.-7 Feb. 1967 (R. Golbach); \2 $ 3 , Horco Molle. San Xavicr Mts., Tucuman, Jan. -June 1966 (L. Stange). Dec. 1967 (C. C. Porter). (Received 7 October 1968.) 5 Acta Hymenopterologica. 2: 1 10. 1969 APENESIA AND DISSOMPHALUS 21 Figs. 1-6. Male genitalia of Apenesia spp., ventral aspect. Fig. 1., A. lilloana n. sp. Fig. 2., A. simulate! n. sp. Fig. 3., A. laevicornis n. sp. Fig. 4., A. spatulata n. sp. Fig. 5., A. lacerata n. sp. Fig. 6., A. spinipes n. sp. 27 BREVIORA No. 311 23 26 27 28 29 31 1969 APENESIA AND DISSOMPHALUS 23 Figs. 7-15. Clypeus of male Apenesia and Dissomphalus. Fig. 7., A. lil- loana n. sp. Fig. 8., A. simulata n. sp. Fig. 9., A. pygmaea n. sp. Fig. 10., A. inca Evans. Fig. 11., A. spatulata n. sp. Fig. 12., A. lacerata n. sp. Fig. 13., A. spinipes n. sp. Fig. 14., D. deformis n. sp. Fig. 15., D. bispinulatus n. sp. Figs. 16-20. Base of abdomen of male Dissomphalus, dorsal view. Fig. 16., D. deformis n. sp. Fig. 17., D. ulceratus n. sp. Fig. 18., D. piiteolus n. sp. Fig. 19., D. microstictus n. sp. Fig. 20., D. inftssus n. sp. Figs. 21-31. Mandibles of male Apenesia and Dissomphalus. Fig. 21., A. lilloana n. sp. Fig. 22., A. simulata n. sp. Fig. 23., A. pygmaea n. sp. Fig. 24., A. laevicornis n. sp. Fig. 25., A. spatulata n. sp. Fig. 26., A. spi- nipes n. sp. Fig. 27., D. deformis n. sp. Fig. 28., D. ulceratus n. sp. Fig. 29., D. piiteolus n. sp. Fig. 30., D. microstictus n. sp. Fig. 31., D. teren n. sp. BREVIORA Museum of Coinpairgitive Zoology Cambridge, Mass. 31 March, 1969 Number 312 ECOLOGICAL OBSERVATIONS ON ANOUS OCCULTUS WILLIAMS AND RIVERO (SAURIA, IGUANIDAE) T. Preston Webster Abstract. Recent observations indicate that the structural niche of Anolis occultus on Puerto Rico is the peripheral foliage of the montane forests from which it has been recorded; probably best considered a canopy species, it descends to ground level only at the few localities where trail- edge vegetation merges with the tree crowns. Structurally, behaviorally, and in coloration it is very well adapted for a life on twigs and leaves. As ex- pected from other studies of Anolis ecology, the spatial preferences of oc- cultus show little overlap with those of the other species recorded from the same locality. In 1963 the first specimen of the very distinctive species Anolis occultus was captured by day at Cerro La Punta in Puerto Rico's Cordillera Central. Subsequent collecting produced about 40 addi- tional specimens from all areas of higher montane forest on the island. All but two of these specimens were collected by Richard Thomas at night along forest trails where the trail-edge vegetation merged more or less intimately with the foliage of the canopy. Thomas suggested (Williams, Rivero, and Thomas, 1965) that occultus is a canopy species with a preference for exposed areas of bare twigs and vines, although after further collecting he sug- gested that "their habitat requirements may not be as narrow as my first experience with the species indicated" (Thomas, personal communication, March, 1968). As Thomas observed only a single specimen during the day, the diurnal ecology of this elusive lizard remained a mystery. Recently 15 additional specimens were obtained 13.6 kilometers south of Palmer, Puerto Rico, in the Sierra de Luquillo. The species was first sought at night; nine specimens were easily lo- cated the night of April 7th. Four slept on long, exposed twigs, two slept on twigs near leaves, and one slept across the upper sur- face of a broad, stiff leaf. Twigs selected for sleeping perches were a quarter inch in diameter or less. Thomas's observations were 2 breviora No. 312 confirmed; occultus clings tightly to its perch while asleep, may squeak loudly when handled, and can grip the twig strongly with its prehensile tail. Sleeping specimens are a light grey and stand out clearly in the beam of a headlamp. The two remaining anoles, asleep in the same small clump of vegetation, one across a leaf, the other on a bare twig, were left for observation in the morning. Both lizards were found asleep at 6:15 A.M. One became active at 6:30. moving sluggishly to a nearby perch that seemed to offer greater exposure to the sun. The second became active a half hour later, but neither moved far for at least an additional hour. While both were in view, a third specimen was discovered when it became backlighted; far more active, it seemed to drink from raindrops on two occasions and prowled with frequent long pauses over an area of largely bare and exposed twigs, eventually wander- ing upward and out oi sight. The remaining two also disappeared in the vegetation, but with the appearance of new individuals or the reappearance of those previously seen a total of six were ob- served, at least oik- staying in sisiht at all times. Movements were slow, cautious, and generally along twigs of small diameter, al- though occasionally the anoles crossed leaves or used larger branches to ascend or descend through the vegetation; after seem- ingly great deliberation, quick and agile leaps from twig to twig and twig to leal were undertaken. Movement for more than a few inches at a time was infrequent; during the intervening pauses the lizard generally lay Hush with a twig or leaf. Much more rapid and extensive changes oi perch occurred when the foliage was briefly disturbed by a gust of wind. At 10: 15 the three specimens remaining in sight were captured without difficulty. When the locality was revisited about noon, an occultus was soon located on a dead twig in a pile of cut branches. It showed the same alternation of pauses with slow movements over short distances observed earlier. After extensive wandering among the twigs it leaped to the upper surface of a fern frond, where it re- mained for perhaps a half hour; although this small brown lizard was difficult to distinguish on a substrate of bare twigs, on the frond it was always conspicuous, and the two pale orange spots at the base of the tail were displayed. Eventually, after a little maneuvering on its leaf, the lizard jumped to another tangle of dead twigs and disappeared. That evening intensive searching yielded six additional speci- mens in an hour and a half; three were on living twigs near leaves, one was on a long dead twig, one was at the tip of a very long descending branch, and a juvenile was on a dead fern. 1969 ECOLOGY OF ANOLIS OCCULTUS 3 On Puerto Rico the genus Anolis is represented by 10 species. All share adaptations for an active, diurnal, arboreal life, are pri- marily insectivorous, and in many cases are of similar size. Rand (1964) used the concept of a structural niche, which for arboreal Anolis is a combination of perch height and diameter as deter- mined by quantitative observation, to separate into three ecological groupings seven of the eight Puerto Rican Anolis recognized at the time. Temperature preference provided an additional niche dimen- sion that permitted Rand to separate the species within each group- ing. It is of interest to relate the structural niche of occultus to those of other Puerto Rican species, especially those recorded from the same locality. Anolis occultus appears to be an inhabitant of the peripheral foliage. Branches and bushes along trails have characterized all the productive collecting localities discovered to date; these some- what artificial conditions give the collector access to vegetation continuous with the more or less high canopy characteristic of Puerto Rico's montane forests. Thomas's (1965) hypothesis that occultus is normally an occupant of the canopy is probably sta- tistically correct, as suitable conditions near the ground occur only infrequently. There is no indication, however, that areas of dead branches and climbing plants are preferred; the specimens observed in this study showed a preference for fairly dense vegetation by day and no well-defined selection of dead or living twigs for sleep- ing perches. The sleeping positions selected in thin foliage or on dead twigs are probably optimal for early morning sunning; one of the two specimens observed to awaken was asleep in moderately dense foliage, but moved early to a more exposed position where it remained motionless for a long time. Rand (1964) has provided a detailed study of the structural niches for the three common Puerto Rican Anolis present at the La Mina area in the Sierra de Luquillo. Of the three species pres- ent at La Mina, A. evermanni uses almost exclusively perches of several inches or more diameter. A. gundlachi, which shows some- what greater preference for shade than evermanni, perches lower but also on tree trunks and branches of large diameter. A. krugi, while primarily found on grasses and similar plants, occasionally perches low in bushes on branches of moderate diameter. Al- though evermanni and krugi were common, gundlachi uncommon, and cristatellus rare at the locality of the recent occultus collection, the only species showing any overlap of structural niche with occul- tus in early April was evermanni. Juvenile evermanni were occa- sionally seen on twigs and small branches, although in general they 4 breviora No. 312 seem to prefer perches of large diameter. This overlap is prob- ably over-emphasized by an examination of trail-margin condi- tions; although, as Rand noted, studies of Anolis ecology are biased by the restriction of accurate observation to the lowest level oi the forest, the available evidence indicates that evermanni lives below the canopy, while occultus is largely a species of the canopy. If the observations of as many as six occultus in a small clump of vegetation can be considered indicative of the population density at this locality, occultus is much commoner than juvenile ever- manni, onl\ one of which was noticed. Thomas recorded gund- Icichi. cristatellus, krugi, and evermanni from localities where he took occultus. Anolis occultus seems well-adapted structurally, in coloration. and behaviorally for a cryptic existence in the peripheral foliage of trees and bushes. Long and slender, with a downward tapering snout, resting specimens merge easily with the outline of a twig and cast little or no shadow ; from the distance of only a few feet a human observer finds the blending of outline of lizard and twig very deceptive. Although its use has only been observed at night, a prehensile tail is presumably of some diurnal advantage to occultus, if only to hold it firmly Hush with a perch. Thomas (1965) described the coloration of live occultus in detail. Speci- mens observed In the present study by day were grey or brown, the degree of patterning \ariable; although individuals crossed leaves on several occasions, and one lingered for a considerable length oi time on a fern frond, the green phase recorded by Thomas was not observed. The orange spots at the base of the tail are conspicuous in some postures; the) may be eye spots to ward olf predation or possibly function in intraspecific communi- cation. Motion is slow and continuous over only short distances. The tendency for occultus to be most active when the vegetation is in general motion suggests that the species seeks to remain incon- spicuous at all times. The selection of exposed surfaces for sleeping may, by warming the li/ard as early as possible on cool mornings, maximize the number of daylight hours during which occultus is alert and active. ACKNOWLEDGMENTS Field work was in part supported by National Science Founda- tion Grant GB6944 to Ernest E. Williams. Thanks are due to Dr. Williams, who sponsored this collection of Puerto Rican rep- tiles and expressed particular interest in the occultus problem; to 1969 ECOLOGY OF ANOL1S OCCULTUS 5 Mr. Richard Thomas, who supplied information on collecting in Puerto Rico; and to Dr. Williams and Mr. Thomas Schoener, who read and provided valuable criticism of this paper. LITERATURE CiTED Rand, A. Stanley 1964. Ecological distribution of anoline lizards of Puerto Rico. Ecol- ogy 45: 745-752. Thomas, Richard 1965. Field observations of Anolis occultus Williams and Rivero. Part II in: E. E. Williams, J. A. Rivero, and R. Thomas, A new anole (Sauria, Iguanidae) from Puerto Rico. Breviora, No. 231: 10-18. (Received 6 November 1968.) BREVIORA Museem of Contiparative Zoology Cambridge, Mass. 31 March, 1969 Number 313 LOUIS AGASSIZ'S NUMBERS FOR STEAMER BLAKE STATIONS OF 1877-78, 1878-79 Myvanwy M. Dick Abstract. Correct Blake station numbers corresponding to Roman numerals used by Louis Agassiz for specimens collected by the Blake, 1877- 1878, 1878-1879. Eschmeyer (1965) demonstrated that frequent errors in locality data for fishes taken during Blake expeditions, 1877-1879, have been included in several major published works on fishes and in the fish catalogue, U. S. National Museum. These errors were caused by interpreting as station numbers Roman numerals in- cluded with the specimens — - numerals intended for other purposes. For example, Goode and Bean (1896: 296-297) reported Cal- lidnymus himantophorus from Blake Station XXX and used data for that station for the station list. "XXX," however, is a number applied to the specimen, apparently by L. Agassiz, the correct station number being 274. Eschmeyer's account of these errors and their perpetuation is excellent and correct, but as no catalogue relating Agassiz's Roman numeral designation to true Blake Station numbers was known to exist, the resulting specific pub- lished errors could be corrected only with difficulty and in many instances not at all. In collating early papers and catalogues relating to the Fish Department of the Museum of Comparative Zoology, a special catalogue of deep sea fish from the Blake expedition was found. Judging by the handwriting, this was prepared by Louis Agassiz. This contains the Roman numerals, which had been incorrectly interpreted as Blake Station numbers, and corresponding Blake Stations and localities. Each number was apparently intended to apply to a specific fish. Occasionally, in the Blake catalogue of 1878-1879, the same number is given in pencil as "Duplicate," with a different Blake Station. Presumably this would indicate fish of the same species but a different locality. This presents the 2 BREVIORA No. 313 possibility of questionable localities for specimens having just the Agassiz numbers, such as "Agassiz V," which would allow seven localities under "duplicate Agassiz V." In the Agassiz catalogue of the Blake expedition of 1877 1878 the Roman numeral is not always accompanied by a Blake Station number; however a locality is given, except in a very few instances where there are entries with "no locality." Several specimens from the Blake collections were sent to Steindachner in Vienna for examination. These were assigned numbers from the Agassi/ catalogue with their locality, but the Blake Station numbers were omitted. Hence material described by Dr. Steindachner would have the Agassiz number and correct locality but would not include a Blake Station number. In general, however, Agassiz's holograph catalogue can be used to relate his Roman numeral designations to Blake Station numbers and hence to the correct data as published b\ Pierce and Patterson ( 1870), Agassi/ (1881), and Smith ( I 889). Agassiz's catalogue is reproduced below. LITERATURE CITED Agassiz. A. I SKI. 1 ist o\~ dredging stations occupied (.luring the year 1 880 by the U.S. Coast Survey Steamer "Blake." Bull. Mus. Comp. Zool.. 8(4): 95-98. ESCHMI HR.W. N. 1965. Station data errors for fishes collected hv the Steamer Make. Copeia.2: 236-238. Goode. G. B.. and I H. Bl vn 1896. Oceanic ichthyology. Mem. Mus. Comp. Zool., 22: 1-553. J'ii R( i . B.. and C. P. Patterson 1879. 1 ist of dredging stations occupied by the United States Coast Survey steamers "Corwin." "Bibb," H.issler," and "Blake," from 1867 to 1879. Bull. Mus. Comp. Zool.. 6 ( I ): 1-15. Smith, S. 1889. lists of the dredging stations of the U. S. Fish Commission. Ann. Rept. Comm. Fish Fish. (1886): 873-1017. (Received 6 December 1968.) 1969 BLAKE STATION NUMBERS CAT. DEEP SEA FISH BLAKE EXP. 1878-79 Present Number I II III IV V VI VII VIII IV dup. Vdup. Vdup. Vdup. Vdup. Vdup. Vdup. Vdup. IX VIII dup. VIII dup. IX dup. IX dup. IX dup. X Xdup. XI XII XIII XIV XIV dup. XIV dup. XIV dup. XV XVI XVII XVIII XVIII dup. XIX XX XXI XXII XXIII XXIV XXV Dredge Number 276 278 230 274 147 240 264 153 222 258 260 150 185 290 148 262 116 290 281 274 275 148 290 297 210 142 248 176 227 285 236 148 281 275 274 274 297 142 167 147 147 191 293 Depth fms. 94 69 464 209 250 164 416 303 422 159 291 375 333 73 208 92 150 73 288 209 218 208 78 123 191 27 161 391 573 159 1591 208 288 218 209 209 123 27 175 250 250 108 to 250 82.. _ Localities Number of Specimens off Barbados off Barbados off St. Vincent off Barbados off St. Kitts off Grenadines off Grenada off Montserrat off St. Lucia off Grenada off Grenada off Nevis off Dominica off Barbados off St. Kitts off Grenada off Jamaica off Barbados off Barbados off Barbados off Barbados off St. Kitts off Barbados off Barbados off Martinique off Flannegans Passage off Grenada off Dominica off St. Vincent off Grenada off Bequia off St. Kitts off Barbados off Barbados off Barbados off Barbados off Barbados Flannegans Passage off Guadeloupe off St. Kitts off St. Kitts off Dominica off Barbados 4 BREVIORA NO. 3 1 3 Present Dredge Depth Localities Number of Number Number fms. Specimens XXVI 220 116 off St. Lucia 1 XXVI! 231 95 off St. Vincent 1 XXVIII 293 82 off Barbados 1 XXIX 293 82 off Barbados 1 XXX 274 209 off Barbados 1 XXX dup. 281 288 off Barbados 1 XXX dup. 274 209 off Barbados XXX dup. 258 159 off Grenada 1 XXX dup. 291 200 off Barbados 3 XXXI 273 103 off Barbados 1 XXX 11 261 340 olTGrenad.i 1 XXXIII 132 115 off Santa Cruz 1 XXXIV 253 92 off Grenada 1 XXXV 136 508 off Santa Cruz 1 XXXV dup. 221 423 off St. Lucia 1 XXXV dup. 284 347 off Barbados 1 XXXVI 280 221 off Barbados 1 XXXVII 151 365 off Nevis 1 XXXVIII 2 1 2 317 off Martinique with parasite 1 XXXIX 121 2393 off Santa Cruz 1 XL 200 472 off Martinique 1 XL] 185 333 off Dominique 1 XLII 21 1 357 off Martinique 1 XLII dup. 222 422 off St. Lucia 3 XLII dup. 212 317 off Martinique 1 XLII dup. 151 365 off Nevis 3 XL 11 288 399 off Barbados 1 \I II dup. 260 291 off Grenada 2 XLII dup. 221 423 off St. Lucia 1 XLII dup 260 291 off Grenada 4 XLII dup. 150 375 off Nevis 3 XLII dup. 261 340 off Grenada I XL1II 266 461 off Grenada 1 XLIV 248 161 off Grenada 1 XLV 104 500 Old Bahama Channel 3 XLVdup. 104 500 Old Bahama Channel 10 ± XLVI 271 458 off Bequia 1 XLVII 188 372 off Dominica 3 XLVIII 227 573 off St. Vincent 1 XLIX 221 423 off St. Lucia 1 XLIX dup. 221 423 off St. Lucia 1 XLIX dup. 190 524 off Dominica L 227 573 off St. Vincent 1 LI 148 208 off St. Kitts 1 1969 BLAKE STATION NUMBERS LII 275 218 Llldup. 275 218 LII dup. 147 250 Llldup. 281 288 LIU 295 180 LUIdup. 295 180 LUIdup. 281 288 LUIdup. 291 200 LIII dup. 274 209 LUIdup. 275 218 LIV 283 237 LV 262 92 LVI 231 95 LVII 132 115 LVIIdup. 132 115 LVIII 129 314 LIX 281 288 LX 297 123 LXI 249 262 LXII 274 209 LXIII 274 209 LXIIIdup. 275 218 LXIII dup. 274 209 LXIIIdup. 172 62 to 108 LXIV 297 123 LXV 297 123 LXV dup. 297 123 LXVI 235 1507 LXVII 222 422 LXVII 222 422 LXVIII 160 393 LXVIII dup. 137 625 LXVIII dup. 151 365 LX1X 161 583 LXX 163 769 LXXdup. 163 769 LXXI 163 769 LXXII 230 464 LXXIII 200 472 LXXIV 295 180 LXXIVdup. 291 200 LXXV 281 288 LXXVdup. 281 288 LXXV dup. 260 291 LXXVI 265 576 LXXVI dup. 265 576 LXXVI dup. 288 399 off Barbados off Barbados off St. Kitts off Barbados off Barbados off Barbados off Barbados off Barbados off Barbados off Barbados off Barbados off Grenada off St. Vincent off Santa Cruz off Santa Cruz off Santa Cruz off Barbados off Barbados off Grenada off Barbados off Barbados off Barbados off Barbados off Guadeloupe off Barbados off Barbados off Barbados off Bequia off St. Lucia off St. Lucia off Guadeloupe off Santa Cruz off Nevis off Guadeloupe off Guadeloupe off Guadeloupe off Guadeloupe off St. Vincent off Martinique off Barbados off Barbados off Barbados off Barbados off Grenada off Grenada off Grenada off Barbados 1 2 1 1 2 20 14 3 4 2 6 BRliVIORA No. 313 P rescn i Dredge Depth Localities Number of Number Number litis. Specimens LXXV11 151 365 off Nevis LXXVIIdup 151 365 off Nevis LXXVIIdup 222 422 off St. Lucia LXXVIIdup 260 291 off Grenada LXXVIIdup 150 375 off Nevis 1 \\\ 11 dup 135 450 off Santa Cruz 2 I XXVII dup 185 333 oil Dominica LXXV11I 291 200 off Barbados 1 XXVIII du] D. 2lM 200 oil' Barbados LXXIX 274 209 oil Barbados 1 \\l\ dup. 274 209 off Barbados LXXIX dup. 291 200 Off Barbados LXXIX dup. 193 169 mi Martinique LXXIX dup. L49 60 to ISO Off Sl. Kills LXXIX dup. [46 24.^ off St. Kitts LXXX 200 4'2 off Martinique 1 \X.\ dup. 2()() 4^2 oil Martinique LXXXdup. 175 MS oil Dominica LXXX dup. 227 573 off St. Vincent LXXXdup. 135 450 off Santa Cruz LXXX dup. 163 769 oil ( iuadeloupe LXXX dup. 227 573 off St. Vincent LXXX1 151 J65 off Nevis 1 VXXI dup. 151 365 oil Nevis LXXXII 205 334 off Martinique LXXX 11 dup 205 334 otl Martinique LXXXII dup 130 451 off Santa Cruz LXXXII dup 186 333 off Dominica LXXXII dup 222 422 off St. Lucia LXXXIII 154 164 off Grenada LXXXIV 182 1131 off Dominica LXXXV 151 365 olf Nevis LXXX\ 1 180 982 olT Dominica LXXXYI du p. 180 982 off Dominica LXXXVII 196 1030 off Martinique LXXXVIII 214 476 off Martinique LXXX\ III I lup. 200 472 off Martinique LXXXVIII ( tup. 163 769 off Guadeloupe LXXXIX 205 334 off Martinique XC 258 159 off Grenada XC dup. 283 237 off Barbados XCdup. 275 218 off Barbados 3 XCI 180 982 off Dominica 1 1969 BLAKE STATION NUMBERS XCII XCII dup. XCII dup. XCII dup. XCII dup. XCII dup. XCII dup. XCII dup. XCII dup. XCII dup. XCIII XCIV xcv XCVI XCVI dup. XCVI dup. XCVII XCVII dup. XCVIII XCVIII dup. XCVIII dup. XCVIII dup. XCIX XCIX dup. C CI CH cm civ CI dup. CI dup. CV CVI CVII CVIII CIX ex CXI CXII CXIII CXIV cxv CXVI CXVII CXVIII CXIX exx 265 576 off Grenada 161 583 off Guadeloupe 135 450 off Santa Cruz 185 333 off Dominica 2 200 472 off Martinique 227 573 off St. Vincent 130 451 off Santa Cruz 195 501 off Martinique 188 399 off Barbados 239 338 off Grenadines 260 291 off Grenada 100 250-400 off Moro Castle, Cuba 185 333 off Dominica 163 769 off Guadeloupe 162 734 off Guadeloupe 2 227 573 off St. Vincent 4 274 209 off Barbados 2 274 209 off Barbados 31 227 573 off St. Vincent 227 573 off St. Vincent 3 143 298 off Montserrat 257 553 off Grenada 262 90 off Grenada 3 262 90 off Grenada 30 132 115 off Santa Cruz 231 95 off St. Vincent 142 27 Flannegans Passage 142 27 Flannegans Passage 295 180 off Barbados 291 200 off Barbados 2 297 123 off Barbados 150 375 off Nevis 190 524 off Dominica 291 200 off Barbados 148 208 off St. Kitts 188 372 off Dominica 258 159 off Grenada 300 82 off Barbados 243 82 off Barbados 151 365 off Nevis 290 73 off Barbados 243 82 off Barbados 276 94 off Barbados 276 94 off Barbados 290 78 off Barbados 243 82 off Barbados 253 92 off Grenada 8 HREVIORA N« 3. 3 1 3 Present Dredge Depth Localities Number of Number Number fms. Specimens CXXI 128 180 off Santa Cruz CXXII 142 27 Flannegans Passage CXXIII 142 27 1 l.mnegans Passage CXXIV 142 27 1 lanne^ans Passage cxxv 271 458 off Bequia CXXVI 278 69 off Barbados < WVII 243 82 off Barbados CXXVIll 300 82 off Barbados CXXIX 132 115 off Santa Cruz 1 \\x 220 1 \b oil St. Lucia ( \\\l 276 94 off Barbados 1 WXII 104 500 Old Bahama Channel ( XXXIII 241 163 off Cariacou c xxxrv 293 82 off Barbados I XXXV 104 500 Old Bahama Channel CXXXN dup. 104 500 Old Bahama Channel ( WXVI 104 500 Old Bahama Channel CXXXV11 104 500 Old Bahama Channel CXXXVIdup. 104 500 Old Bahama Channel ( WW III 104 500 ( )ld Bahama Channel < XXXIX 109 1554 off Caya de Moa CXL HI 1200 la!. 19°07'N, Ion. 74 52'W ( XI 1 150 375 off Nevis CXI. II 288 399 off Barbados ( XI III 150 375 off Nevis CXI l\ 256 370 off Grenada CXLV 284 347 off Barbados CXLV1 283 237 off Barbados CXLVII 205 surface off Martinique 2 CXLVIII 201 565 off Martinique 2 CXLVIIIdup. 201 565 off Martinique 3 CXLIX 212 317 off Martinique 2 CL 204 476 off Martinique 2 ( 1 1 217 398 off St. Lucia CLII 217 398 off St. Lucia CLIII 222 422 off St. Lucia CL1V 124 580 off Santa Cruz ( 1 V 207 826 off Martinique CLVI 182 1131 off Dominique CLVII 190 524 off Dominique CLV1II 190 524 off Dominique 2 CLIX 185 333 off Dominique CLIX dup. 185 333 off Dominique CLIX dup. 188 372 off Dominique 4 1969 BLAKE STATION NUMBERS CLX 188 372 off Dominique 1 CLXI 236 1591 off Bequia 1 CLXII 180 982 off Dominica 1 CLXIII 173 734 off Guadaloupe 2 CLXIV 769 off Guadaloupe 1 Blake Exp. 1877-78 CLXV 29 955 lat.24°36'n,lon.84°05'w 1 CLXVI 29 955 lat.24°36'n,lon.84°05'w 1 CLXVII 12 36 lat.24°46'n,lon.83°16'w 1 CLXVIII 36 84 lat.23°13'n,lon.89°10'w 1 CLXIX 12 36 lat.24°46'n,lon.83°16'w 1 CLXX 29 955 lat.24°36'n,lon.84°05'w 1 CLXXI 11 37 lat.24°43'n,lon.83°25'w 5 CLXXII 29 955 lat.24°36'n,lon.84°05'w 2 CLXXIII 29 955 lat.24°36'n,lon.84°05'w 1 CLXXIV 31 1850 lat.24°33'n,lon.84°23'w 1 CLXXV 186 88 off Alligator Keys 1 CLXXVI 37 35 Alacran Shoals, Yucatan 9 CLXXVII 37 35 Alacran Shoals, Yucatan 1 CLXXIX 47 321 lat.28°42'nlon.88°40'w 4 CLXXX 6 137 lat.24°17.5'n Ion. 82°09'w 1 CLXXXI 47 321 lat.28°42'nlon.88°40'w 1 CLXXXII 135 off Key West 1 CLXXXIII 270 Santaren Channel 1 CLXXXIV 128 to 240 Two miles off Havana 1 CLXXXV 45 101 lat.25°33'nlon.84°21'w 1 CLXXXVI 32 95 lat.24°52'nlon.88°05'w 1 CLXXXVII 177 off Havana 1 CLXXXVIII 40 off Cary's Fort Reef 1 CLXXXIX 29 955 lat.44°36'nlon.84°05'w 5 CXC 46 888 lat. 25°43'nlon. 84°47'30" 1 CXCI 44 539 lat.25°33'nlon.84°35'w 1 CXCII 36 84 lat. 23°13'nlong. 84°16'w 25 CXCIII 36 84 lat. 23°13'nlong. 84°16'w 25 CXCIV 175 off Havana 1 cxcv 40 off West Reef I CXCVI 98 "off Panfii Reef" 1 CXCVII 42 "W. of Tortugas" 1 CXCVIII 117 Illegible 1 CXCIX 37 W. of Tortugas 1 cc 306 W. of Tortugas 1 CCI 68 W. of Tortugas 1 10 BRFA'IORA NO. 3 1 3 P rescn I Number ecu CCill CCIV ccv CCVI CCV11 Dredge Depth Localities V 'umber of Number jms. Specimens 37 W of Tortugas 118 off Alligator Reef 31 1920 hit. 24 33' Ion. 84 23' 43 339 lat. 24°08'lon. 82 51' 35 804 lat. 23 54-46" Ion. 88°58' 37 35 4 miles w. of whale rock w. of Alacran Reels CCV11I 29 955 lat. 24 36' Ion. 84 05' CCIV 11 37 lat. 24 43'lon. 83c25' ( ( \ 45 off French Reef CCX1 35 off Cargoforl Reef ( ( XII 320 off Cargofort Reef ( ( XIII 413 off ( ape San \ntonio ( ( \l\ 96 loc? ( ( XV 35 W. of Tortugas ( ( XVI 1 19 lat. 26 31'lon.85°03' ( ( \\ II 13 "42 3V4 N. W. off Havana 1 ( \\ III 47 321 lat 28 42' Ion. 88 40' ( ( XIX 30 968 lat 24 33' Ion. 84 05' < [ XX 2" 955 lat. 24°36' Ion. 84 05' * 1 XXI 29 955 lat. 24 36' Ion. 84 05' ( ( XXII 47 J21 lat. 2S 42' Ion. 88 40' ( 1 XXIII 84 lat 23 13' Ion. 89 10' ( ( XXIV 11 37 lat. 24 43' Ion. 83 25' ( ( \\\ 47 321 lat. 28 42' Ion. 88 '40' ( ( \\\ 1 12 J6 hit. 24 34' Ion. 83 16' CCX.W II 12 36 lat. 24 34' Ion. 83 16' ( 1 \\\ 111 12 36 lat. 24 34' Ion. 83 16' ( ( XXIX 14 850 lat. 23 18' Ion. 8221' ( ( XXX 36 84 lat. 2r 13' Ion. 89° 10' ( ( XXXI 29 955 lat. 24°36' Ion. 84 05' ( ( XXXII 37 35 Alacran Shoals, Yucatan 1 ( XXIII 29 955 lat. 24 36'lon. 84°05' CCXXXIV 10 37 lat. 24°44'lon. 83°00' CCXXXV 3 243 to 450 off Havana CCXXXVI 36 84 lat 23 13' Ion. 89 10' ( ( XXXVII 12 36 lat. 24°34'lon. 83° 16' CCXXXII1 29 955 lat. 24 36'lon. 84 05' CCXXXIX 10 37 lat. 24°44'lon. 83°00' CCXL 29 955 lat. 24'36'lon. 8405' CCXLI 36 84 lat. 23 13'lon. 89 10' CCXLII 36 84 lat. 23 13' Ion. 89 10' CCXLIII 11 37 lat. 24"43'lon. 83*25' CCXL1V 36 84 lat. 23° 13' Ion. 89 10' 1969 BLAKE STATION NUMBERS 11 CCXLV CCXLVI CCXLVII CCXLVIII CCXLIX CCL CCLI CCLII CCLIII CCLIV CCLV CCLVI CCLVII CCLVIII CCLIX CCLX CCLXI CCLXII CCLXIII CCLXIV CCLXV CCLXVI CCLXV1I CCLXVIII CCLXIX CCLXX CCLXXI CCLXXII CCLXXIII CCLXXIV CCLXXV CCLXXVI 3 450 to 243 off Havana 1 29 955 lat. 24°36'lon. 84°05' 1 12 36 lat. 24°34'lon. 83° 16' 9 36 84 lat. 23°13'lon. 89°10' 1 11 37 lat. 24°43'lon. 83°25' 1 11 37 lat. 24°43'lon.83°25' 1 37 35 Alacran Shoals, Yucatan 5 37 35 Alacran Shoals, Yucatan 2 11 37 lat. 24°43'lon. 83°25' 1 37 35 Alacran Shoals, Yucatan 1 36 84 lat. 23° 13' Ion. 89° 10' 9 47 321 lat. 28°42'lon. 88°40' 1 36 84 lat. 23°13'lon. 89°10' 1 11 37 lat. 24°43'lon. 83°25' 1 36 84 lat. 23°13'lon. 89°10' 2 36 84 lat. 23°13'lon. 89°10' 3 36 84 lat. 23° 13' Ion. 89° 10' 1 14 lat. 23°18'lon. 82°21' 2 11 37 lat. 24°43'lon. 83°25' 1 36 84 lat. 23°13'lon. 89°10' 2 29 955 lat. 24°36'lon. 84°05' 1 29 955 lat. 24°36'lon. 84°05' 1 29 955 lat. 24°36'lon, 84°05' 1 29 955 lat. 24°36'lon. 84°05' 1 no loc. 1 31 1850 lat. 24°33'lon. 84°23' 1 no loc. 1 no loc. 1 no loc. 1 BREVIORA Museumi of Comparative Zoology Cambridge, Mass. 31 March, 1969 Number 314 THE CRANIAL ANATOMY OF THE PERMIAN AMPHIBIAN PANTYLUS Alfred Sherwood Romer Abstract. A detailed description is given of the skull structure of the early Permian tetrapod Pantylus, using both whole specimens and a skull serial sectioned and reconstructed in wax plates. This study definitely proves that Pantylus, often thought to be a reptile, is a microsaurian amphibian, and furnishes further proof that microsaurs cannot be considered reptilian ancestors. INTRODUCTION Pantylus, known in the past mainly from a heavily-built, heart- shaped skull carrying a powerful dentition, is an early Permian tetrapod from the Texas red beds whose systematic position has been open to considerable doubt. Cope, who obtained the first materials of this animal, originally considered it to be a batrachian (1881), but later classed it among the Reptilia (1892, 1896), as did a long series of later workers, including Case (1911, 1912, 1915), Huene (1913), Broom (1913), Williston (1916a, 1916b, 1925), and Wilson (1951). Two decades or so ago, however, in considering the nature of the Paleozoic microsaurs, it became obvi- ous to me that it was, as far as known, basically similar in structure to members of the microsaurian family Gymnarthridae, and hence should be placed in the order Microsauria (Romer, 1950). My conclusion has, I think, been generally accepted by recent workers (cf., for example, Dechaseaux, 1955; Gregory, Peabody, and Price, 1956; Tatarinov, 1964). However, it seems important to add as much as possible to our present inadequate knowledge of its struc- ture in order to attempt to fix its systematic position. But the importance of such study goes further. Most microsaurs are very poorly known anatomically. If (as I hope to demonstrate) Pantylus is a true microsaur, a study of its cranial structure may prove of value in fixing the phylogenetic position of that group as a whole. Microsaurs have often been claimed to be related to the 2 brhviora No. 314 captorhinomorph cotylosaurs and to be possibly ancestral to the reptiles, or to at least part of that class (ef.. for example, Williston. 1908; Westoll. 1942a, 1942b; Olson, 1947; Huene, 1948; Vaughn, I960, 1962; Brough and Brough, L967). Another possibility that has been suggested is that the microsaurs may be ancestral to modern amphibians, particularly the urodeles (Romer, 1950, etc.), although this suggestion is far from generally accepted (ct\, for example. Parsons and Williams. 1963). The pedigree of the microsaurs is in doubt. I have, since 1933, some- what arbitraril) combined the microsaurs with the aistopods and nectrideans in the Lepospondyli, in an expanded use of that term. But the interrelationships of the three groups are far from certain. And while the derivation of the l.abvrinthodontia from the rhipi- distian crossoptervgians seems clear, there has been little evidence to tie any of the three lepospondyl orders into this evolutionary series. Previous studies; materials. Cope's original materials consisted o( two imperfect skulls; from these he was able to describe much of the roof pattern and part oi the palate. later studies of these speci- mens by Case (1911) and lluene (1913) added little; however, here, as frequently, Broom (1913) was able to give greater depth to analysis of the specimens available to him. Mehl. in 19 12. described a fragmentary skull collected for the University o\' Chicago in 1908 by Paul C. Miller, but added little o\ interest. The latter collector, in expeditions of 1913 to 1916. obtained a series of specimens, in- cluding several skulls and some postcranial materials, from the Mitchell Creek locality; Williston ( 19 Km) described much of this material, giving an adequate account of the skull roof (except the temporal region), some further data on the lower jaw and palate. and a few scattered details regarding the braincase and postcranial skeleton. No further materials were found until, in 1939. Mr. A. Witte of Henrietta, in charge of a WPA group under the auspices of the Texas Bureau of Economic Geology, came upon a "nest" of speci- mens near Windthorst. Archer County. Texas; this suite, now en- tered as No. 40.001 (1-8) in the Texas Memorial Museum. Austin, includes several skulls and considerable postcranial materials. In Mr. Wittc's possession is a further small skull from this locality. Post-war trips to Texas from the Harvard Museum of Comparative Zoology (abbreviated as MCZ below) yielded further specimens — two skulls from the Archer City bonebed. a partial skull from Montaeue County, and two incomplete crania from southern Archer County. 1969 CRANIAL ANATOMY OF PANTYLUS 3 Study of this newer material, together with re-examination of older Chicago specimens, now makes it possible to give a nearly complete account of the skeletal anatomy of Pantylus. The post- cranial skeleton is now being studied by Dr. Robert L. Carroll of McGill University. Both Dr. Carroll and I are much indebted to Drs. E. C. Olson and R. Zangerl of Chicago, Drs. John A. Wilson and Wann Langston, Jr., of Austin, and Mr. Witte for the loan of materials. Also, I gratefully acknowledge financial aid for the preparation of material from National Science Foundation Grant GB 500. For cranial anatomy, some of the older Chicago and American Museum specimens were restudied. Valuable were, as noted, Har- vard skulls MCZ 2040 and 3302. The latter is incomplete and broken, but the parts preserved show excellent detail, including some braincase structure. MCZ 2040 is complete, except that the roof is broken off posteriorly on one side. It shows surface struc- tures perfectly, and the break is valuable in that it reveals much of the braincase. Several of the Texas Memorial Museum specimens show aspects of skull anatomy. Most important was No. 40,001 ( 1 ), which, with the consent of Prof. John A. Wilson, was serially sectioned and reconstructed, enlarged X 6, in wax plates. The sur- faces of each section were preserved by the cellulose "peel" method, and at each grinding two peels were taken. Although this sectioned specimen was extremely valuable in revealing internal structure, and is the main "document" on which Figures 3 - 12 are based, it proved disappointing in two regards. ( 1 ) Subsequent to burial, it was in- vaded by calcite, obviously crystallized out of ground water. This crystallization caused multiple small fractures in the skull, which have rendered details of sutures in the dermal bones difficult to determine. (2) Before burial, the skull received one or more vio- lent blows, which, apart from shattering a fraction of the roof (a matter of small moment), jarred loose the vomers, the epiptery- goids, the basisphenoid plus parasphenoid, and most of the otico- occipital elements of the braincase. For the most part, these ele- ments are still present in the specimen, lying, disarticulated and more or less battered, between the lower jaws. Their restoration and proper positioning in the braincase and palate presented obvious difficulties. Fortunately, most needed data on these disarticulated or demolished elements are available in MCZ 2040 and 3302. Stratigraphy and systematics. Cope's original specimens were collected by Jacob Boll in 1 880 from the "Big Wichita." In his first expedition, two years earlier, Boll had explored no higher in the Texas beds than exposures in the Belle Plains Formation of the 4 breviora No. 314 Wichita Group, between the site of the modern Diversion Dam on the Big Wichita River and the exposures along the Little Wichita south of Dundee. In 1880, however, he went somewhat further west, and there is reason to suppose that he collected at the Mitchell Creek locality from which Paul C. Miller later collected many valu- able amphibian specimens for S. W. Williston (Romer, 1928, 1935). This locality lies high in the Clyde Formation, very generally re- garded as the uppermost formation of the Wichita Group except for the overlying Lenders limestones. Miller's 1908 specimen, de- scribed by Mehl, is entered merely as from the "Big Wichita" but is probablv from Mitchell Creek. His later materials, described by \\ illistOll m 1916, are all definitely from Mitchell Creek. All the early described specimens, thus, are probably from a single locality and horizon near the summit of the Wichita Group. I he newer materials extend the range stratigraphically and topo- graphically. W ith two exceptions, all are from the Putnam Forma- tion, considerably lower in the Wichita Group (and in early days thought by many to be part of the Cisco Group of the Carboni- ferous). The Texas Museum specimens, and the individual later collected b\ Mr. Witte, were found in an exposure in Section 55, Block 3. of the Clark and Plumb Subdivision, about 4 miles north of Windthorst, Archer County; the horizon is about the middle of the Putnam Formation. One Harvard specimen, MCZ 2041, is from the J. J. Prather Survey, about 5 miles northwest of Stone- burg. Montague County; this is at about the same horizon as that of the Texas Museum specimens. Harvard skulls MCZ 2040 and 3302 are from a bonebed about a mile southwest of Archer City, Archer County, which is close to the summit of the Putnam Formation. Two final specimens, MCZ 1913 and 3610. are a poorly preserved skull and a snout fragment from Section 1834, Texas Emigration and Land Co.. north of the West Fork of the Trinity River in south- central Archer County and just below the presumed boundary be- tween the Moran Formation and the overlying Putnam. It now appears, hence, that Pantylus was in existence for at least most of Wichita time. Cope's original specimens were given the name Pantylus cordatus; no other species properly pertaining to the genus has been described. Cope in 1 896 described as Pantylus co'icodus a fragmentary speci- men, mainly a tooth row, from Coffee Creek in the Arroyo Forma- tion of the Clear Fork Group. However, these teeth are bulbous, pointed structures, quite unlike those of other Pantylus specimens. As various later writers have noted, they do not appear to belong to Pantylus (Case, 191 1; Gregory et al., 1956), and the specimen is 1969 CRANIAL ANATOMY OF PANTYLUS 5 perhaps a caseid. A jaw fragment from Coffee Creek to which Cope gave the manuscript name Pantylus tryptichus is probably a captor- hinid (Cummins, 1908; Case, 1911; Gregory et al, 1956). Wil- liston in 1913 gave the name Ostodolepis brevispinatus to a series of small vertebrae from Coffee Creek. In 1916 he noted the simil- arity of these vertebrae to those of Pantylus. "Ostodolepis," as de- scribed by Case (1929), is a gymnarthrid microsaur of a different nature, but as noted by Carroll and Baird (1968), there is no guarantee that Case's specimen is related to the type. It is possible that the stratigraphically lower Putnam and Moran materials are specifically distinct from the earlier described speci- mens from the Clyde. There is some variation in skull size, in general amounting to about 20 per cent, most skulls ranging in measured or estimated length from 72 to 90 mm; but this variation may be related to age or sex. The Witte specimen has an estimated length of about 36 mm, and one Chicago specimen from Mitchell Creek is but 39 mm long; but these are surely young individuals, presumably cospecific with the larger individuals found with them. There are dental differences, in the description and figures of earlier writers, regarding the number of teeth in the marginal rows, noted later, and the pattern of toothplate teeth. But one may strongly suspect that the patterns figured are in some cases diagram- matic and not entirely trustworthy. Quite surely age (and sex?) factors enter into variations in toothplate dentition. There may be individual differences, as well. For example, the skull sectioned by me has a large central tooth in both upper and lower toothplates, but Williston mentions a specimen in which such a tooth was pres- ent above but not below, and figures (how accurate I do not know) of other specimens show no such enlarged tooth in either upper or lower dentition. DESCRIPTION Skull roof (Figs. 1, 2, 7, 12). The skull of Pantylus is very mas- sively built, and very broad and low — its width, which, from a rounded snout, spreads out greatly in the cheek region, is nearly as great as the length. There is no otic notch. The cheek in the jugal region extends downward so as to cover externally a considerable area of the lower jaw. Skull height, even allowing for the depth of the overhanging cheek region, is little more than a third of the length. The broad roof is nearly flat, although bending sharply downward at the margins of the "table," and there is a somewhat overhanging snout. The roof bears, as in most early or primitive BRFVIORA No. 314 amphibians, patterns of ridges and intervening valleys radiating from the ossification centers of the dermal elements. There is no evidence of grooving for lateral line canals. The small skull in Mr. Wine's collection shows aearly straight sutures between roofing Fig. 1. Dorsal surface of skull, based on \l( / 2040; right posterior area restored. ■ 1*4. Abbreviations for tins and following figures: (;. angular; ar. articular; a\p. anterior splenial; ba, basal articulation of braincase with palatal structures; ho. basioceipital: bs, basisphenoid; c, coronoid; ch, choana. . exoccipital; /. frontal: jo. fenestra ovalis; fv, foramen for transverse vein in pituitary region; /', jugal; /, lacrimal; in. maxilla: n, nasal: op, opisthotic; p, parietal; pal, palatine; pur. prearticular; pf. postfrontal; pm. premaxilla; po, postorbital; pp. post- parietal; pr, prootic; prf, prefrontal: ps, parasphenoid; psp, posterior splenial; pt. pterygoid; l line striae on its dorsal surface. The epimeral areas o\' the metasomal segments are more or less rounded, except lor that of the segment of let: 4. which is truncated. The segment o\ leg 5 < I g. 2) is 104 • 161 ,». Between this seg- ment and the genital segment there is a distinct ventral interseg- mental sclerite (Fig. 3). The genital segment is 180 /< long, in dorsal view divided by lateral constrictions into a broad anterior hall' (width 187 ") with convex lateral margins and a much nar- rower and unexpanded posterior half (width 120 // ) with almost straight margins. In lateral view (Fig. 3) the thicker anterior half is separated from the posterior half by an abrupt transverse dorsal constriction (indicated also m Fig. 2). The areas o\' attachment of the egg s.ics are located laterally (Fig. 3). each area ( Figs. 4 and 5 ) bearing two naked setae 20 u and I 1 <> in length and a spini- form process about 8 ti long. I he three postgenital segments are 81 X 97 ft, 55 X 83 ,.. and 83 ■ 78 u from anterior to posterior. I he posteroventral margin of the anal segment is smooth. The caudal ramus (big. 6 ) is elongated. 148/< in length, its great- esl width proximally 31 ». and its least width distally 19 ,,. The 1 The specific name dentifer, from Latin dens = a tooth, and fero = to Hear, refers to the toothlike process on leg 5 in the female. 1969 COPEPOD FROM A JAMAICAN STARFISH 3 ratio of length to width (greatest dimensions) is 4.77: 1. The outer lateral seta is 78 fi and the dorsal pedicellate seta 29 /*, both of them naked. The outermost terminal seta is 81 /x, finely barbed in its distal half. The innermost terminal seta is 86 /x, with a few minute barbules near its tip. The two median terminal setae are 176 [i (outer) and 330 p (inner), both naked and inserted a little ven- trally. There is a minute setule 6 /x long on the outer proximal area of the ramus. The ornamentation consists of a few small hairs. The dorsal surface of the prosome and both dorsal and ventral surfaces of the urosome bear numerous hairs (sensilla) and re- fractile points. The ratio of the length of the prosome to that of the urosome is 1.1:1. The egg sac (Figs. 1 and 7) is an elongated oval, about 420 X 200 fi, extends as far as the anal segment, and contains 7-13 eggs, each approximately 100 /x in diameter, but somewhat variable in shape. The rostrum (Fig. 8) has a broad and rather truncated postero- ventral margin. The first antenna (Fig. 9) is 7-segmented, but a sclerotization on the ventral surface of the third segment (Fig. 10) suggests an in- tercalary segment. The lengths of the segments (measured along their posterior non-setiferous margins) are: 32 (48 /x along its an- terior margin), 75, 31, 33, 32, 20, and 18 p respectively. The for- mula for the armature is 4, 13, 6, 3, 4 + 1 aesthete, 2 -f- 1 aesthete, and 7+1 aesthete, as usual in the genus. All the setae are naked. The second antenna (Fig. 1 1) is 3-segmented, without a trace of division of the third segment. The first two segments are short, and each bears one seta. The longer third segment bears the usual inner group of three setae, is ornamented with a diagonal row of small spinules on its proximal posterior surface, and has distally three recurved claws and four setae. One claw (50 it along its axis and inserted terminally) is much larger than the other two claws (both about 19 p, one inserted on the subterminal anterior surface of the segment, the other subterminally near the large claw). One of the setae has a swollen base, the others are slender. All the elements are naked. The labrum (Fig. 12) has two divergent lobes, each with a small inner pointed process and a distal hyaline lamella. The mandible (Fig. 13) has on its concave margin a row of long slender spinules; its convex margin bears proximally a group of minute spinules fol- lowed by a large hyaline dentiform process and a row of smaller teeth. The long terminal flagellum has a row of spinules along the proximal half of its inner margin. The paragnath (Fig. 14) is a 4 breviora No. 315 small hairy lobe. The first maxilla (Fig. 15) bears four setae, three terminal and one subterminal. The second maxilla (Fig. 16) has an unarmed first segment. The second segment bears a surficial posterior seta and an inner setifonn spine. The terminal lash has on its convex side a large, weakly-selerotized tooth followed by a row of long slender spinules. and on its concave side a few barbules; across the base oi the lash there is a diagonal row of small spinules. The maxilliped ( Fig. 1 7 ) is 3-segmented. The slender first segment bears two minute distal inner spinules. The second segment is ex- panded medially and bears two naked setae and numerous spinules. The small third segment bears a barbed spine, a hyaline seta, and a minute spinule; the segment terminates in a large spiniform pro- longation with an attenuated tip. The ventral surface between the maxillipeds and leg 1 is slightly protuberant. A sclerotized line connects the bases of the maxil- lipeds (Fig. 18). Legs 1-4(1 igs. 19, 20. 21. and 22 ) have the following armature (the Roman numerals indicating spines, the Arabic numerals rep- resenting setae ) : P, coxa 0-1 Pa coxa 0-1 IV coxa 0-1 P, coxa 0-1 The inner seta on the coxa oi legs 1-3 is long and plumose, but in leg 4 this seta is short ( 15 p ) and naked. The inner side of the basis in leg 1 is smooth, but in legs 2-4 bears a row of hairs. The endopod of leg 4 (Fig. 22) is much shorter than the exopod. the ratio being about 1 :2.3. The first segment is 34 X 29 p, with its inner seta 105 fx. The second segment is asymmetrical, 43 X 29 //. and armed with an inner seta 110/4. and two terminal setae 61 fi (outer) and 75 /«. (inner). All four cndopodal setae have very fine distal barbules. The outer margins of both segments bear a row of hairs, and the second segment has a row of very small spinules on its distal margin. Leg 5 (Fig. 23) has a prominent, toothlike, distally-directed process on the inner margin of the free segment. This segment is 39 n long. 34 p in greatest width at the level of the tooth, and 46 p basis 1-0 exp 1-0 1-1 III. 1. 4 enp 0-1 0-1 1. 5 basis 1-0 exp 1-0 i-i III. 1. 5 enp 0-1 0-2 I. II. 3 basis 1-0 exp I-o I-I III. I. 5 cnp 0-1 0-2 I. II. 2 basis 1-0 exp 1-0 1-1 II. 1. 5 enp 0-1 2. 1 1969 COPEPOD FROM A JAMAICAN STARFISH 5 in greatest diagonal dimension. The two terminal setae are 91 p, and 39 ^, and the seta on the body near the insertion of the free seg- ment is 31 ft. All three setae are naked. Near the outer side of the free segment there is a slight oblique dorsal ridge with a series of extremely small points (spinules ?) . Leg 6 is probably represented by the two setae near the attach- ment of each egg sac (Fig. 5 ) . The color in life in transmitted light is opaque, the eye red. Male. — The body (Fig. 24) has a prosome more pointed an- teriorly than in the female. The length (without the ramal setae) is 1.18 mm (1.11-1.21 mm) and the greatest width is 0.35 mm (0.31- 0.41 mm), based on 10 specimens in lactic acid. The ratio of the length to the width of the prosome is 1.74: 1. The segment of leg 5 (Fig. 25) is 55 X 1 12 ^t. Between this seg- ment and the genital segment there is no ventral intersegmental sclerite. The genital segment is 156 X 160 /*, with rather flattened lateral margins in dorsal view. The four postgenital segments are 78 X 99 fx, 73 X 81 p, 62 X 68 /x, and 62 X 62 p from anterior to posterior. The caudal ramus resembles that of the female, but is smaller, 110X 25/*. The rostrum, first antenna (with no aesthetes added), second antenna, labrum, mandible, paragnath, first maxilla, and second maxilla are like those of the female. The maxilliped (Fig. 26) is slender and 4-segmented, assuming that the proximal part of the claw represents the fourth segment. The second segment bears two setae and two rows of spinules. The strongly recurved claw, 159 fx along its axis, bears two very unequal proximal setae and is divided about midway. The area between the maxillipeds and leg 1 resembles that in the female. Legs 1-4 are similar to those in the female, with the same seg- mentation and armature, but the second segment of the endopod of leg 4 (Fig. 27) is more symmetrical. Leg 5 (Fig. 28) has a free segment 26 X 13 //., without an inner toothlike process. The two terminal setae are 72 p and 32 it, and the adjacent seta on the body is 32 p. Leg 6 (Fig. 29) consists of the usual posteroventral flap on the genital segment, bearing two naked setae 32 /x and 44 p. The spermatophores (Fig. 30), attached to the female in pairs, are each about 120 X 48 p., not including the neck. The color in life resembles that of the female. 6 BREVIORA No. 315 COMPARISON WITH RELATED SPECIES S. dentifer may easily be distinguished from all other species in the genus by the presence of the toothlike process on leg 5 in the female. The genital segment of the female also has a configuration unlike that of any other species. In most species o\' Stellicola the caudal ramus is not more than twice as long as wide. Two species, however, have a relatively long caudal ramus. In S. gracilis (Thompson and A. Scott, 1903). the ratio of length to width oi the ramus is 4: 1. and in S. affinis Humes and Ho. 1967. it is 6.3-7:1 (in the female) and 7.5:1 (in the male). I he new species is readik separated from these two species. both oi which have a single claw on the second antenna and. in addition, differ in the two points mentioned in the previous para- graph. S. dentifer seems to be most closely related to the four West Afri- can species described b\ Humes and Cresse) ( 1958), S. frequens, S. astro pectinis, S. luidiae, and S. hiatus, all of which similarly have three claws on the second antenna. Unlike most other species of Stellicola, the inner edge of the free segment of leg 5 in these four species is somewhat swollen or irregular; a toothlike process is ab- sent, however. The twenty-one species of Stellicola may be grouped geographi- cally on the basis of the number o\' claws on the second antenna. Those from the Indian and western Pacific oceans ( 15 species) have one such claw. The only known species from Europe and the Medi- terranean Sea. S. clausi ( Rosoll, 1 889 ). has two claws (see Bocquet and Stuck. 1962). The five species from the tropical Atlantic Ocean (four from West Africa and one from Jamaica) have three claws. SUMMARY The new species Stellicola dentifer is associated with the asteroid Luidia clathrata (Gray) in Jamaica. The copepod may be recog- nized within the genus by the prominent toothlike process on leg 5 in the female. Four West African species of Stellicola appear to be related to the new Jamaican form. LITERATURE CITED Bocquet. C, and J. H. Stock 1962. Copepodes parasites d'invertebres des cotes de la Manche. IX. CyclopoYdes associes a Marthasterias glacialis (L.). Arch. Zool. Exp. Gen., 101 I Notes et Revue) (2):79-91. 1969 COPEPOD FROM A JAMAICAN STARFISH 7 Humes, A. G., and R. F. Cressey 1958. Four new species of lichomolgid copepods parasitic on West African starfishes. Bull. Inst. Fr. Afrique Noire, 20, ser. A (2) : 330-341. Humes, A. G., and J.-S. Ho 1967. New species of Stellicola (Copepoda, Cyclopoida) associated with starfishes in Madagascar, with a redescription of S. caeruleus (Stebbing, 1900). Bull. Brit. Mus. (Nat. Hist.), Zool., 15(5): 201-225. KOSSMANN, R. 1877. Entomostraca (1. Theil: Lichomolgidae). In: Zool. Ergeb. Reise KUstengeb. Rothen Meeres, erste Halfte, IV, pp. 1-24. Rosoll, A. 1889. Uber zwei neue an Echinodermen lebende parasitische Cope- poden: Ascomyzon comatulae und Astericola clausii. S.-B. Math.-Naturw. CI. Ak. Wiss. Wien, 97 (Abt. I, Heft 4): 188- 202. Thompson, I. C, and A. Scott 1903. Report on the Copepoda collected by Professor Herdman, at Ceylon, in 1902. Rept. Govt. Ceylon Pearl Oyster Fish. Gulf of Manaar, part I, suppl. rept. no. 7, pp. 227-307. (Received 14 January 1969.) 8 HREV10RA No. 3 1 5 Figures 1-8. Stellicola dentifer n. sp.. female: 1. dorsal (A); 2, urosome, dorsal (B); 3, segment of leg 5 and genital segment, lateral (B); 4. area of attachment of egg sac. dorsal (C): 5, area of attachment of egg sac, lateral iC ); 6, caudal ramus, dorsal (D); 7. egg sac, dorsal (E); 8. rostrum, ven- tral (F). 1969 COPEPOD FROM A JAMAICAN STARFISH 9 Figures 9-18. Stellicola dentifer n. sp., female: 9, first antenna, anterodor- sal (G); 10, third segment of first antenna, posteroventral (H); 11, second antenna, posterior (D); 12, labrum, ventral (D); 13, mandible, posterior (C); 14, paragnath, posterior (C); 15, first maxilla, posterior (C); 16, sec- ond maxilla, posterior (C); 17, maxilliped, antero-inner (C); 18, area be- tween maxillipeds and leg 1, ventral (G). 10 imrvioRA No. 3 1 5 ^ / 21 J 22 19 20 . Figures 19-22. Stellkola dentifer n. sp.. female: 19. leg 1 and intereoxal plated anterior (G); 20. leg 2. anterior (G): 21. leg 3. anterior (G): 22. leg 4 and intereoxal plate, anterior (G). 1969 COPEPOD FROM A JAMAICAN STARFISH 11 Figures 23-30. Stellicola dentifer n. sp. Female: 23, leg 5, dorsal (C), Male: 24, dorsal (E); 25, urosome, dorsal (B); 26, maxilliped, anterior (D); 27, endopod of leg 4, anterior (G); 28, leg 5, dorsal (C); 29, leg 6, ventral (D); 30, spermatophores, attached to female, lateral (G). BREVIORA Museum! of Cooiparsitive Zoology Cambridge, Mass. 30 April, 1969 Number 316 CYTOTAXONOMIC STUDIES ON SOME UNUSUAL IGUANID LIZARDS ASSIGNED TO THE GENERA CHAMAELEOUS, POLYCHRUS, POLYCHROIDES, AND PHENACOSAURUS, WITH BEHAVIORAL NOTES G.C.Gorman1 R. B. Huey2 E. E.Williams INTRODUCTION Abstract. Chromosome data are presented for four species of iguanid lizards. Phenacosaurus heterodermus has a diploid number of 36, with 12 metacentric microchromosomes and 24 microchromosomes. Chamaeleolis porcus also has 12 metacentric macrochromosomes. The exact number of microchromosomes was not ascertained, but it is quite similar to that in Phenacosaurus. The karyotypes found in these two species resemble the primitive iguanid condition and that most frequently found in alpha Anolis. Polyckrus fetnoralis has an apparent diploid number of 26, with 10 pairs of acrocentric and one pair of submetacentric macrochromosomes, and two pairs of microchromosomes. Polychrus peruvianas has 20 acrocentric macro- chromosomes and 8 microchromosomes (2n = 28). A karyotype with low diploid number (2n = 30 or less) and few metacentric macrochromo- somes is very unusual in iguanid lizards — previously reported only for Polychrus marmoratus. Its presence in the two presently studied Polychrus confirms their very close relationship inter se, but gives no clue to their relationship to other members of the family Iguanidae. P. peruvianas has previously been assigned to the monotypic genus Polychroides. However, osteological and cytological data show its very close relationship to Poly- chrus, and we formally propose the synonymy of Polychroides with Poly- chrus. Notes on behavior and ecology are appended for the four species. Although there may be a nomenclatural problem surrounding the use of the name P. femoralis, our Peruvian animal resembles P. femoralis from Loja, 1 Museum of Vertebrate Zoology, University of California, Berkeley, Cali- fornia 94720. - Biological Laboratories, University of Texas, Austin, Texas. 2 BREVIORA No. 316 Ecuador. This is the first record of an apparent Polychrus femoralis from Peru. We have recentl) obtained several poorl) known iguanid lizards Oil Held trips to Peru (RBH). and Cuba and Colombia (GCG). Although our observations and chromosome data arc meagre, the) provide insight into the relationships of these lizards. Field and behavioral notes are appended because of the paucit) of in- formation on these species and because we have the first record of an apparent Polychrus femoralis? from Peru. The genus Chamaeleolis is endemic to Cuba and the adjacent Isle of Pmcs. [.on^ considered monotypic, Garrido and Schwartz (1968) have demonstrated that there are two distinct species. Both are arboreal, moderatc-si/cd lizards (snout-vent of adult males 155 mm), quite closel) related to Anolis. The) share with anoles extensible clew laps and adhesive lamellae on the toes but differ in a number of OSteological features, the most conspicuous of which is a parietal ercst that extends tar back over the neck (Etheridge, 1960), giving the animals a marked resemblance to true old-world chamaeleons. Phenacosaurus is an iguanid genus with ver) close affinities to Anolis. Dunn ( I W44 ) listed a series ol characters by which the genus was supposed to differ from Anolis (e.g. a prehensile tail, a heterogeneous scutcllation with large and small scales intermixed, a dorsal crest of enlarged scales), but Etheridge (PHiO) pointed out that all of these characters occur individual!) in a number of species of Anolis. However, because of a number of osteological peculiarities (low number of sternal ribs, high number of paraster- nal chevrons), Etheridge retained the genus. Phenacosaurus is known predominantly from high altitudes (1800-3500 m) in the Eastern Andes of Colombia. There are currentl) four recognized species, but richteri and paramoensis are undoubtedK synonyms of I', heterodermus, and there is at least I he single specimen reported here runs down i<> P. femoralis in the ke\ provided bv H. W. Parker (1935: 516). Examination of comparative ma- terial (all of it femoralis h\ Parkers key) indicates that there may he as man\ as three distinguishable populations. \ specimen in the United States National Museum from Guayaquil, the type locality of femoralis. has markedK larger scales than Parker's l.oja specimens, which closeh resemble the Peruvian animal. Pending revisionarv studies, we believe it to be prefer- able to use the name femoralis for all this material, including our Peruvian animal. 1969 CYTOTAXONOMY OF IGUANIDS 3 one undescribed species from Ecuador (J. D. Lazell, Jr., personal communication). Polychrus is a genus of arboreal lizards from Central and South America (including Trinidad) comprising about five distinct species (Parker, 1935). Etheridge (1960) compared 22 charac- ters in Anolis and Polychrus (excluding characters unique to Anolis or common to the family Iguanidae) and found that 20 were shared by Anolis and Polychrus. He wrote, "Although there appears to be a close correspondence between Polychrus and the anoles, there are striking differences. Most of these differences, however, are features which are either unique to the anoles or unique to Polychrus and therefore tell us little more than that the anoles are separated from other members of the family, including Polychrus, by a wide morphological gap" (Etheridge, 1960: 111). He also pointed out that Polychrus and Chamaeleolis are highly arboreal, "and unusual in their habit of slow, deliberate move- ment. The similarities of these genera may well be parallel adapta- tions to a similar way of life, yet the occurrence of multiple parallel developments may, in itself, be an indication of relationship be- tween the groups in which it occurs" (Etheridge. 1960: 113). The monotypic genus Polychroides was erected by Noble (1924) when describing a new species of iguanid lizard (P. peruvianus) from the provinces of Cajamarca and Piura, northern Peru. Noble's comments on the affinities of the new genus were quite confused and are here quoted verbatim. On p. 109 in "Remarks" following the generic diagnosis he wrote: "The generic status of many of the slow-moving arboreal iguanids is very uncertain. The species described below cannot be referred to either Enyalius, Enyalioides or Polychrus as at present defined. It seems most closely related to Polychrus with which it agrees in its femoral pores, large head scales, subequal third and fourth toe and its sacculated lung. T have seen both Polychrus and Polychroides alive and have been struck by their great similarity in behavior. The pronounced nuchal crest of the latter readily distinguishes it from the former." On p. 11 0, however, in the "Remarks" which follow descrip- tion of the species he stated: "This species is closely allied to Enyalioides festae Peracca from which it differs in its larger head scales and gular sac, also in certain differences in the scutation of the digits and head. I would not hesitate to refer it to the genus Envalioides were it not that this procedure would require a consid- erable modification of our present conceptions of that genus." 4 BREVIORA No. 316 Etheridge I I960) was unable to find an) skeletal differences be- tween Polychrus and Polychroides in \-ra\ studies of iguanid gen- era, and he therefore cited peruvianus in his list of material exam- ined as another species of Polychrus. Onl\ the external charac- ter of a dorso-nuchal cresl distinguished peruvianus from other Polychrus. Chromosomes have been useful in elucidating relationships of iguanid lizards. I he majority of species studied have a karyotype consisting of six pairs of metacentric macrochromosomes and eleven or twelve pairs of microchromosomes (Gorman, Atkins and Holzinger, 1967; Gorman and Atkins, ll><>7. 1968). One species, Plica plica, showed apparent centric fissioning with lour pairs of acrocentric macrochromosomes. I he chromosomes o\ the single species of Polychrus examined. /'. marmoratus, were so different irom all other species of iguanids ol which karyotypes are known that it was difficult to relate this genus to am of the others (Gor- man, Atkms, and Holzinger, 1967). Female /'. marmoratus have a diploid number of 30, consisting ol 20 acrocentric macrochromo- somes and 10 acrocentric microchromosomes. Males have a diploid number of 29, with l metacentric and 19 acrocentric macrochromosomes, and 9 acrocentric microchromosomes. In both Anolis (Gorman, llM7) and Sceloporus (Lowe, Cole, and Patton, 1967), species have now been found with high numbers of acro- centric macrochromosomes. I hese speeies. however, are charac- terized h\ high diploid numbers, greater than 2n JO, the formula which appears primitive lor the family [guanidae (Gorman, et al., 1967 I I he probable explanation tor these high numbers is centric fission of the metacentric macrochromosomes (Gorman, Baptista, and Bury, 1969). MATERIALS AND METHODS Chromosome preparations were made b\ tissue culture of whole blood following methods alread) outlined; b\ culture of marrow ob- tained from the femur and treated in the same fashion as blood; and b\ direet testis preparations (see Gorman, et al.. 1967). The specimens used were one female Polychrus peruvianus (Mu- seum o\' Vertebrate Zoology, no. X2X34). one female Polychrus femoralis (M.V.Z. 82835), one male Chamaeleolis porcus (Mu- seum of Comparative Zoology, 100472). and one male Phena- cosaurus heterodermus (M.C.Z. 104409). Chromosome spreads were photographed and karyotypes ana- lyzed as best we could. Unfortunatelv. for Chamaeleolis. Polychrus peruvianus, and P. femoralis, we obtained few mitotic divisions. 1969 CYTOTAXONOMY OF IGUANIDS 5 and we are unable to present fully definitive karyotypes or diploid numbers, though the material permits interesting comparisons. Numerous divisions from the testis of Phenacosaurus were studied. RESULTS AND DISCUSSION Only four clearly resolved metaphase plates were found in P. peruvianas, all with 28 chromosomes, all acrocentric. There is not a sharp break in size between macro- and microchromo- somes; however, 20 might be termed macrochromosomes and 8 microchromosomes (Fig. la). This is very similar to the karyo- type of female Polychrus marmoratus, and this is only the second species of iguanid reported to date that has a karyotype of all acrocentric chromosomes. So few cells were seen that we cannot safely assume that the diploid number is 28 — but it is quite clear that the karyotype of P. peruvianas resembles that of P. marmora- tus to such a great extent that considering these forms as members of separate genera obfuscates their close relationship. We formally propose the synonymy of Polychroides with Polychrus. Few divisions were obtained in our specimen of P. femoralis — but, again, there are enough data to establish both its close rela- tionship to the other Polychrus in the similarity of the specialized karyotype and its uniqueness among the species thus far examined. The diploid number appears to be 26. (This is based upon only four metaphase plates; additional spreads were seen, but, because of overlaps, prec:se counts could not be made.) In all cells, there is one pair of submetacentric chromosomes (Fig. lb). Within the genus Polychrus, a submetacentric chromosome had previously been seen only in the male of P. marmoratus, where it is quite clearly the Y chromosome (Gorman, et al., 1967). In P. femo- ralis also, the largest chromosome pair is clearly subacrocentric with minute short arms present. There is a total of 1 1 pairs of macrochromosomes and 2 pairs of microchromosomes in the cells that were carefully analyzed. The karyotype of P. femoralis is most likely derived from a marmoratus-\\ke karyotype of 2n=30. Centric fusions of microchromosomes to macrochromosomes would account for the reduction in diploid number and the appear- ance of submetacentric and subacrocentric chromosomes. Numerous metaphase plates were observed in Chamaeleolis porcus, but we lack definitive information on the number of micro- chromosomes. It is quite clear that the karyotype resembles the "typical" iguanid in having six pairs of metacentric macrochromo- somes and a series of more than 20 microchromosomes (Fig. 2). I!RI \ IOR \ No. 3 Id (! 5 ■ 1 3 1 1 ■ « a /I '• •»«•»'»« X* Oil"* Fig. I. Karyotypes of Polychrus. a., P. peruvianus female. There are 28 acrocentric chromosomes Rone marrow preparation, Giemsa Stain, b., P. femoralii female, ["here are 26 chromosomes; one pair nop row. far right) is submetacentric. Leukocyte culture, Giemsa stain. Onl\ testis preparations were made for Phenacosaurm heter- odermus. There were numerous meiotic and mitotic divisions. In meiosis we can clearly see six large macrobivalents and 12 con- siderably smaller microbivalents (lis:. 3). In mitosis, it was quite clear that the microchromosomes were metacentric. Thus, the diploid number is 36, with 24 microchromosomes. 969 CYTOTAXONOMY OF IGUANIDS Fig. 2. Mitotic metaphase of Chamaeleolis porcus male showing the "typical iguanid" condition of 12 metacentric macrochromosomes, and a sharp break in size between the macro- and microchromosomes. Among Anolis, Etheridge (1960) defined two major groups termed alpha and beta. Chromosome studies (Gorman, et al., 1967) have shown that the primitive alpha karyotype is identical with that of the primitive iguanid condition ( 12 metacentric macro- chromosomes, 24 microchromosomes). In a recently proposed classification of iguanids on the basis of structure of the caudal vertebrae (Etheridge, 1967), Chamaeleolis and Phenacosaurus were placed in a group with the alpha anoles; chromosomes sup- port this classification. Polychrus remains a puzzle within the iguanids. Now that three species have been studied it is quite clear that they are close to each other inter se, but distinctly different from anoles and anoline genera such as Chamaeleolis, Phenacosaurus, and Anisolepis (Gor- man, et al., 1967). Since this paper was originally written, we have received the following new data. 8 Bkl \ IORA No. 3 I o On November 26, 1968, Dr. John Wright oi the Los Angeles County Museum collected a male Polychrus peruvianus at 3.5 km east of the junction between Bagua Grande and Bagua Chica, 15.5 km west of Bagua Grande, Ama/onas Dept.. Peru, at an altitude of 2.0(H) feet. (Field catalog number P-c>^ 7 ; to be deposited in L.A.C.M ) I es;es were mineed and allowed to settle in a hypotonic citrate solution (as no centrifuge was available), the citrate was removed with an e\e-dropper. and fixative was added. I his field prepara- tion was then brought baek to the United Stales, where slides were made some two months later. Dr Wright has kindb consented to our examination of the slides and Utilization of the data. At diakinesis there are 13 bodies. Fig. 3. Diakinesis in Phenacosaurui heterodermus. There are six large bivalents, and 12 microbivalents. n = IS. Large round black area is a sperm head, not part of the meiotic figure. 1969 CYTOTAXONOMY OF IGUANIDS 12 bivalents, and a trivalent. The male diploid number is thus expected to be 27. One mitotic metaphase was seen, and 27 chromosomes were counted. Position of the centromere could not be ascertained. One clear metaphase II cell had 14 chromo- somes, all acrocentric. These data are consistent with the sex chromosome situation known in Polychrus marmoratus, in which the males have a diploid o \ 4Mf Fig. 4. Diakinesis of Polychrus peruvianus male. There are 13 bodies, 12 bivalents, and a sex trivalent ( arrow ) . 10 BREVIORA No. 3 Id number one lower than the females, the Y chromosome is meta- centric (not demonstrated for P. peruvianus), and a sex bivalent is clearl) evident at meiosis (Gorman, Atkins, and Holzinger, L967). The finding of 12 bivalents and a trivalent in /'. peruvianus, with an expected male 2n o\ 27. gives further strength to our assump- tion that female 2n 28 (the latter figure having been arrived at with the examination of relatively lew cells). Figure 4 illustrates diakinesis in /'. peruvianus. NOTES ON POLYCHRUS POLYCHRUS PI Rl V IANUS Five specimens of Polychrus peruvianus were collected: MYZ S2S34 F.I Arenal. Rio Huancabamba, elevation 3000 82388 ft.. 7 km V 50 km 1 of Olmos, Dept. Caja- 83678 marea. Peru. 83679 MVZ 82413 (skin and skeleton). Imgo. Rio I leubamba. elevation 3000 ft., 30 km S, 41 km E of Bagua, Dept. Amazonas, Peru. I 1 \renal is on a terrace above the Rio Huancabamba. The hillsides are dr\ and rocky, and the terrace has a predominance of mesquite (Prosopis) interspersed with cultivated areas. All specimens were collected on mesquite branches. I he I ineo individual was captured b\ Dr. Carl Koford at nighl on a horizontal branch o\ an unidentified tree. Stomach contents included several Hymenoptera and a leaf fragment. I he largest ovarian follicles were less that 2 mm in diameter. This localit) is much wetter than El Arenal. Bromeliad-laden trees up to 40 feet are found on the steep hillsides, and dense second-growth vegetation covers the valle) floor. Body coloration in life I MVZ 82834) was medium yellow-green with a narrow, pale yellow band running from the eye to the groin and olive-drab saddles on the dorsum and tail. Small irregular yellow marks were spaced between the saddles. Some capacity to darken the body colors was observed in captivity. The tail of this specimen was slightly prehensile. P. femoralis gave no evidence of this ability. Both species emitted a nasal salt solution in captivity (see Templeton. 1967 ). P. peruvianus has a more protrusive gular flap than P. femoralis. In addition, only P. peruvianus possesses a row of raised scales forming a mid-gular crest. 1969 CYTOTAXONOMY OF IGUANIDS 11 POLYCHRUS FEMORALIS On 31 August 1967, one gravid female of this species was cap- tured near Molino, devation 2300 ft., 21 km E, 7 km N of Olmos, Dept. Lambayeque, Peru (5.9 S/79.60 W). This specimen (MVZ 82835) represents a considerable range extension from the record in Loja (Parker, 1932) and an apparent Peruvian record. She was oriented head down on a vertical branch of a plant about 5 ft. tall and remained motionless until capture. The locality is in a narrow river valley. Deciduous woodland alternates with open areas cleared for farming. She was kept alive and placed in a runway with a temperature gradient beginning 18 September. Water was sprinkled daily at the cool end. On 27 September she laid 12 eggs at the cool, moist end of the runway. The eggs were removed and placed in an incu- bator. Egg size eight hours later averaged 19.6 mm in length and 1 1.3 mm in width. The shell texture was soft and leathery. Mold developed on the eggs by 4 October and they subsequently decayed. In captivity movement was rare. When prodded, she exhibited a leaping form of locomotion described by Boker (1935) as "bipedal leaping" for Polychrus marmoratus. Davis (1953) ob- served a similar method of locomotion in Corythophunes cristatus. in life she was light yellow-green with pronounced brown saddles on the dorsum. The capacity for color change in this specimen was less marked than in P. peruvianas. Behavior in Captivity Davis (1953) outlined a sequence of four defensive responses used by Corythophanes cristatus against apparent predators. These can be grouped into two general categories: I. Passive defense A. Behavioral and morphological camouflage B. Catalepsy to eliminate all movement not associated with a positional change and thus decrease the probability of detection H. Active defense A. Active postural movements to increase the apparent size of the animal B. Aggressive biting as a last resort Davis stated that the defensive response given was geared to the relative immediacy of the threat to the animal's safety. The more immed:ate the threat, the more active the response. 12 BREVIORA No. 316 Both Polychrus peruvianus and /'. femoralis exhibited this sequence of defensive traits in captivity hut did differ in degree of response. Since onl\ one specimen of each species was ob- served, these notes should be considered tentative. Behavioral and morphological camouflage is well marked in both species. The colors and patterns render detection oi these animals difficult in their arboreal habitats. Their relative im- mobility and capacit) for color change accentuate this conceal- ment. In both the held and captivity, /'. femoralis relied more on immobility than on other defensive responses. Behavior similar to the cataleps) oi walking sticks | Phasmidae) has been reported for several arboreal lizards (reviewed in Davis) and was present in these two species as well. /'. peruvianus could be easily induced into this behavior b\ forcibl) altering its position, but both used it when coining to rest after voluntary movement. When matched with other types o\ camouflage, Davis felt that this behavior aided in concealment. In response to a threat, main animals arc recorded to increase the apparent size o\ the bod\ or parts of the bod) b\ behavioral posturing. I his behavior is manifest in social encounters, as well. Postural movements common to both Polychrus species involve lateral compression ol the body, expansion of the throat fan. and head bobbing. I he plane of expansion is sagittal and presented broadside for both, as :s the ease in CorythophaneS. The expanded area is thus displayed to maximum advantage. /'. peruvianus would often gape when posturing. If approached closely, she would lunge forward, maintaining balance by the two hind legs and tail. She would bite vigorously if given the opportu- nity In contrast. /'. femoralis could rarely be induced to bite. But it would often attempt to escape b\ leaping. Of the two species o\ Polychrus under discussion, the behavior of /'. peruvianus was more similar to that n\ Corythophanes than was that of /'. femoralis. It is perhaps of significance that /'. femoralis, having a much less pronounced gular Map than the other two species, relied less on active postural movements and more on passive camouflage. NOTES ON CHAMAELEOLIS Although rare in collections. Chamaeleolis is widely distributed throughout Cuba. Through the help of Orlando Garrido. we ob- tained a juvenile male C. porcus from La Florida. Sabanilla 1969 CYTOTAXONOMY OF IGUANIDS 13 Oriente Province, and a larger (but subaduit) male from La Casimba, eight kilometers west of Maisi, Oriente Province. Wilson (1957) published a note on the behavior of a captive Chamaeleolis. Although he called this C. chamaeleontides (sic), the specimen has been subsequently re-identified by Schwartz as porcus. Wilson reported, "its behavior was strikingly chamaeleon- like and quite different from that of several species of Anolis (including the giant A. equestris Merrem) which the author has observed in captivity." He stated that it was very sluggish even when freshly captured, and that it could be left on a laboratory table without much danger of its wandering. Often, it would re- main in the same spot for hours or even days without changing its position. When confronted with food in the form of living in- sects, it would move deliberately and without hurrying. Experi- ments using a live male Anolis equestris and a cardboard model resembling a male Chamaeleolis failed to elicit any aggressive response. Only once, in the year that he had the specimen, did it extend its dewlap — and this was without any apparent external stimulus. Our observations are not at all in accord with this descrip- tion. Our specimens would actively jump toward prey (crickets, grasshoppers, and mealworms). They would often scamper away when left out in the open. Garrido and Schwartz {op. cit.) point out that C. porcus is "at least capable of quick and decisive action. " They give a detailed account of the escape behavior of one speci- men encountered in the field. Our Chamaeleolis were not so quick as Anolis, but certainly did not show the incredibly slow move- ments of true chamaeleons. Furthermore, they were quite aggres- sive. They would display to each other by extending the dewlap fully, opening the mouth and protruding the tongue. They also displayed in similar manner to A. equestris. When handled, they would extend the dewlap and often would bite. Unfortunately, no films were taken of the displays It appears, then, that the social behavior of Chamaeleolis has been misunderstood. Its reported sluggishness and immobility appear to be facultative rather than obligatory and may well be associated with its cryptic coloration. The ability to move quite quickly, and the fact that ano!e-like territorial responses were easily elicited implies that the behavior and social structure of Chamaeleolis may be quite similar to that of most anoles. 14 BRE\ IORA No. 3 16 NOTES ON PHENACOSAURUS Previous notes on Phenacosaurm have dealt with specimens in captivit) (Osorno-Mesa and Osorno-Mesa | 1946], on shedding of skin, incubation of eggs, feeding habits, etc., and Kastle 1 1965], a detailed ethological stud) with particular emphasis on social behav ior ). On Jul\ 10. 1968, Dr. Jorge Hernandez, oi the Universidad Nacional. Bogota, led Gorman and J. F. Lynch on a one-da) trip specificall) to collect Phenacosaurus in Chia (Cundinamarca), a semi-rural suburb of Bogata (at about 35 km distance, alt. ca. 2700 m) and collected between II am and 2:30 pm. The da) was overcast, with intermittent showers, and cold. A thermometer was not available, but the temperature for most of the time was probabh not in excess of 14 ( . the mean temperature of that immediate area. I he roads were lined with shrubbery, often blackberries, which separated either pasture-land or homes. I luce Phenacosaurus were in the first blackberry bush ex- amined. I wo. a male and a female, were collected. In the course of the afternoon, 14 of these supposedly rare lizards were collected. Most often, more than one was taken at a time, usually in groups of two. sometimes of three. One pair was actually in bodil) contact the male's chm was touching the tail of the female. All but two of the li/ards were on blackberrs bushes, generally 1-3 feel above the ground, and usualK on a bare branch. It is likelx that as main were missed as were caught — especially tin se in leaf) parts of the bushes. I he escape response is to drop down, often clinging to a branch with the prehensile tail, and then to slink slowl) away. I he most usual perch position was almost horizontal; onl) one individual was perched vertically, head down In review of bod) temperatures in reptiles. Brattstrom ( 1965) listed 18.0 C. as the minimum voluntar) temperature tor an iguanid lizard. This was l ma notata, which, in fact, has a mean bod) temperature of 38.6 when active, and the IX probably represents an unusual reading. Normally, iguanids living in cold temperatures bask in sunlight until warmed to a preferred body temperature of 33 ! 3 . An extreme example was reported bv Pearson (1954) for Liolaemus multiformis at high altitudes in Peru. He found that the intense isolation enables these species 1969 CYTOTAXONOMY OF IGUANIDS 15 to warm to body temperatures of 35°, with shade temperatures as low as 11°. Phenacosaurus was remarkable for an iguanid lizard because it was normally active at low temperatures with no opportunity to warm up. We consider it quite possible that its preferred body temperature is well below that of any other iguanid reported to date. ACKNOWLEDGMENTS We thank Dr. Leonard Atkins, Mr. Fernando Ortiz, Mr. Bruce Bury, Miss Ida Leone, and Mrs. Charlotte Kayavas for technical assistance in the chromosome work, which was supported in part by Children's Bureau Project No. 906 and NSF Grant GB-6944. Field work in Cuba was supported by Sigma Xi, the Graduate Department of Biochemistry, Brandeis University, and Academia de Ciencias of Cuba. Special thanks are due to Sr. Miguel Jaume and Sr. Orlando Garrido for their help in all our work in Cuba. Field work in Peru was supported by the Museum of Vertebrate Zoology. The help of Dr. C. Koford, Mr. R. B. Hamilton, and Mr. F. McCollum in the field, and Drs. Hans and Maria Koepcke and Mr. Robert Conly, our hosts in Peru, is greatly appreciated. Field work in Colombia was jointly spon- sored by the Museum of Vertebrate Zoology and NSF Grant GB-6944. We thank Mr. James F. Lynch for help in collecting and for his observations, and we are particularly indebted to Dr. Jorge Hernandez C, of the Universidad Nacional in Bogota. He brought us directly to the Phenacosaurus locality and provided facilities at the University for making chromosome preparations. Comparative material permitting provisional identification of the Peruvian Polychrus femoralis was loaned by Miss A. G. C. Grandi- son (British Museum, Natural History) and Dr. James Peters (United States National Museum). We are grateful for their help. LITERATURE CITED Borer, Hans 1935. Einfuhrung in die vergleichende hiologische Anatomie der Wirheltiere. I. Jena, G. Fischer, xi + 228 pp. Brattstrom, Bayard H. 1965. Body temperatures of reptiles. Amer. Midi. Nat., 73: 376-422. Davis, D. D. 1953. Behavior of the lizard Corythophanes cristatus. Fieldiana: Zool., 35: 3-8. 16 brfviora No. 316 Dunn, E. R 1944. [Tie lizard genus Phenacosaurus. ( aldasia, 3: 57-62. Etui riix.i . R 1960. The relationships oi the anoles (Reptilia, Sauna, Iguanidae): An interpretation based on skeletal morphology. Ph.D. Thesis. Dept. of Zoology, Univ. of Michigan (Univ. Microfilms, Ine.. Ann \i boi . Mich. ). 1967. Lizard caudal vertebrae Copeia, 1967:699-721. GARRIDO, ().. AND A. SCHW \R I / 1968. Cuban lizards ol the genus ( hamaeleolis. Quart. Jour. Florida tcad Sci., 30: 197-220. GORM \N. ( , ( 1967 Studies on the evolution and zoogeograph) of Anolis, Ph.D. rhesis, Depl of Biology, Harvard University GORM W I i ( . \M> I . \ IklNs 1967. The relationships of tnolis of the roquet species group (Sauria Iguanidae). II. Comparative chromosome cytolog) Sysl /ool.. 16: 137-143 1968. New karyotypic data on 16 species of tnolis from Cuba, Ja- maica, and the ( ayman Islands Herpetologica, 24: 13-21. GORM VS. I i ( I \ I kiss. \\|> I HOLZINGER 1967. New karyotypic data on 15 genera of lizards in the family Iguanidae, with a discussion of taxonomk and cytologies] im- plications ( Oogenetics, 6: 286 299 Gorman, G. ( .1 Baptista, and R. B Hi ki 19hl> ( hromOSOmes and SCeloporine relationships with special refer- ence to the horned lizards Mammalian ( hromosome Newslet- ter. 10 6-10. k \STLE, WLRM R 1965. /in Ethologie des \nden- tnolis Phenacosaurus richteri. Zeit- schnft fur rierpsychologie, 22: 751-769. Lowe, C. H .. C. J Cole, and J. L. Patton 1967. Karyotype evolution and speciation in lizards (genus Scelo- porus) during evolution of the North American desert. Syst. /ool. 16: 296-300. Noble, G. K 1924. New lizards from northwestern Peru. Occ. Pap. Boston Soc. Nat. Hist.. 5: 107-113. Osorno-Mi sa, Hi rnando. ind Ernesto Osorno-Mesa 1946. Anotaciones sobre lagartos del genero Phenacosaurus. Caldasia, 4: 123-130. P.KRKhR. H. W. 1932. Some new or rare reptiles and amphibians from southern Ecua- dor. Ann. Mag. Nat. Hist.. (10) 9: 21-26. 1969 CYTOTAXONOMY OF IGUANIDS 17 1935. The frogs, lizards and snakes of British Guiana. Proc. Zool. Soc. London. 1935: 505-530. 1938. The vertical distribution of some reptiles and amphibians in southern Ecuador. Ann. Mag. Nat. Hist., (II) 2: 438-450. Pearson, Oliver P. 1954. Habits of the lizard Liolaemus multiformis multiformis at high altitudes in southern Peru. Copeia, 1954: 111-116. Templeton, J. R. 1967. Nasal salt gland excretion and adjustment to sodium loading in the lizard Ctenosaura pectinata. Copeia, 1967: 136-140. Wilson, E. O. 1957. Behavior of the Cuban lizard Chamaeleolis chamaeleontid.es (Dumeril and Bibron) in captivity. Copeia, 1957: 145. (Received 5 November 1968.) BREVIORA Museum of Cooipsirative Zoology Cambridge, Mass. 30 April, 1969 Number 317 RELATIONSHIPS OF TWO CRETACEOUS LIZARDS (SAURIA, TEIIDAE)1 Richard Estes Abstract. Peneteius aquilonius, n. gen., n. sp., from the late Cretaceous of Montana, is a small teiid lizard with complex cheek teeth. It most closely resembles the modern South American teiids Dicrodon and Teius and also shows resemblance to the large Cretaceous teiid Polyglyphanodon from Utah. Chamops segnis Marsh, from the late Cretaceous of Wyoming, re- sembles the modern teiid Callopistes mac u kit us. Meniscognathus altmani Estes, from the same locality as Chamops, may be related to the modern teiids Cnemidophorus and Kentropyx. These resemblances indicate that three distinct groups of primitive teiids living today in South America were in existence in North America during late Cretaceous time. INTRODUCTION In 1964 I described teiid lizards of modern type from the late Cretaceous Lance Formation of Wyoming. These included: (1) Chamops segnis, a species suggested to resemble the Recent Crocodilurus-Tupinamhis line in dental adaptations (see below), although it differed from all other so-called "macroteiids" (Van- zolini and Valencia, 1965) by having a parietal foramen; and (2) Meniscognathus altmani, which bore resemblances to the Ken- tropyx-Ameiva line in both dental apparatus and externally-con- cave mandibles. Two other forms, (3) Leptochamops denticulatus and (4) Haptosphenus placodon, were less clearly related to modern forms. Polyglyphanodon sternbergi, a large late Cretaceous lizard from Utah, was originally described by Gilmore (1940). In 1942 he placed it in a separate family Polyglyphanodontidae, but stated 1 Fossil vertebrates from the late Cretaceous Hell Creek Formation, Mon- tana: Contribution No. 2 (Contrib. No. 1 is Estes, 1965, Copeia, No. 1, pp. 90-95.) 2 breviora No. 317 that only tooth characters separated it from the Iguanidae. Hoff- stetter (1955) was the first to note the resemblance of Polygly- phanodon to the teiids and in 1962 suggested that it be referred to the Teiidae. A smaller, related genus Paraulxnhanodon was also named by Gilmore (1940, 1943). These animals are currently being restudied by Mr. William MacLean, 3rd. A vertebrate fauna recentl) recovered from the late Cretaceous IIJI Creek Formation, Montana, resembles that from the Lance Formation (Sloan and Van Valen, 1965; Estes, Berberian. and Meszoely, ms.). A single dentar) from the Hell Creek Formation sample belongs to a teiid lizard of unusual type and is described here. I interpret this fossil as in some ways intermediate between Polyglyphanodon and the modern genera Teius and Dicrodon, It is probably closer to the latter genera and provides an indication that a third major living "maeroteiid" line was already in ex- istence in late Cretaceous time in North America. ORDER SAURIA SUBORDER SCINCOMORPHA Family Teiidae Peneteius aquilonius, n. gen., n. sp. Holotype. MCZ (Museum o\' Comparative Zoology, Harvard University) 3612. fragmentary right dentar) with four complete teeth and the bases of four others (Fig. 1 ). Horizon and locality. West half section 9, T 22 N, R 43 E, McCone County, Montana; Hell Creek Formation. Collected by MCZ part) in 1964. Etymology. Latin, pent, almost; aquilonius, northern. Diagnosis. Differs from Recent Teius and Dicrodon in having tooth crests nearly transverse, but the lateral cusp anterior to medial cusp rather than posterior. Differs from fossil genera Paraglyphanodon and Polyglyphanodon in lacking transverse ex- pansion of tooth and in having external crests of principal cusp less well developed. Description. Jaw fragment relatively delicate; Meckelian groove wide, indicating a large splenial; bony separation between Mecke- lian canal and more lateral canal (for vascular and nervous struc- tures) set far medially, reducing depth of Meckelian groove. Teeth subacrodont, becoming molariform posteriorlv; most posterior (broken) tooth evidentlv the largest; tooth bases subcircular, relatively thin-walled; sulcus dentaUs absent. Tooth crowns antero- posteriorly compressed into crests; crests essentially transverse 1969 CRETACEOUS TEIID RELATIONSHIPS and formed by two main cusps, the labial one relatively the higher; both cusps connected by a transverse crest. Main cusps closer together and difference in height less pronounced in more anterior teeth, but cusp axis remains transverse; crests extending anteriorly and posteriorly from main cusps, forming slight basins on each side of crown; faint depressions present lingually on each side of main lingual cusp. Discussion. The widely-open Meckelian groove in combination with the unusual tooth crowns and heterodonty indicate relation- ship of Peneteius to the Teiidae. Closest resemblances within that family are to Recent Dicrodon and Teius from South America 0 1mm Fig. I. Peneteius aquilonius, n. gen., n. sp., MCZ 36 12; a, labial, b, lingual, c, occlusal view of fragmentary left dentary. KRIVIORA NO. 3 1 7 and the fossil Paraglyphanodon and Polyglyphanodon from the Cretaceous of Utah. Paraglyphanodon is the smaller of the two Eossil genera. Teeth oi the two described species /'. utahensis and P. gazini show a morphological series that could lead to the strongl) transverse, crested tooth condition seen in Polygtyphano- don, and it is possible that Paraglyphanodon is onl) a small in- dividual or the young of the former. The most complex and cuspate teeth in an\ Recent teiid occur in Teius teyou; Figure 2 shows occlusal views of teeth of pertinent living and fossil species. I he 1 Lg 2 Crown patterns of tend teeth. All arc posterior teeth of letl dentaries. ./. hur»di>n guttulatum, MCZ 111415; />. /). heterolepis, M( / 12329; c, Teius teyou cyanogaster, \l< Z J9982; d, I teyou, M( Z 43351b; e, Peneteius aquilonius, n. gen., n. sp.. MCZ 3612; f, Paraglyphanodon uta- hensis, United States National Museum 15668; i\ P. gazini, USNM 16580; h. Polyglyphanodon rternberghi, USNM 15477. a-g X about 16, h X about 9. two species of Dicrodon are quite distinct in tooth patterns and in man) other characteristics as well (Schmidt. 1957); Schmidt's species D. hohnbergi has been synonymized with P. guttulatum by Fugler ( 1967). The Recent species figured here all differ from Peneteius aquilonius in orientation of the two major cusps and in lacking a faint depression on the lingual side of the crown. The tooth pattern of Peneteius is distinct" from the Utah Cretaceous genera in lacking strongly-curved, prominent labial crests that may (Polyglyphanodon) or'may not (Paraglyphanodon) connect with the secondary cusp. Resemblances are shown to the modern genera 1969 CRETACEOUS TEIID RELATIONSHIPS 5 in the latter character, especially to Teius teyou and Dicrodon heterolepis. Additional similarities with the former are the ap- parently small number and relatively large size of the teeth. The basin-crest structure on tooth crowns of Recent species and Pene teius recalls anterior teeth of Paraglyphanodon (Gilmore, 1942, fig. 22), but the similarity is not great. However, resem- blance to the Utah Cretaceous forms is shown in the more trans- verse rather than oblique orientation of the tooth cusps. Restudy of the Utah fossil forms must precede further speculation on the affinities of Peneteius. THE RELATIONSHIPS OF CHAMOPS Chamops segnis Marsh (1892) is relatively common for a late Cretaceous lizard and is known from the Lance Formation (Wyom- ing), Hell Creek Formation (Montana), and Wapiti Formation (Alberta). Estes (1964) noted that Chamops was "quite probably ancestral to both Crocodilurus and Tupinambis.'" Comparison of Chamops and Callopistes (a genus not available to me in 1964) requires some modification of that statement. Callopistes maculatus (MCZ 2751) is close to Chamops in several ways, principally in the relatively deep shape of the maxilla (Fig. 3c,d), the more normal (less conch-like, or curved) quadrate shape, and relatively elongate parietal (Estes, 1964, rig. 49). The parietals and quadrates are referred to Chamops on the basis of both size and frequency, as well as on their generally teiid appearance; Chamops is the largest and most common teiid in the Lance Formation. The tooth row of Callopistes maculatus is more heterodont than that of Chamops and fewer teeth are tricuspid; I interpret both heterodonty and bicuspid teeth as specialized features. How- ever, the nasal in Callopistes maculatus extends further forward on the maxilla than in Tupinambis and is thus more like the con- dition in Chamops (Estes, 1964: 107). The latter, Tupinambis, and C. maculatus share a pointed lateral premaxillary process of the maxilla (Fig. 3). The dentary of Chamops is relatively deeper than that of Tupinambis of equal size and resembles the propor- tions seen in C. maculatus. Tooth number is essentially the same in all these genera, contrary to my statement in 1964 (p. 107), which was based on only a few individuals. Facial elongation is characteristic of many "macroteiids" and is most extreme in Cnemidophorus. The latter does not differ from Ameiva in facial elongation, although I so stated in 1964 (p. 108); examination of a large series shows considerable size variation in BRLYIORA No. 317 Fig. 3. a. Crocodilurtu lacertinus, American Museum of Natural His- tory 46290; />. Tupinambu nigropunctatus, Los Angeles County Museum R-74; c, Cluunops segnis. University of California Museum of Paleontology 46033, restored dorsally from UCMP 46094 and other specimens; d, Cal- lopistes maculatus, MCZ 2751; not to scale, all reduced to a common length. 1969 CRETACEOUS TEIID RELATIONSHIPS 7 this character and the two genera are probably synonymous, as indicated by recent studies (Gorman, pers. comm. and ms. 1968). Tupinambis nigropunctatus and Callopistes maculatus of equal size show the former slightly exceeding the latter in facial length; Crocodilurus resembles the latter. Kentropyx calcaratus resembles Cnemidophorus in this feature; my statement to the contrary in 1964 (p. 108) was based on a misidentified skeleton. In summary, the Recent Callopistes maculatus appears to be the closest relative of Chamops segnis. Tupinambis nigropunctatus is the most primitive member of that genus and is close to Chamops but appears more advanced than the latter and Callopistes in a number of features. The maxilla of Crocodilurus is relatively less high than that of Chamops and the former seems to be less closely related to the latter than it is to Tupinambis. CONCLUSIONS Current study of "macroteiids" by Gorman, Presch, MacLean, and myself is in general agreement with Vanzolini and Valencia (1965) in separating two major subgroups: one including Callo- pistes, Tupinambis, Crocodilurus, and Dracaena; the other formed of Ameiva, Cnemidophorus, Kentropyx, Teius, and Dicrodon. The latter two genera possess distinctive, crested, cuspate cheek teeth. Peneteius aquilonius, n. gen., n. sp., from the late Cretaceous of Montana, has similar teeth and is probably related to the Recent Teius-Dicrodon line. Chamops segnis, from the late Cretaceous of Wyoming, Montana, and Alberta, appears to be related to the Recent species Callopistes maculatus. The latter two species are probably more primitive, on the basis of high maxilla and less well-developed heterodonty, than are Tupinambis, Crocodilurus, or Dracaena. With the possible relationship noted above of the fossil genus Meniscognathus to the Ameiva-Kentropyx group, it thus appears that three distinct groups of "macroteiids" living today in South America were in existence in North America during late Cretaceous time. ACKNOWLEDGMENTS I am grateful to Mr. William P. MacLean, 3rd., Mr. William Presch, and Drs. Arthur C. Echternacht and George Gorman for comments. Figures 1 and 2 are by Mr. Laszlo Meszoly. This re- search was supported in part by National Science Foundation Grant GB-7176. 8 breviora No. 317 REFERENCES CITED ESTFS, R. 1964. Fossil vertebrates from the late Cretaceous Lance Formation. eastern Wyoming. Univ. Calif. Publ. Geol. Sci . 49: 1-180. FUGLER. C. 1967. Geographic variation in Dicrodon guttulatum Dumeril and Bibron of the Ecuadorian and Peruvian littoral. Journ. Alabama Acad Sci . :i«: 322. Gil MORI . ( 1940. New fossil lizards from the Upper Cretaceous of Utah. Smith- sonian Misc. ( oil.. 99: 1-3. 1942. Osteology oi Polyglyphanodon, an Upper Cretaceous lizard from l tan. Proc. I S Natl, Mus., H<»: 11-26. 1943. Osteolog] Of Upper Cretaceous lizards from Utah, with a description of a new species. Proc. U. S. Natl. Mus., 93: 2<>l>- 214 Hoi i mi i ii r. R \i><* Squamates de i\pe moderne. /". J. Piveteau, ed.. Traiic; de Paleontologie (Paris, Masson ci ( ie.), •>: 606-662. 19h2 Revue des recentes aquisitions concernanl l*histoire el la sWcmatiquc des squamates. Colloq. Internat.. Prob. Pal. no. 104: 243-2 M \ksh. O. 1892. Notice of new reptiles from the I aramie Formation. Amcr. Journ. Sci., (3) 13: 449-453. S( HMII>I . k 1 1>57. Notes on lizards of the genus Dicrodon. Fieldiana: Zool., 39: 65-71. Sloan, R . \m> I Van Vai i N 1965. Cretaceous mammals from Montana. Science. 118: 220-227. Vanzolini. P., and J. Vali n< i \ 1965. The genus Dracaena, uith a brief consideration of macroteiid relationships (Sauria. leiidae). Arq. Zool.. 13: 7-35. (Received 5 February 1969.) BREVIORA Mmsemnti of Comparative Zoology Cambridge, Mass. 30 April, 1969 Number 318 LEUCOLEPIDOPA SUNDA GEN. NOV., SP. NOV. (DECAPODA: ALBUNEIDAE), A NEW INDO-PACIFIC SAND CRAB Ian E. Eflford1 Abstract. Leucolepidopa gen. nov. is described along with the only known species in the genus, Leucolepidopa sunda sp. nov. The single specimen is from the Sunda Strait between Sumatra and Java. The posi- tion of this new genus is discussed and the relationships of all the genera within the family Albuneidae are reviewed. Leucolepidopa is most closely related to Austrolepidopa and Lepidopa. The Danish Kei Island Expedition collected a single specimen of a new sand crab during dredging operations in the Sunda Strait between Sumatra and Java. This new crab is from an undescribed genus closely related to Austrolepidopa, recently described from Australia (Efford and Haig, 1968), and to an entirely American genus, Lepidopa (Efford, in ms.). In this paper I describe the new species and present a preliminary outline of the probable evolutionary relationships within the family so that the position of the new genus can be seen. Leucolepidopa gen. nov. The type and only known species of this genus is Leucolepidopa sunda. This new genus is in the family Albuneidae and is closely related to Austrolepidopa and Lepidopa. It can be distinguished from other genera in the family by ( 1 ) the antennae having three articles in the flagellum and (2) the eyeplates possessing long setae on the dorsal surface. The following characters are not specific to the genus but can be used to separate it from others within the family: (3) a pleural expansion is present on the fifth abdominal segment; (4) the lateral spine is on the carapace and above the linea anomurica; (5) the scaphocerite is short, only just 1 Department of Zoology, University of British Columbia, Vancouver 8, B. C, Canada. 2 BREVIOR \ No. 3 IS extending up to the base of the fourth antenna! segment; (6) the rostrum is smoothly convex and lacks a subrostral spine; (7) there are no terminal spines on the anterolateral lobes; and (N) the anterolateral lobes cam six or seven teeth. The generic name refers to the whiteness of the carapace and to the close relationship oi this genus with the genera Lepidopa and Austrolepidopa. 1 I l ( (H I 1MDOPA SUNDA SP. NOV. Holotype. A male. 7mm carapace length and 8mm carapace width (Fig. 1. 1-8, and Fig. :. 1-6). Collected on 29 July 1922 from 75° 6' 10" S. 105° 44' E in the Sunda Strait between Sumatra and Java, bv the Danish Kei Island Expedition. The specimen was collected b\ dredging from a sand-shell bottom in 40 m o\ water. Deposited m the /oologisk Museum. Koben- havn. I he specific name sunda is derived from the name o\' the geo- graphical location. Description. The carapace has a rostrum that is gently rounded and lacks spines or teeth. It is Hanked on either side b\ a shallow. gently curved ocular sinus and by rounded anterolateral lobes, the edges o\ which cam six or seven teeth. From the antero- lateral lobes the anterior edge oi the carapace curves backwards and outwards in a smooth sigmoid curve to the large, distinct, lateral spines. I here are no spines along the anterior edge of the carapace as there are in most other members of the family. I lie edge is lined with long, branched setae, which gradually be- come shorter toward the lateral spines. From the lateral spines the carapace widens sligluK back to the midline, behind which the edges arc straight and converge. The posterior end of the carapace is smoothly rounded, with a deeply rounded medial con- cavity. The carapace has an extensive pattern of setae. The basic pattern is similar to that of both Austrolepidopa and Lepidopa; the main difference is that the M-shaped pattern across the an- terior end of the carapace extends up to the anterior edge and some way back along the sides. There is a smooth area in the middle of this pattern on either side. The setae in this area are longer than those on the posterior part of the carapace. Those lining the lateral spine groove are very long and branched, and extend beyond the anterior edge of the carapace. The posterior half of the carapace is almost equally setose, but here the setae 1969 NEW INDO-PACIFIC SAND CRAB 3 are in small groups rather than in one continuous expanse. In contrast, the posterior edge of the carapace, round the cavity, is free of setae and very smooth. The groove running down the side of the carapace stops before it reaches the posterior edge and does not follow the edge round as in Lepidopa. The carapace is chalky white with some iridescence. The first abdominal somite is shorter than wide, and the exposed posterior part forms a raised crescent-shaped area lined along its anterior edge by a dense row of setae. The second somite has broad pleural expansions with slightly concave anterior edges. The edges are lined with long setae that all face forward. Those along the lateral and posterior edges arise from a groove that runs round parallel with the edge. There are no setae along the edge of the central part of the plate. The third, fourth, and fifth seg- ments are also winged, but unlike those of Lepidopa, all the pleural expansions are rather straight, narrowing towards the ends, and both the anterior and posterior edges of the expansions have grooves lined with forward-projecting setae. The setae near the outside tips of the pleural expansions of the fifth segment are particularly long and, in this case, project laterally, away from the abdomen. One noticeable character is the presence of distinct bumps along the posterior edge of the fourth pleural expansion. The sixth segment is shorter than wide and narrows towards the anterior edge. It has a distinct waist, and the posterior edge is lined with backward-projecting setae. The telson is broad and pear-shaped. The eye-plate is quite distinct, as the dorsal surface is covered with long; setae. In Austrolepidopa and Lepidopa the upper sur- face of the plate is smooth, although there may be setae around the edges. The plate is broadest about one third from the proximal end. The inner edge is fairly straight and the outer edge convex. From the broadest point the eyes narrow towards the rounded distal end. The edge of the plate is stepped, and long setae are attached at each step. No cornea or eye-spot is visible. The antennules have a broad basal segment. The second seg- ment is broad, partiallv compressed, and has a large, right-angled bulge on one side. The third segment is about as long as the second and widens towards the distal end. The two first antennule flagella of the holotype have been broken off so that their length is unknown; however, the lower part of the first flagellum sug- gests that in this species they are very long, as in Lepidopa. No second flagellum is present on the antennule. 4 breviora No. 318 The antennae are unusual because the setae are arranged terminally or along the inner side of the segments and they are not found scattered around on other parts as they are in other genera. This gives the antennae a rather bare appearance. The scaphocerite is very small and hardly overlaps the base of the fourth segment. It has a group oi very long setae arising from the outer surface. The fifth segment is shorter than the fourth; it is cylindrical and terminates in a tlagellum oi three long cylindri- cal articles. The last article is longer than the other two. All three articles have areas of vet) long setae on their distal ends. In the holotype the setae are broken oil' the terminal article. I iie mandible is strong, smooth!) shaped, and carries a three- merited palp. There is a \er\ sharp cutting edge, two distinct teeth at the distal end. and three similar teeth at the proximal end. The maxillula was not examined. The maxilla is similar to those of others in the family, although I did not examine the scaphognathite. The firs! maxilliped is Battened and leaf-like; the endopod consists of two distinct segments. The second maxil- liped is similar to others found in the family; the exopod is single and well developed The third maxilliped has a very well- developed expansion o\' the antcrodoisal aid o^i the carpus, which extends over the propodus and reaches the junction oi the pro- podus with the dactylus. The dactylus is over half as long as the propodus. The exopod is a single, long, cylindrical segment. 1 he pereiopods resemble those found in other members o\' the family. The first is subchelate with a sickle-shaped dactylus. The s.des oi the dactylus are smooth, and the upper surface is covered with long setae that point towards the proximal end of the seg- ment. The propodus has a distinct cutting edge lined with very short, blunt setae and terminating in a large, hooked tooth. The side is covered with rows of long setae. 1 he second and fourth pereiopods have broad, blunt, sickle-shaped dactyla. The third pereiopod has a long, smoothlv-curved dactylus without any dis- tinct basal projection; the carpus is very much expanded at the anterior end. with the anterior half of the upper surface covered with a dense mat of setae. The fifth pereiopod is long and chelate. The genital pore is round. Relationships. To examine the relationships of Leucolepidopa with the other members of the family, I used characters that seem to vary little between species within a genus, although they do vary between genera. These were: (1) the number of articles in the flagellum of the antenna; (2) the presence or absence of pleural expansions on the fifth abdominal segment; (3) the 1969 NEW INDO-PACIFIC SAND CRAB 5 presence of the lateral spine above or below the linea anomurica; (4) the length of the scaphocerite; and (5) the shape of the rostral region of the carapace. When the genera are examined using these characters, it becomes clear that some of the charac- ters always occur together. The presence of a pleural expansion on the fifth abdominal somite is always associated with a short scaphocerite and with a rostrum that is quite clearly convex in shape. The opposite three characters are associated with the absence of a pleural expansion on the fifth abdominal somite. These two groups can be used as the basis for separating the family into its evolutionary groups. I should mention here that Blepharipoda and Lophomastix are so distinct from the other genera in the family Albuneidae that they form a side branch off the main stream, and were not included in the analysis. The family can be divided into three groups (Fig. 3). The first, the Lepidopa group, contains Lepidopa, Austrolepidopa, and Leucolepidopa. Leucolepidopa is most clearly related to Austro- lepidopa, as shown by the similar shape of the eye-plates and by the stiff, straight, pleural expansions on the fifth abdominal somite compared to the rather delicate curved ones in Lepidopa. They differ in the length of the antennal flagellum, the presence of setae on the dorsal side of the eye-plate, the absence of a second flagellum to the antenna, and the setal pattern on the carapace. It is assumed that Leucolepidopa is more specialized than Austro- lepidopa, as it has only three articles in the flagellum, whereas both of the other genera have seven or eight. Austrolepidopa is closely related to Lepidopa, as is shown by the similar lamellated eye-plates, the antennal flagellum of seven or eight articles, as well as the elongated carpus of the third maxilliped, short scapho- cerite, pleural expansion, and convex rostrum common to the three species. A second group, the Albunea group, has only two species, Albunea and Stemenopa. The former genus occurs throughout the tropical regions and is rather generalized in form. Stemenopa, recorded from only one location in Western Australia, is a very specialized form with extremely long eye-stalks. It closely relates to Albunea, as is shown by the absence of a pleural expansion on the fifth segment, the concave rostrum, and the long scaphocerite. Zygopa is specialized in having both fused eye-plates and a single article in the antennal flagellum; it occupies a place some- where between the other two groups. According to Holthuis (1960), it is more closely related to the Lepidopa group because of the position of the lateral tooth. Possibly more weight should 6 bri-viora No. 318 be put on its similarity to the Albunea group in its long scapho- cerite, concave rostrum, and pleural expansion of the fifth ab- dominal segment, rather than its similarity to the Lepidopa group in having the lateral spine above the linea anomuriea. Until we have more information, we must assume that a closer relationship is suggested by three characters held in common than by only one. which places Zygopa closer to the Albunea group than to the Lepidopa group. The final group in the family is the rather separate off-shoot of Blepharipoda and Lophomastix. Apart from the position of Zygopa, the genera fall rather easily into these separate groups. These relationships could be tested, and more information obtained to confirm or reject the tentative position of Zygopa, b\ comparing the larvae. At the present time. however, we know the larvae only of Lepidopa and Blepharipoda, so that we are still some \\a\ from understanding the evolution of the family Albuneidae. ACKNOWLEDGMENTS I would like to thank Dr. I orben Wolff for allowing me to describe this specimen and Dr. Ernst Mayr, Dr. Herbert Levi, and Miss Alice Bliss for their help during my stay at the Museum ol Comparative Zoology. LITERATURE CITED Ej-kord, 1. E. (in ms. i. The species of sand crabs in the genus Lepidopa (Decapoda: Albuneidae i. I I I < »KI». I. E., AND J. HAIG 1968. Two new genera and three new species of sand crabs (Decapo- da. Anomura. Albuneidae) from Australia. Aust. J. Zool., 16: 897-914 HOLTHl IS. L. B. I960. Notes on American Albuneidae (Crustacea, Decapoda, Ano- mura) with the description of a new genus and species. Proc. K. Nederiand Akad. Wet., Ser. C, 64 (1): 21-36. (Received 5 November 1968.) 1969 NEW INDO-PACIFIC SAND CRAB Fig. 1. Holotype of Leucolepidopa sunda gen. nov., sp. nov., male cl. 7 mm. 1. Dorsal view of the eye-plates and carapace with the setae pat- tern shown. 2. Dorsal view of the abdomen and telson. 3. Left first pereio- pod. 4. Right third pereiopod. 5. Left fourth pereiopod. 6. Left fifth pereiopod showing round genital pore. 7. Left second pereiopod. Scale 1 mm, except in 1 and 2, where it is 0.5 cm. 8 BREVIORA NO. 3 1 S Fig. 2. Holotvpe of Leucolepidopa sunda gen. now. sp. nov, 1. Base of antennule. 2. Dorsal view of left eye-plate. 3. Left antenna. 4. Left second maxilliped. 5. Left third maxilliped. 6. Left first maxilliped. 7. Left maxilla with the scaphognathite missing. S. Left mandible. Scale 1 mm. 1969 NEW INDO-PACIFIC SAND CRAB LEUCOLEPIDOPA (31 5TEM0N0PA 171 ZYGOPA III SPINE B E 1 0 W LINEA ANOMURICA SPINE ABOVE LINEA ANOMURICA SPINE ABOVE LINEA ANOMURICA NO PLEURAL EXPANSIONS ON 5IH ABDOMINAL SEGMENT. ROSTRUM CONCAVE. SCAPHOCERITE LONC. PLEURAL EXPANSION ON 5TH ABDOMINAL SEGMENT. ROSTRUM CONVEX. SCAPHOCERITE SHORT. BIEPHARIPODA GROUP BLEPHARIPODA IOPHOMASTIX ANCESTRAL ALBUNEIDAE FAMILV HIPPIDAE HIPPA EMERITA MASTIGOCHEIRUS ANCESTRAL HIPPOIDEA Fig. 3. Evolutionary relationships within the family Albuneidae. Num- ber of articles on the antennal flagellum is given for each genus. BREVIORA Museumi of Comparative Zoology Cambridge, Mass. 30 April, 1969 Number 319 COMPETITIVE EXCLUSION AMONG ANOLES (SAURIA: IGUANIDAE ON SMALL ISLANDS IN THE WEST INDIES' A. Stanley Rand2 Abstract. Each of the Greater Antilles has been a largely independent center of anole radiation and there has been little exchange of species between them. Together they are the source areas from which the other islands of the Antilles have been colonized. The area and number of anole species known from 122 of these smaller islands on 44 banks are tabulated. There are two kinds of small islands in the West Indies. The islands of one class show a correlation between area and number of anole species, and the larger islands may have up to 4 or 5 species. Islands of the other class do not show an area effect and never have more than 1 or 2 species. The first class includes the islands of the Great Bahama Bank and the islands fringing the Greater Antilles and lying on the same bank as one or another of them. The second class includes the Lesser Antilles and the small isolated islands of the Caribbean. The difference between the two classes seems to be a result of their Pleisto- cene history and the way in which they received their anole faunas. The islands of the first class were all connected or very close to a source area during the Pleistocene sea level minima and so probably received their anoles overland and more or less at the same time. The area effect now seen was probably produced by differential extinction subsequent to a sea level rise and separation of the various islands. The islands of the second class must all have been colonized over an appreciable water gap. The absence of islands with more than 1 or 2 species in this second class cannot be due to ecology, size, distance, or species of anoles involved, since some of the islands are large, diverse, and no further from a suitable source than some fringing islands are today. The data suggest that a first species finds it relatively easy to colonize an island. A second species finds it difficult, and a third may find it impossible, even though three species may coexist on a very small and ecologically impoverished island if they have colonized it overland. The exclusion effect occurs not because of the dif- ficulties in reaching the island, but because of the difficulties in establish- ment. The latter are the result of inter-specific competition in which an 1 Research supported in part by NSF Grant GB-6944. - Smithsonian Tropical Research Institute, Balboa, Canal Zone. 2 breviora No. 319 alread) established species can exclude an invader even though if both were established together the) migl I coexist, it is suggested that competitive exclusion ma) also explain the small number of exchanges of species among the Greater Antilles. This paper deals with the factors influencing the number of species of Anolis per small island in the West Indies. Darlington (1957) relates fauna! size and complexity in the West Indies to island area and to distance from the postulated source area o\ Central America. However true these generalizations are for the fauna as a whole, they must he modified for the genus Anolis: first, because the West Indies have been a center o\' speciation and have not received their modern fauna directly from Central America and, second, because there is a class o\' small West Indian islands that have only one or two species i^\' Anolis regard- less of area or distance from source islands. The following dis- cussion o\' the distribution of a single genus in one area is an ap- proach to the theoretical problems o\' the zoogeography of archi- pelagos that I hope is complementary to the more mathematical treatments o\' Hamilton. Barth. and Rubinoff ( 1964), MacArthur and Wilson ( 1967). and others. I here are approximately 3.000 islands in the West Indies. Many of the islands stand on more or less extensive banks and are sepa- rated from one another or from one o\' the Greater Antilles by onl) shallow water. When the sea level was lowered during the last Pleistocene Emaciation, the banks must have largely been exposed. At that time the islands on each bank must have been connected to one another by dry land, so that an exchange of land faunas was possible. The anole faunas o{ the islands on each bank arc very similar to one another. The water between the banks is ver) deep, and there is no possibility that islands on different banks were connected during the Pleistocene. Duruig the interglacial period that preceded the last glaciation. the sea level was higher than it is at present, and the maximum probably was high enough to submerge completely many of the low-lying West Indian islands and exterminate any anoles living there. Anolis is represented in the West Indies by about 100 species, more species than are known for any other genus of land verte- brates in the area. These species may be the results of as few as two overwater invasions, one from Central America into Jamaica or Cuba and the other from South America into Puerto Rico or perhaps Hispaniola. Here and below the conclusions of Etheridge (I960) and Williams (personal communication) on the relationships within the genus Anolis have been followed. 1969 COMPETITIVE EXCLUSION 3 Each of the four Greater Antilles has been a largely independent center of evolution for anoles. Cuba has 23 species of Anolis, Hispaniola 21 species, Jamaica 7 species, and Puerto Rico 10 species. Despite the fact that several of these species have crossed large water gaps to live on distant small islands, only one species is shared by two of the large islands. Even in the more distant past, the number of colonizations between the Greater Antilles was small. Only eight such events are necessary to explain the relationships between living forms: four between Puerto Rico and Hispaniola; two between Hispaniola and Cuba; two between Cuba and Jamaica. On the four large islands, speciation has been largely within and not between islands. The Greater Antilles, with their older and richer anole faunas, are probably the source areas from which the smaller (some only recently habitable) West Indian islands have been colonized. An anole invading one of the Greater Antilles from a small island is probably even rarer than the invasion of one of the Greater Antilles by an anole from another. Anoles have repeatedly crossed water gaps. They have not only reached and spread throughout the West Indies, but one West Indian species has reached the coast of Central America and another the Bay Islands of Honduras. It has been suggested that anoles may be blown from island to island in hurricanes or float across in rafts of vegetation carried by ocean currents. Per- haps they travel in the masses of debris and vegetation that are torn loose from a hurricane-lashed shore and blown out to sea. The heavy rains that accompany hurricanes would reduce the danger from salt water during the journey. On the assumption that the large islands have been the source areas, most but not all colonizations have been from east to west, which is the general direction of both ocean currents and prevailing winds, as well as the storm track of many hurricanes (see Fig. 1 ). Data on area and number of Anolis present have been collected for 126 small islands on 44 banks. Anole distributions from Barbour (1937), Cochran (1934, 1941), Grant (1932a, 1932b, 1936, 1937), Grant and Roosevelt (1932), Oliver (1948), Ruibal (1964), Underwood and Williams (1959) have been supple- mented by the collections in the Museum of Comparative Zoology and unpublished information from the files of their curator. Dr. Ernest E. Williams. Mr. J. D. Lazell, Jr., and Dr. Albert Schwartz generously shared the results of their extensive field experience. Thanks are due to all three. Information on island areas was taken from the Columbia Lippincott Gazetteer of the World and imi \ loRA No. 3 1 9 1969 COMPETITIVE EXCLUSION 5 U. S. Hydrographic Office charts. The data are given in Table 1 . Almost all the major islands are included; the number of smaller ones could be increased greatly, but there is no suggestion that this would change the picture. The smaller islands of the West Indies fall into four geographical categories: (1) the islands fringing the Greater Antilles; (2) the Lesser Antilles; (3) the isolated islands of the Antilles; and (4) the islands of the Bahamas. Each of these groups shows a some- what different pattern of anole distribution. ( 1 ) The fringing islands of the Greater Antilles stand on a bank with one or another of the four Greater Antilles and are separated from it by only shallow water. Most are close to the associated large island, but some of the Virgin Islands are almost 100 miles from Puerto Rico. These islands vary ecologically from very simple to moderately complex. I have data on 44 islands in this group, all inhabited by anoles. The fringing islands show a definite area effect, i.e., size of island and number of species are directly correlated (Fig. 2). Many are low, and most must have been submerged during the last Pleistocene sea level maximum. During the last Pleistocene sea level minimum, these islands must have become hills on the coastal plains of the Greater Antilles and been inhabited by the common lowland species of Anolis. After the sea level rose, the larger islands (more than 25 square miles) kept their original three to five species, but on the smaller islands there appears to have been a tendency for species to become extinct. The number that remains is related in part to island area, and the very smallest islands (less than 1 square mile) frequently now have only a single species left. On none of these islands has there been any apprecia- ble differentiation except for the endemic, A. roosevelti, on Culebra. (2) The Lesser Antilles extend in a 500 mile arc from just east of the Virgin Islands south almost to Trinidad and Tobago. Many reach altitudes of several thousand feet and show a variety of climatically different habitats with rich floras. I have data on 33 islands on 16 different banks, all inhabited by anoles. Comparing these islands among themselves there is no apparent area effect. Five of the banks have islands with two species, the islands on the remaining 1 1 banks have only a single species (excluding certain introductions by man). The Lesser Antilles have been colonized by two different stocks. One, the bimaculatus species group, including wattsi, has spread throughout the Northern Lesser Antilles, crossing at least nine water gaps. It probably 6 BRLVIORA No. 319 came from the Greater Antilles, though the ancestral species has either disappeared or changed so much that it is unrecognizable. The stock has differentiated on ever) island, and taxonomists differ on how many populations have reached the species level. On several islands these anoles show marked geographical varia- tion within the island ( l.a/cll. 19d2. 1964). Though the bimacula- tus group has been in the Lesser Antilles a long time, there seem to have been very lew cases of double invasions. On only the three northernmost banks o\o two species (one of them wattsi) occur together. The second stock, the roquet group, has invaded the Lesser Antilles from the south, crossing at least seven water gaps, but it is not known to be closel) related to an) living South American species. There are two banks on which two species of this group coexist. The degree of differentiation between islands is less in this group than in the bimaculatm group. (3) The isolated islands are those islands or small groups of islands scattered throughout the West Indies and separated from all other islands b\ \cr\ deep water. Most are low and quite simple ecologically. Some arc \cr\ far from any source area. Others are quite close. I have data on I I islands, on 9 banks, all inhabited In anoles. ( I here are also a vcr\ lew. very small. isolated cays [e.g. Pedro Cays, Aves Island] that are uninhabited; the) are the onl) islands in the West Indies known to lack anoles.) These islands show no area effect. One island has two species ol anoles. the others a single species. Most o[ these islands were probabl) submerged during the Pleistocene sea level maximum and colonized overwater since their emergence. All of these islands are occupied b) species that are coiispecilic with or clearly derived from a species now living on one of the Greater Antilles. The one exception is acutus on St. ( roix, probabl) derived from a Puerto Rican stock but not from one of the living species. (4) "The Bahamas are a large group of islands stretching for some six hundred miles in an elongate cluster from Florida, along the northeast coast oi Cuba, almost to Hispaniola. They are all low islands with little ecological diversity and were probably all submerged during the last Pleistocene sea level maximum. The islands lie on a number of banks o\ various sizes. I have data on 53 islands on 15 banks, all inhabited by anoles. The largest bank is the Great Bahama Bank. During the Pleistocene sea level minimum, its many islands must all have been part of a single very large island lying next to Cuba. The present islands show an area effect very like that of the fringing 1969 COMPETITIVE EXCLUSION 7 islands. The larger islands on this bank are inhabited by four species of anoles, three from Cuba (sagrei, carolinensis , and angusticeps) and one from Hispaniola (distichus). These species must have reached the bank overwater and then probably spread through it overland. The remaining Bahamas lie on a number of smaller banks. Of the 14 banks for which I have data, the islands on 10 are occupied by a single species and on 4 by two species. Many of these islands are much larger and at least as varied ecologically as some of those on the Great Bahama Bank that are occupied by four species of anoles. The Bahamas on other banks that lie close to the Great Bahama Bank are occupied by one or two of the anoles that occur on the Great Bahama Bank. The Bahamas farther southeast are oc- cupied by a single endemic species, scriptus. The ancestors of scriptus may have reached the Bahamas in a single long overseas crossing from Puerto Rico, where its very close relative, cristatellus , occurs, though it may have come from a now extinct form on Hispaniola. A. scriptus does not occur with any other anole, though it lives on West Plana Cay only ten miles from East Plana Cay, which is occupied by A. carolinensis. From this survey it appears that, on the basis of the size of their anole faunas, there are two kinds of small islands in the West Indies. The first class of island includes the fringing islands of the Greater Antilles and the islands of the Great Bahama Bank. These clearly show an area effect and only the smallest have a single-species fauna (Fig. 2, upper). The second class of island includes all of the Bahamas off the Great Bahama Bank, the Lesser Antilles, and the isolated islands (Fig. 2, lower). These show little if any area effect, and even the largest islands have only one or two species. The difference in numbers of anoles between these two types of islands cannot be explained on the basis of differences in their ecological diversity or carrying capacity. The Lesser Antilles, with only one anole species, are larger, much richer, and more varied ecologically than are many fringing islands with three or four species of anoles. Within the Bahamas there seems to be no dif- ference in the ecology between the islands on the Great Bahama Bank, where four species coexist, and those on other banks, where only one or two species occur together. The difference cannot be in behavioral or ecological peculiarities of the species involved, for the species that replace one another in some of the Bahamas occur 8 BRE\ IORA No. 319 ♦ ♦ I ♦ i ♦ o o : ♦ • : T3 C co c •5 c 3 -a . oc co c CO — £ E m ea u c co x 5= 2 k -a o c CO u — c ed «- CO « O -7 a O .SJ _£ _ u u - C O .2 £ 1-1 I ° £: X. II c ■ M -3 "" C C H CO CO S o c ill - \j Z. - —, to ~ IS! - " ( r) O X. E E 3 < E CO x: - QC —> CO u u. u DO •j: ^^ ■-> -3 O s I- o m t ro (M S3ID3dS JO H38WflN 1969 COMPETITIVE EXCLUSION I _o l_ l_ I < CD O ^°~~«.E°y *^i a 3 /*, R SS g a I « ' 5 .E ^ Q.T3 ~ .E E ~ t? •5 -3 M -E ~ - ^E£g.iE »5 B «-i in •— *- 0 _C Q t4~l *o '35 ca c d "2 *• ' p u x: o ^ E .§ | 8 8 * -8 8 ^m § s s J s - "rt-cec.sSuE «w B o 2 „. > o I- X o "O "o ^ -° oa - b o cu2 y S E ox: 5T ^ 5 B 2 £ u g ° £ g «B w y 09 c "B ■« ESE I 8 8 £ Jg o £ E u « «2 * „ o, « „> y . .a e c p 1 0.25 Dead Man's ( hest 1 0.25 Sail Ke\ 1 0.25 o Santiago 3 0.125 1 0.125 1 esser Antilles: 33 islands on 16 banks Vnguilla 2 35.0 St. Martin 2 35.0 St Bartheln 1 9 s Vntigua 2 108.0 Barbuda 2 62.0 Saba 1 5.0 St. Eustatius 2 7.7 St. Kitts 2 68.0 Nevis 2 50.0 Redonda 1 0.5 Montscrrat I 38.0 1969 COMPETITIVE EXCLUSION 15 Island (those indented under another lie on the same bank) Desirade Guadeloupe Les Saintes, Terre de Bas Terre de Haut Marie-Galante Dominica Martinique St. Lucia Barbados St. Vincent Grenada Quatres Carriacou Mustique Petit Mustique Petit Nevis Baliceaux Petit Martinique Frigate Ronde Caille Diamond Isolated islands; 1 1 islands on 9 banks. St. Croix Buck Grand Cayman Mona Cayman Brae Little Cayman Navassa Swan (Larger) Swan (Smaller) Desecheo Alta Vela Bahamas; 53 islands on 15 banks. Great Bahama Bank (15 islands) Andros Eleuthera Cat Island Long Id Great Exuma Mangrove Cay New Providence Little San Salvador Stocking Id Number of Island area A nolis species in sq. mi. 1 10.5 1 687.0 1 3.0 1 2.5 1 58.0 1 305.0 1 427.0 1 233.0 (2 introduced) 1 166.0 2 133.0 2 120.0 1 0.5 2 8.0 1 2.5 1 0.125 1 0.125 1 0.75 1 1.0 1 0.25 1 1.0 1 0.25 1 0.125 1 82.0 1 0.75 1 71.0 1 21.0 1 13.0 2 9.0 1 1.5 1 1.0 1 0.5 1 0.75 1 0.5 4 500.0 4 164.0 4 160.0 4 130.0 4 108.0 3 85.0 4 58.0 1 4.25 1 4.0 16 » BRFVIORA NO. 319 Island (those indented under Number of Island area another lie on the same bank ) Anal is species in sq. mi. South Bimini 4 3.0 North Cat Caj : 0.75 Margaret Island, Ragged Island Group 0.75 Flamingo Cay 0.5 Knife Cay 0.062 ( io.it Caj 0.062 \cklins 185.0 Crooked Id 76.0 Long Cay 7.0 Castle Kland 1.5 Cotton Caj 0.5 Atwood Caj 10.0 ( Irand ( aicos 40.0 North Caicos 40.0 Providenciales 30.0 1 ast c aicos 300 so ith ( aicos 8.0 West ( laicos 7.0 Pine Caj 1.0 < otton c aj 2.0 Elbow i 1.0 Conception 4.0 1 ast Plana ( B) 3.0 Great Inagua 560.0 Sheep ( a) 0.25 1 ittle Baham i Bank Gr. Ahaco hi 776.0 1 i and Bahama 430.0 1 ittle Ahaco 27.0 Elbow 1 2.0 Pcnsacola ( B 1.0 Water ( a\ . Grand Bahama 1.0 Strangers ( aj 0.75 Moraine Caj 0.062 I ittle Inagua 45.0 Mavaguana Id 96.0 Booby Caj 0.25 Rum Cay 2 29.0 Grand Turk 8.0 Salt Ca\ 2.5 Cotton Ca\ 0.5 I .one Cay 0.25 Great Sand Cay 0.25 Watlings Id 2 60.0 West Plana Cav 1 3.5 BREVIORA Museum of Comparative Zoology Cambridge, Mass. 10 June, 1969 Number 320 TAIMANAWA, A NEW GENUS OF BRISSID ECHINOIDS FROM THE TERTIARY AND RECENT INDO-WEST-PACIFIC WITH A REVIEW OF THE RELATED GENERA BRISSOPATAGUS AND GILLECHINUS Robert A. Henderson and H. Barraclough Fell Abstract. Taimanawa gen. nov. is erected to include two New Zealand fossil species, T. pulchella sp. nov. (taken as the type species) of early Miocene age and T. greyi (Hutton) of Oligocene age, as well as a poorly known extant species from the Kei Islands, Indonesia, T. mortenseni sp. nov., which had hitherto been tentatively referred to Plagiobrissus Pomel. In spite of possessing an internal fasciole, Taimanawa is referred to the Bris- sidae, with which it agrees in other morphological features. The closely related genera Gillechinus Fell and Brissopatagus Cotteau are re-evaluated using type material of the type species; they are maintained as distinct, but most of the species previously referred to Brissopatagus are here transferred to Gillechinus. INTRODUCTION In the course of a preliminary survey of the New Zealand Ter- tiary Spatangoida by one of us (H. B. F., 1947-55), the genus herein named Taimanawa was recognized as new. However, the two fossil species referable to the genus seemed to be congeneric with an unnamed and poorly known form from the Kei Islands, west of New Guinea, which had tentatively been referred to Plagio- brissus by Mortensen (1951). Therefore, it was deemed advis- able to defer publication of the material until the fossils could be compared with the suspected living species. In the interim, the responsibility for describing the entire New Zealand collection of Tertiary spatangoids has been assumed by the second author. In consequence, this report has been prepared as a joint project. Through the kindness of Dr. F. Jensenius Madsen (University Zoological Museum, Copenhagen) it has been possible to study fragments of two specimens from the Kei Islands and to confirm 2 BREVIORA No. 320 that the) represent a living species of Taimanawa. I his species, together with /. pulchella (the type of the genus) and /. greyi (Hutton), which was described from New Zealand a centur) ago, are all treated here. I wo additional species occur in the New Zealand Tertiary, Inn as the) add little to the definition of the izenus. the) will he described in a monograph of New Zealand Tertian Spatangoida now in preparation. I he closest relatives o\ taimanawa are the genus Gillechinus Fell, 1964, based on G. cudmorei Fell, and a number of species which previousl) have been referred to Brissopatagm Cotteau, 1863. Brissopatagm is based on li. caumonti Cotteau. which is known from a single. poorl\ -preserved specimen. Recently Phillip i 1966) has synonymized Gillechinus with Brissopatagus. In order to evaluate this contention and to determine the relationship o\ Brissopatagus and Gillechinus to Taimanawa, type material of li. caumonti and G. cudmorei have been compared. Paratypes oi the forme: were supplied In Mi I V Darragh (National Museum o\ Victoria), and the holotype o\ the latter was kindl) lent to us b) Professor J. Roger (Laborator) o\ Paleontology, University of Paris). \s .i result o\ this study, Brissopatagus and Gillechinus are here maintained as distinct, and most o\' the species previousl) referred to Brissopatagus have been regrouped in Gillechinus. Tai- manawa is perhaps the direct descendant of Gillechinus; it is less closel) related to Brissopatagus. I he authors wish to thank Dr. ( \ Fleming and Mr. I. \\ Keyes (New Zealand Geological Survey), Mr. .1. D. Campbell (Otago University), Mr. J. A Grant-Mackie (Auckland Univer sit\). Mr. D. R. Gregg (Canterbur) Museum), and Dr. P. Kier (Smithsonian Institution) for the loan o\ specimens under their charge, and Mr. R .1. hosier (Hughesdale, Victoria, Australia) for a gift ot specimens of Gillechinus cudmorei. The project was under- taken during tenure at Harvard Universit) o\ a Frank Knox Memorial Fellowship and a Taranaki Post-Doctoral Fellowship by one of us ( R. A. H.). The research was sponsored by a National Science Foundation grant, GB-3532. Repositories. The repositories iA type and other specimens are indicated by the following abbreviations: New Zealand Geological Survey ( F.C. ): Geolog) Department. Otago Universit) (OU.); Geology Department. Auckland University (E.): and Canterbury Museum (zfe. ). Localities. Localities of the New Zealand fossil specimens have been arranged in order of the sheet districts of the one mile to the 1969 INDO-WEST-PACIFIC ECHINOIDS 3 inch topographic map series (New Zealand Department of Lands and Survey, series N.Z.M.S. 1 ) on which they appear. The sheet districts are designated "N" and "S" for the North and South Islands and are arranged numerically from north to south on each island. Sheet Fossil Numbers of the New Zealand Fossil Record Form System (e.g. SI 64/496) are recorded for those localities to which they have been allocated. Stratigraphy. The sequence of New Zealand Tertiary stage di- visions relevant to the stratigraphic ranges of T. pulchella and T. greyi, listed in order from youngest to oldest, are as follows: Whaingaroan, Duntroonian (Oligocene); Waitakian, Otaian, Awa- moan (Lower Miocene). Order Sparangoida Claus, 1 876 Family Brissidae Gray, 1855 Genus TAIMANAWA nov. Type species. T. pulchella sp. nov. Other included species: T. mortenseni sp. nov.; T. greyi (Hutton). Diagnosis. Large, flattened forms with peripetalous, subanal, and internal fascioles. Frontal ambulacrum deeply depressed, cross- ing the ambitus in a deep, narrow frontal notch. Paired petals long, narrow, shallowly depressed, and with subparallel pore series. An- terior pair widely splayed, posterior pair less so. Primary tubercles crenulate, restricted to the posterior plate series of the paired inter- ambs within the peripetalous fasciole. Labrum very short, extend- ing only halfway along the first plate of the adjacent ambulacra. Etymology. The name Taimanawa is derived from the Maori words tai (= sea) and nianawa (= heart); it is here treated as a Latin feminine first declension noun. Description. Test large, flattened, cordiform in outline, and al- most as wide as long. Aboral surface gently convex, oral surface flattened, ambitus sharp. Apical system at or near the axis of maxi- mum height, usually slightly pre-central and ethmolytic, with four gonopores. Peristome crescentic, labiate, and anterior; periproct situated on the truncated posterior test surface. Well-developed peripetalous and subanal fascioles and a clear but less well-devel- oped internal fasciole. Neither the peripetalous nor the internal fasciole continuous across the frontal ambulacrum. Both primary and secondary tubercles perforate and crenulate. Aboral surface: Frontal ambulacrum apetaloid, deeply depressed, crossing the ambitus in a narrow, deep frontal notch; ornamented 4 BRE\ [OR \ No. 320 with fine, dense miliary tubercles and perforated by minute, conju- gate pore pairs. Paired petals long, narrow, and weakly depressed. with subparallel pore series. Anterior petals widely splayed, at an angle oi approximatel) 120 : posterior petals less so. tunning an angle of approximate!) ("»<> . Each pore pair weakly conjugate and separate.! from the next In a single row of miliary tubercles. Inter- ambs with a broad, low ridge extending along or near the median suture oi the plate series; ridges oi the paired interamhulaera sub- tend gently sloping, flattened surfaces posteriorly to the depression ol each petal. Posterior plate series of the paired interamhulaera bear prominent fields of primar\ tubercles located within the peri- petalous fasciole. Tubercles surrounded by large, shallow scrobi- cules; those oi each field generall) arranged in a reticulate pattern with discernible rows both parallel and perpendicular to the axis oi the adjacent petal. Median suture of the posterior interamh bordered on each side by a single or double row of well-spaced, coarse secondary tubercles. Haeh interamhulaera] plate of the two series bordering the frontal ambulacrum bears ;i small triangular field iA course secondary tubercles. I he remaining intcrambulacral areas of the aborul surface bear tine, dense seconders and miliary tubercles. Oral surface: Plastron amphisternous, weakl) keeled, and ex- tending anteriorl) almost to the peristome; it bears coarse, dense secondary tubercles arranged in lines radiating from the posterior oi the keel. Subanal fasciole broad and rcniform. enclosing ;i densely tuberculate area with an arcuate row ol conjugate pore pairs on each side Labium \er\ short, extending only halfway along the first plate oi the adjacent ambulacra, twice as wide as long, and with a few secondary tubercles. Remaining interambu- laeral areas bear well-spaced secondary tubercles. Ambulacra! areas not depressed, naked except for scattered miliary tubercles; those margining the plastron are conspicuous, being half as wide as the plastron itself. Secondary spines bear prominent longitudinal ribs and are akin to those oi most other spatangoids. Remarks. Judging from the faseioles alone. Taimanawa would be placed in the Loveniidae alongside Breynia, which is the onl) other spatangoid genus with peripetalous, subanal. and internal faseioles. However, its other characters show that it is unrelated to Breynia and that it cannot be referred to the Loveniidae in spite of possessing an internal fasciole. In general morphology it is very similar to the undisputed brissid genera Gillechinus, Plagiobrissus, Brissopatagus, Macropneustes, and Eupatagus. Its long, narrow petals with subparallel pore series, short labrum. erenulate primar\ 1969 INDO-WEST-PACIFIC ECHINOIDS 5 and secondary tubercles, and the absence of ampullae set it apart from the loveniids and necessitate its inclusion in the Brissidae. Taimanawa, by virtue of its internal fasciole, is unique among the Brissidae. Its nearest relative is Gillechinus, which also has primary tubercles restricted to the posterior plate series of the paired interambulacra within the peripetalous fasciole and which has similar petals. Most members of Gillechinus possess flattened in- terambulacral surfaces subtended posteriorly from ridges of the paired interambs into confluence with the depressions of the paired petals, as in Taimanawa. However, Taimanawa, in addition to possessing an internal fasciole, is distinguished by its shorter labrum (Fig. 1 ), its much more deeply depressed frontal ambulacrum, and its much stronger frontal notch. It is interesting to note that the earliest species of Taimanawa, from the late Eocene of New Zea- land (as yet undescribed), resembles contemporaneous Gillechinus in possessing a slightly depressed frontal ambulacrum and a rela- tively weak frontal notch. Such a relationship strongly suggests that Taimanawa was derived directly from Gillechinus. The genera Eupatagus, Gymnopatagus, Macropneustes, Plagiobrissus, and Brissopatagus all bear some resemblance to Taimanawa. Besides lacking an internal fasciole, they are distinguished by possessing primary tubercles on both plate series of at least the posterior paired interambs (with the exception of E. ibericus Lambert). Eupatagus, Plagiobrissus, and Brissopatagus possess a weak frontal groove and frontal notch, and even in Macropneustes and Gymnopatagus, where these structures are better developed, they are not as strong as in Taimanawa. Eupatagus and Gymnopatagus are further dis- tinguished by possessing spatangiform petals with curved pore series, Plagiobrissus by possessing lateral branches of the subanal fasciole, and Brissopatagus by the concave depressions of its an- terior paired interambulacra. The presence of an internal fasciole in Taimanawa. was initially a perplexing feature, since this structure has not previously been reported from any member of the Brissidae and, indeed, Morten- sen's (1951) diagnosis of the family excludes forms with an in- ternal fasciole. However, it will be recalled that a similar perplexity arose in reviewing New Zealand species of the spatangid genus Paramaretia (Fell, 1963), though in this case the fasciole proves to be a transitory feature that disappears in the adult. The rela- tionship of fascioles to environmental features and the loss or ac- quisition of fascioles during the life-span of certain spatangoid echinoids have recently been studied by R. H. Chesher (private (-, KK 1 \ lORA No. 320 communication), and the whole subject has now been considerably clarified b) Ins recent publication ( 1968). Most of the New Zealand Tertiai \ specimens of Taimanawa are from detrital limestones and from sandstones that were deposited in comparatively shallow water during the early and mid- Tertian transgression of the New Zealand region. In contrast, the extant species is recorded by Mortensen ( 1951 ) from two deep-water sta- tions (260 and 268 meters); the sediment type is known lor onl\ o\w of the stations and is recorded In Mortensen ( 1 c> 2 3 ) as mud and shells. I he bottom temperature of the two stations is not known but is probabl) near 20 (' and therefore some s -lo C higher than the mean annual temperature of surface water oil the southeast coast of the South Island. New Zealand, where Taiman- awa was abundant in OligOCene and earliest Miocene time. Such a relationship is consistent with the higher temperatures interred for the mid-Tertiar) marine environment of the New Zealand region on the basis ol U'smI echmoids ( I ell. 1956), and b) Bcu ( 1966) and others on the basis of Other fossil groups KEY TO SPECIES OF TAIMANAWA I (2i( renulation ol the primary tubercles strongl) developed, contiguous with the mamelon, the platform therefore aborted ; 2(1) Crenulation of the primar) tubercles restricted to the peripher) of the platform, widel) separated from the mamelon mortenseni ) (4) 23-40 primary tubercles in each interambulacral field, 22-2^ well developed pore pairs in the anterioi petals K" w 4(3) 12-16 primar] tubercles in each interambulacral field, 18-19 well developed pore pain in the anterior petals pulchella TAIMANAWA PULCHELLA sp. nov. Plate 2, fig. 2; Plate 3; Plate 4. Figures 1 , 2a, 2c. Holotype. OU. 8590, Karitane, Otago; SI 55/538. Paratypes. S75: E. 310. Burnt Hill. Trelissick Basin. mid-Canter- bury; S75 517. SI 27: EC. 405, Ngapara, south Canterbury; SI 27/ 561. S155: OU. 8540, Puketeraki, Dunedin district. Otago'. E. 302. 304, Waikouaiti North Head. Matanaka Beach. Otago: SI 55 540. S163: EC. 42h. Blackhead. Dunedin district. Otago; SI 63/495. OU. 4736. 8566a, b, 8567b-d, 8568a-d, Blackhead. Dunedin dis- trict. Otago. SI 64: EC. 437. Green Island district. Otago; SI 64/ 496. OL I. 8585, New Zealand Cement Co. pit, Burnside. Otago. Locality unknown: EC. 67 I . 969 INDO-WEST-PACIFIC ECHINOIDS Figure 1 . Comparison of the peristomial regions of Gillechinus and Tai- manawa. a) Gillechinus alabamensis (Cooke), U.S. National Museum 562467; x3.5. b) Gillechinus cudmorei Fell, Museum of Comparative Zo- ology 4147; X3.5. c) Taimanawa pulchella sp. nov., holotype, OU. 8590; X2.5. 8 BREVIOR \ No. 320 Diagnosis. Adult tests compressed, with a moderately depressed Frontal ambulacrum and a strong frontal notch; 18-19 well-devel- oped pore pairs in the anterior paired petals. 12-16 primary tubercles in each lateral intcrambulacrum. Description, lest up to 120 mm in length, cordiform in outline, almost as wide as long, with a sharp ambitus. Aboral surface shallowly arched, oral surface flattened; height breadth ratio ap- proximately o.27. Apical system central to slightly anterior in position. King on the axis of maximum height of the test and morphologically typical of the genus. Peristome crescentic, an- terior, and margined b\ a prominent lip. IVriproel lies on the truncated posterior surface of the test; no specimen is sufficiently well preserved to show its shape. Peripetalous fasciole thin, not indented between petals. Internal lasciole thin, of variable width, located near the median suture of the anterior mterambs and form- ing a tight loop posteriori) around the apical sWem. Neither the peripetalous nor internal fasciole is continuous across the frontal ambulacrum. Subanal fasciole well developed and reniform. \boial surface: Frontal ambulacrum of moderate depression; frontal notch prominent, as wide as deep in adult tests. Paired pel. ils straight, lightly depressed; anterior pair generally slightly longer than those posterior. Pore pairs weakly conjugate and o\ate. Adult tests have IN-IC> pore pairs in the anterior petals and 15-16 in those posterior. Primary tubercles restricted to the posterior plate- series of lateral intcrambs within the peripetalous fasciole. Adult tests with 12-lh tubercles m each such interambulacral field. Other features ol the aboral surface typical Of the genus. Oral surface: Labium very short, reaching only halfway along the first plate of the adjacent ambulacra. I here are eight pores m side the subanal lasciole. arranged in two lines of lour parallel to the lateral portions of the fasciole. In other characters the oral surface is characteristic o\ the genus. Remarks. The most distal row {)\ primary tubercles in the inter- ambulacral fields, .is in the other species i)\ Taimanawa, is gen- erally Hush with the distal end of the adjacent petal. I he anterior intcrambs of OU. 8540, however, have two additional rows o\ tubercles nearer the ambitus than the distal tip <^ the adjacent petal. Ljoing an unusually high number o\ tubercles (22) compared to that oi more normal specimens ( 12-16). One specimen (EC. 671 ) has a conspicuous boring perforating the test at its anterior margin. evidently the result oi an attack by a large, predators gastropod. . Lower Miocene; Waitakian — Otaian. OU. 8585 is known 1969 INDO-WEST-PAC1FIC ECHINOIDS 9 to be of Waitakian age and EC. 405 has the age limits of Dun- troonian — Waitakian. The remaining specimens, with the excep- tion of E. 310, are from the Caversham Sandstone, which is largely of Otaian Age although Awamoan microfaunas are known from its uppermost horizons. The holotype is known to be of Otaian Age, and the other specimens from the Caversham Sandstone are most likely of Otaian Age also. TAIMANAWA GREYI (Hutton) Plate 1 ; Plate 2, fig. 1 . Figure 2d. Brissus greyi Hector, 1870: 192 {nomen nudum). Eupatagus greyi; Hutton, 1873: 41. Holotype. EC. 682, Cobden, north Westland; S44. Additional material. N51 : E. 296, near Waikawau Stream mouth, west Auckland; N5 1/677. S37: EC. 480, 481, Woodpecker Bay, north Westland; S37/528. S44: EC. 458, Cobden Limestone, Grey- mouth, north Westland; S44/465. EC. 543, Cobden Limestone Quarries, Greymouth, north Westland; S44/476. zfe. 286, 287, 290, Cobden Limestone, Greymouth, north Westland. S102: EC. 407, Kakahu River, south Canterbury; S102/1. OU. 8548b, Hang- ing Rock, Opihi River, south Canterbury. Sill: E. 250, Lower Pareora Gorge, south Canterbury; SI 11/683. OU. 8606, Gordon Valley, south Canterbury, zfe. 284, Pareora Gorge, south Canter- bury. S127: EC. 615, Pigeon Rock, Waitaki Valley, south Canter- bury; SI 27/37 1. EC. 473, 474, Waihao River, south Canterbury; SI 27/376. EC. 472, Waitaki Valley, south Canterbury; SI 27/447. EC. 471, Awamoko Creek, Oamaru, south Canterbury; SI 27/450. E. 246, 248, Taylor's Road, Oamaru, south Canterbury; S127/639. OU. 8546, 8547, Waitaki Limestone, Duntroon, south Canterbury. OU. 8542, Ngapara, south Canterbury. OU. 8598, 8601, 8602. Earthquakes, Duntroon, south Canterbury. S136: OU. 8545, Weston, south Canterbury. Locality unknown: zfe. 278, Canter- bury, unlocalised. Diagnosis. Adult tests compressed, with a deeply depressed fron- tal ambulacrum, 22-25 well-developed pore pairs in the anterior petals, and 23-40 primary tubercles in each interambulacral field. Description. Test up to 1 34 mm in length, cordiform in outline, almost as wide as long, with a sharp ambitus. Aboral surface shallowly arched, oral surface flattened; height/breadth ratio approximately 0.25. Apical system varies in position from slightly 10 BR F MORA No. 320 anterior to slightl) posterior oi center; it lies on the axis i>t maxi- mum test height, and its morpholog) is typical o\ the genus. Peri- stome crescendo, anterior, and margined by a prominent lip. Anal region not preserved on any available specimens. Peripetalous fasciole thin, not Strongl) indented between the petals. Internal fastiole extending lor a short distance near the sutures of the two plate series of interamhs 2 and 3 and forming i tight loop pos- tcriork around the apical system. Neither the internal nor the peripetalous fasciole closed across the frontal ambulacrum. t ■ ft --- ft*^ \ a j -> - *&%* '1? I igure 2 a) Primar) tubercles of Taimanawa pukhella sp. run., holotype, OU. 8590; x7. b) Primary tubercles of Taimanawa mortenseni, holotvpe: X7. c) Plates of ambulacrum V of Taimanawa pulchella sp. nov.. holotype, OU. 8590; x7. d) Apical system of Taimanawa greyi (Hutton). OU. 8598; X7. 1969 INDO-WEST-PACIFIC ECHINOIDS 11 Aboral surface: Frontal ambulacrum deeply depressed, termi- nating in a conspicuous frontal notch that is as wide as deep on adult tests. Paired petals lightly depressed; anterior pair straight or gently curved towards the frontal ambulacrum and generally slightly longer than the posterior pair, which are straight or gently curved away from the posterior interamb. Pore pairs ovate and weakly conjugate; adult tests have 23-25 well-developed pore pairs in each anterior petal and 20-24 in each posterior petal. Fields of primary tubercles restricted to posterior plate series of the lateral interambs inside the peripetalous fasciole; 23-40 tubercles in each field on adult tests. Other features of the aboral surface as de- scribed for the genus. Oral surface: As described for the genus. The anal region is poorly preserved on all the available specimens; the presence of a subanal fasciole cannot be confirmed, and the nature of the tuber- culation within the anal fasciole is not known. Remarks. The material, although abundant, offers but a single, indifferently preserved oral surface. The petals of different speci- mens are somewhat variable in length, breadth, and the degree to which they are depressed; the anterior petals of several specimens of about the same test width (1 14-119 mm) vary in length from 37 to 42 mm and vary in width from 5.0 to 6.5 mm. The number of primary tubercles in the interambulacral fields is also variable. Three specimens from the Cobden Limestone, Greymouth (zfe. 286, 287, 290), and a single specimen registered as from Ngapara, Oamaru (OU. 8542), differ from the remainder of the material in possessing more numerous primary tubercles (35-40 in each inter- ambulacral field) and in bearing up to four rows of well-spaced, coarse secondary tubercles situated adjacent to and running parallel with the ambitus of the anterior interambs. The matrix of OU. 8542, a hard, fine-grained, muddy limestone, contrasts with the softer, coarse-grained, almost pure limestone that forms the matrix of other echinoids from Ngapara; it closely resembles the matrix of the Greymouth specimens. It may be that the locality of OU. 8542 has been incorrectly recorded. Although the holotype has never been figured, it is known to be the specimen used by Hutton in the erection of the species. It is badly crushed and shows few of the characters of the species, but as it has at least 25 pore pairs in ambulacrum IV and approxi- mately 36 primary tubercles in interambulacrum 4, it is un- doubtedly conspecific with the other specimens herein referred to T. greyi. 12 brf.viora No. 320 Taimanawa greyi closely resembles /'. pulchella, to which it was probably ancestral. It is distinguished by its petals, especially the anterior pair, having more numerous pore pairs (Fig. 3). Further differences include the deeper frontal goove and more numerous primary tubercles oi T. greyi. Age. Oligocene — lowermost Miocene; Whiangaroan — Waita- kian. The exact age of the holotvpe is not known. From its local- ity and matrix, it can confidently be assigned to the Cobden Lime- stone, which ranges in age from \\ hiangaroan to Waitakian. TAIMANAWA MORTENSENI sp. nov. Plate 5, fig. 1 . Figure 2b. Plagiobrissus sp. ? Mortensen, 1951, p 503, pi. 39, tig. 3. Holotype. Fragments of a specimen from Station 32 of the Danish Expedition to the Kei Islands ( 1922), between Little Kei and Tajando Islands at a depth o\ 260 meters. Ciliated by the University Zoological Museum. Copenhagen. Paratype. Fragments o\ a second specimen are represented in the collection from the same station. Description, Known from fragments onl\. the largest of which comprises the anterior half o\ an ahoral surface, lest huge, and appears to be approximated as wide as long. Aboral surface in- Bated and ambitus more hroadU rounded than in other members o\ the genus. Peripetalous fasciole well developed, not indented be- tween the petals and not continuous across the frontal ambulacrum. Internal fasciole thin. o\ variable width, bordering the frontal am- bulacrum for half the distance from the broken margin of the test, slightb anterior of the apical system, to the peripetalous fasciole. Subanal fasciole well developed and. as far as can be judged. reniform. Vboral surface: Frontal ambulacrum deeply depressed, apetaloid. terminating in a pronounced frontal notch that is approximately as wide as deep. Amb plates bear a fine, dense, secondary and miliar) tuberculatum and are perforated by minute, conjugate pore pairs. interior petals slightly depressed, splayed at an angle of some 120 . and weakk tlexuous. being concave away from the frontal ambulacrum proximallv and concave towards the frontal ambu- lacrum distally. Each bears 25 ovate, weakly conjugate pore pairs; adjacent pairs are separated by a single row of secondary and mili- ary tubercles, and the interporiferous zone bears scattered miliary tubercles. Anterior plate series of the frontal interambulacra tumid, subtending a weakly concave surface posteriorly into confluence 1969 INDO-WEST-PACIFIC ECHINOIDS 13 with the depressions of the frontal petals. Primary tubercles re- stricted to the posterior plate series of the anterior interambs lying within the peripetalous fasciole, surrounded by prominent scrobi- cules, and very weakly crenulate. The partially preserved anterior plate series of interamb 4 devoid of primary tubercles, suggesting that tuberculation of the posterior lateral interambs is akin to those anterior. Frontal ambulacrum margined by a zone of coarse sec- ondary tubercles. The remaining aboral interambulacral surfaces bear a dense miliary and secondary tuberculation. Oral surface: Nature of the plastron and labrum unknown. In- terambs bear well-spaced secondary tubercles; ambulacra naked except for scattered miliary tubercles. The area enclosed by the subanal fasciole bears dense secondary tubercles and is perforated on each side by seven conjugate pore pairs. 150 140 130 _* 120 Ui ~* 110 CL 100 90- 80- 70 60 50 / / / / / / • / / / / i • / / / / / x / • / / / / / / • • / / / T. pulchella / T. greyi X / / / / / / / H 15 16 17 18 19 20 21 22 23 24 25 26 pore pairs Figure 3. Plot of the number of fully developed pore pairs in each plate series of the anterior petals against test width for Taimanawa pulchella sp.. nov. and T. greyi (Hutton) showing the separation of the two species. Ordinate values have been estimated for many points and are accurate only to ± 5 mm. 14 imrvioRA No. 320 Remarks. Mortensen (1951) also recorded fragments ot this species from Station 44 o\ the Danish Expedition to the Kei Islands ( lc>22). but the material is poor (Dr. F. Jensenius Madsen, private communication) and has not been examined in the preparation of this report. /'. mortenseni is conformable in all its principal char- acters with the fossil T. pulchella and I . greyi but represents a discrete species. I he possession of an internal fasciolc and a strong frontal notch shows that mortenseni must be referred to Taiman- awa. Indeed, the \er\ close resemblance between the mid- 1 ertiar\ New Zealand species and the present form, which is living in the Kei Island area ^\ the Indonesian region, is remarkable. I he holotype represented b\ the most complete set of fragments would have measured approximately 140 mm in width. 1 he test height cannot be accurately estimated, but the aboral surface is more inflated than that of either /. pulchella 01 / greyi, and the ambitus is more rounded than in either o\' the fossil species. I he anterior petals resemble those of /. ( species. H. caumonti Cotteau ( 1863: 144. pi. s. figs. 3-7); b) subsequent designation (Cotteau 1886: 135). Brissopatagus Cotteau, 1863: 144; Pomel, I S K 3 : 32. pars; Duncan and Sladen, ISS4: 226. pars; Duncan, 188 > 250, pars; I amberl and I hiery. 1924: 491. pars; Cooke, 1942; 57, pars; Mortensen, 1951: 453. fnus: ( ooke, 1959: 92, pars; Phillip. 1966: I 14. pars; Fischer, 1966: 5S4. pars. Brissospatangus ( otteau, 1XX6: 135. pars. nun Brissospatangus, ( otteau, 1890: IS. Diagnosis, rest small, with the test height approximately halt the test length, bearing siibanal and peripetalous fascioles. Frontal ambulacrum narrow and Lightl) depressed, forming a narrow, shal- low frontal notch. Petals lightl) depressed with subparallel pore series; anterior pair widely splayed, posterior pair less so. Anterior interambs with conspicuous depressions confluent with depressions of the anterior petals. Primary tubercles present on the anterior plate series of the frontal interambs. 1969 INDO-WEST-PACIFIC ECHINOIDS 15 Remarks. Brissopatagus is a poorly defined genus. It was pro- posed for two incompletely known species: B. caumonti from the Eocene of France; and B. javanicus Cotteau (1863: 144), origi- nally described as Spatangus? sp. by Herklots (1854: 13, pi. 3, figs. 2, 2a, 2b). The latter was reported from the Eocene of Java by Cotteau, but Gerth (1922: 520) revised the age to middle Miocene. Each of the species is apparently known from a single specimen; that representing B. javanicus is an internal mould and shows no tuberculation, whereas that representing B. caumonti is an eroded, somewhat deformed specimen. So far as their mor- phology can be observed, the two species show good agreement in test shape and in the nature of the ambulacra. B. caumonti, the type species by subsequent designation, forms the basis of Brissopatagus and is redescribed and refigured below. The specimen shows traces of a peripetalous and a subanal fasciole which have not hitherto been recorded. Cotteau's figures of the specimen, as typical of fossil echinoid illustrations of that time, are idealized. The distribution of primary tubercles is of particular significance. Those shown in Cotteau's figures are fictitious; the original specimen shows traces of a few primaries on the anterior plate series of the anterior paired interambs, but the remainder have been removed by erosion. Brissopatagus, in the sense of the type species, is closely related to Eupatagus, with which it has been grouped as a subgenus by some authors (Dames, 1878; Cooke, 1959). It is also closely related to Macropneustes. The only character by which it can adequately be differentiated from these two genera is the depres- sions of the anterior interambs. The genus is also affiliated with Gillechinus Fell (based on G. cudmorei Fell, 1964, p. 213, pis. 1. 2), which is discussed in detail below. Phillip (1966: 114) has proposed that Gillechinus be treated as a synonym of Brissopatagus. A comparison of the type species of the two genera shows at once that such a view is untenable. G. cudmorei lacks the interambu- lacral depressions of B. caumonti, an essential character of Brisso- patagus, as Fischer (1966) indicates. Further, G. cudmorei is clearly distinct from B. caumonti in that its primary tubercles are restricted to the posterior plate series of the paired interambs. If Brissopatagus is to be taken as including forms lacking interambu- lacral depressions and possessing primary tubercles on the anterior plate series of the paired interambs, then it is transitional to Eupa- tagus through species like E. lamberti Fourtau and E. cordiformis Duncan and Sladen and is indistinguishable from Macropneustes. 16 BRJ V IORA No. 320 Subsequent to its erection, a number of Species have been re- ferred to Brissopatagus. Several possess characters of both Brisso- patagus and Gillechinus, so that their eorreet generic assignment is not immediately apparent. For the reasons given in the discus- sion of Gillechinus, those with primary tubercles restricted to the posterior plate series o\ the paired interambs are referred to that genus. The remaining species are II. colligoni Lambert ( 1933: 37. pi. 4. fig. 21) from the Eocene of Madagascar and II. SUlldaicus Bohrn (1882: 365, pi. 2. figs. 2a. b) from Madura Island north o\ Java. 11. sundaicus is no older than Miocene in age. as no Lower lertiarv rocks are known to crop out on Madura Island (van Bemmelen, 1949: 106). This species, like 11. javanicus, possesses distinct interambulacral depressions, but both species are repre- sented b\ internal moulds alone, so that the nature oi their tuber- culatum is not known and the) cannot be referred to cither Brisso- patagus or Gillechinus with certainty. B. collignoni was described from crushed material that shows primary tubercles on both plate series of the paired interambulacra. and thus it agrees with Brisso- patagUS rather than Gillechinus. However, Us interambulacral depressions are \er\ wcaklv developed, and it mav prove to be best included in EupatagUS 01 Vf/i © ~ 2 o ^ -2 3 c u_ c ™ cl, a. <~ * R T3 . Q-, 2 •• 3 O > >/i © q >/-i © q >/-i i/i © >, 5 - « g c ni *** ® t' iri u-; u-; i-; ^ t^ ■" 5 0^"^ "S ^ "3 +-* C *a _C 55 « 5 G. «* < z ca c u ©>/1so^>o©i^, ©o c'?D"'o^'o q. o\ — x}:t^odvd— '06 — ooTf'06 «o.o « <- W Q_ — r<1r<1-^r>ir X! ~ h .S-.S T3 O — ~ u z - 'ONooOOtOO^-, 1/1 ■ - 4§ C , Cn ? ca c8 < £ « jc .2 2 2 Cm ©on— .'^/i©qqqq/i /i©osO\oorJoo>/iosr~-r/1/isOsO^OsOsOl^'- . „ X £ o , O -3 — ^ ^ •£ O c « '£ U | J|a5^£ ij < |! qpirjnqqoi;^qovi w ^ -o •- °* g zb oq 2 ^ ■^■'vdsdrn>^>ovdvdvovdr^ri x; ^ q,13 n ^l << £ o_.2 ~ 'C x ^ S q, qqqqoqoooqovi "> =a « > .22 <-> .£ Hco —- u ~ O- c C 5 2 X - -s | tL, oo©/i©©o©i/i u 3 j2 tJ M c 33 r/isosot^sosor~ — < ™ u, u " P 1: o ti o* Z -So 4) CO . C u i_ ■- xi « 2 ©©© o© ^.tv^fe.SuSc <5 ^ O 3 O r~- oo <^ ^ Tt vo Tt wi os rt vd >o ^ ob 1? o &> -a & so © © — 0 0 c 2 voo\vDtm^a\^t- ©©— "S E T» " _o u. i*i«^/ii/ii/it-->>osoi: -^u^y^T:^ OOOO-^-Tj-SDOOOOOOTt-OOOO^ ~ O i« *« E u . ■ ■ . • • S =L-'CO'-x«x: 0 D O O U^^ D uD D 5 ,• « " -2 g S f OOUUUOOOWOO 5 ^c.E-u^* " c u 22 BREVIORA No. 320 LITERATURE CITED Bl \1\11 i » N, R. W. VAN 1449. The Geologv of Indonesia, Vol. I A. General Geology. Govern- ment Printing Office, The Hague, 732 pp. Bl I . A. G. 1466. Sea temperatures in New Zealand during the Cenozoic Era as indicated by molluscs. Trans. R. Soc. N.Z.. 4(9): 177-187. BOHM, V 1882. Einige Tertiare Fossilien von der lnsel Madura nordlich von Java. Denk. kais. Akad. Wissen., 45(2): 359-372. pis. 1-4. < \m i \. L. 1930. Revision des echinides du nummulitique du Departement des 1 andes. Actes Soc. Linn. Bordeaux, 82: 5-69. pis. 1-4. ( HI SHKR. R. H. 1968. The systematic^ oi sympatric species m West Indian spaiangouK Stud. Trop. Oceanogr., Miami. 7: vii -f 168, 35 pis. COOKJ . C. W. 1942. Cenozoic irregular echinoids oi eastern United States. J. Paleont., L6( I): 1-62. pis. 1-8. 1959. Cenozoic echinoids of eastern United States. Prof. Pap. U.S. Geol. Surv., 321: 1-106, pis. 1-43. Cotteau.G. H. 1863. Echinides fossiles des Pyr6nees. Congr. Scient. fiance. Bor- deaux: 1-160. pis. 1-9. iss> Paleontologie francaise. Terrain Tertiaire. Eocene, ser. 1. I. 1889. 672 pp.. 200 pis. 1890- Echinides Eocenes de la Province d'Alicante. Mem. Soc. Geol. 1891. France, ser. 3, 5(2): 1-107, pis. 22-32. Dami s. W. 1878. Die Fchiniden der Vicentinischen und Veronesischen Tertiaei ablagerungen. Palaeontographica, 25: 1-100, pis. 1-11. Die M RSON. R E., AND W. S. W. Kl W 1917. The fauna of a medial Tertiary formation and the associated horizons of northeastern Mexico. Proc. California Acad. Sci., ser. 4. 7: 125-156. pis. 17-26. DuN< w P. M. 1889. A revision of the genera and great groups of the Echinoidea. Linnean Soc. London Jour.. Zoology. 23: 1-3 11. I)l N( \n. P. \1 . \m> W. P. Sladen 1882- Fossil Echinoidea of western Sind and the coast of Baluchistan 1886. and of the Persian Gulf, from Tertiary formations. Mem. Geol. Surv. India. Palaeont. Indica. ser. 14. 1: 1-392, pis. 1-58. Fell. H. B. 1956. Tertiary sea temperatures in Australia and New Zealand from the evidence of fossil echinoderms. Proc. XIV Intern. Cong. Zool. Cophenhagen, 1953: 103-4. 1969 INDO-WEST-PACIFIC ECHINOIDS 23 1963. The spatangid echinoids of New Zealand. Zool. Pub. Vict. Univ. Wellington, 32: 1-8, pis. 1-6. 1964. New genera of Tertiary echinoids from Victoria, Australia. Mem. Nat. Mus. Victoria, 26: 211-215, pis. 1, 2. Fischer, A. G. 1966. Asterozoa-Echinozoa. ///: R. C. Moore ed.. Treatise on Inverte- brate Paleontology, Pt. U, Echinodermata 3. Univ. of Kansas Press, pp. 543-640. Fourtau, M. R. 1908. Sur quelques echinides Eocenes d'Egypte nouveaux ou pen connus. Bull. Inst. Egyptien, ser. 5, 1: 205-220, pis. 1, 2. Gerth, H. 1922. Die Fauna von Java. Echinodermata, Echinoidea. Samml. d. geol. Reichs Mus. Leyden, 2(4) : 497-520, pi. 52. Herklots, J. A. 1854. Description des restes fossiles d'animaux des terrains Tertiaires de File de Java. Quatrieme partie, Echinodermes. E. J. Brill, Leide, 24 pp., 5 pis. Hector, J. 1870. Catalogue of the Colonial Museum, Wellington, New Zealand. Wellington, 235 pp. Hutton, F. W. 1873. Catalogue of the Tertiary Mollusca and Echinodermata of New Zealand in the Collection of the Colonial Museum. Govern- ment Printer, Wellington, 48 pp. Lambert, J., and P. Thiery 1909- Essai de Nomenclature Raisonne des Echinides. Libraire 1925. Ferriere (Chaumont), 607 pp., 15 pis. Lambert, J. 1933. Echinides de Madagascar communiques par M. H. Besaire. Annls. Geol. Surv. Mines Madagascar, 3: 1-49, pis. 1-4. Phillip, G. M. 1966. Notes on three recently proposed Australian echinoid genera. Proc. Linn. Soc. New South Wales, 91(2): 114-117. POMEL, A. 1883. Classification Methodique et Genera des Echinides Vivants et Fossiles. Adolphe Jourdan (Alger), 131 pp., 1 pi. MORTENSEN, T. 1923. The Danish expedition to the Kei Islands 1922. Saertryk Vi- densk. Medd. Dansk. naturh. Foren., 76: 55-99, pis. 1-3. 1951. A Monograph of the Echinoidea, 5(2), Spatangoida II. C. A. Reitzel, Copenhagen, 593 pp., 64 pis. Sanchez Roig, M. 1951. Faunula de equinodermos fosiles del Terciaro, de termino Municipal de Moron, Provincia de Camaguey. Mem. Soc. Cubana Hist. Nat.. 20(2): 37-64, pis. 23-40. (Received 18 September 1968.) 24 HRI MORA NO. 320 '<• ":v { \ 3 i ^s / IM Ml 1 Figs. 1. 2. Taimanawa greyi (Hutton). dorsal view (Fig. 1 >. lateral view (Fig. 2). OU. 8598. test length 135.5 mm: < K% ). 1969 INDO-WEST-PACIFIC ECHINOIDS 25 .jr.* ■* * >-j J ■ '■■■ 4 J f -!• L *"•* '•*-'*«HH -»■■;•.-/ *-.». PLATE 2 Fig. 1. Taimanawa greyi (Hutton), ventral view. OU. 8598. test length 133.5 mm; (X%). Fig. 2. Taimanawa pulchella sp. nov., lateral view. Holotype, OU. 8590. test length c 110.0 mm; (x%). 26 imi \ [OR \ No. 320 ' | ' _/ f /•"*'' m ,^Aj >• PI \ I I J / manawa pulchella sp. nov., dorsal view. Holotype, 01 8590, tesl length < 1 10.0 mm; < ■ 1 >• 969 INDO-WEST-PACIFIC ECHINOIDS 27 w>- •; -^ ! w -■; PLATE 4 Taimanawa pulchella sp. nov., ventral view. Holotype, OU. 8590, test length c 1 10.0 mm; ( x 1 ). 2S HKl \ [OR \ No. 320 V 4 i "V, \ / U 1,17 jk* »» • 4. ^"ViV • " * 4 1969 INDO-WEST-PACIFIC ECHINOIDS 29 PLATE 5 Fig. 1. Taimanawa mortenseni sp. nov., dorsal view. Holotype, frag- ments of a specimen from Station 32 of the Danish Expedition to the Kei Islands (1922), University Zoological Museum Copenhagen, width of fragment 105 mm; (Xcl). Figs. 2-4. Brissopatagus caumonti Cotteau, dorsal view (Fig. 2), ventral view (Fig. 3), lateral view (Fig. 4). Holotype, 409- 1A- Co 1-2- b 132, Ecole des Mines, Paris, test length 40.5 mm; (Xcl). BREVIGRA Museum of Comnpairative Zoology Cambridge, Mass. 10 June, 1969 Number 321 OBSERVATIONS ON THE AGONISTIC AND BREEDING BEHAVIOR OF LEPTODACTYLUS PENTADACTYLUS AND OTHER AMPHIBIAN SPECIES IN VENEZUELA Juan A. Rivero and Andres Eloy Esteves1 Abstract. Observations were made of breeding aggregations of Lepto- dactylus pentadactylus, Phyllomedusa trinitatis, Engystomops pustulosis, Phrynohyas venulosa and Hyla minuta in northeast Venezuela. L. penta- dactylus makes hollows near the water's edge and produces a foam nest where the eggs are deposited. During amplexus and oviposition, the male moves the feet in alternate, coordinated movements over the cloacal openings of the embracing pair, apparently assuring, in this way, proper fertilization and distribution of the eggs, besides beating the foam into the proper consist- ency. During amplexus, the male shows definite agonistic behavior towards other members of his species or to any other object that may touch him. Quite often fights between males occur. These may last several minutes and consist of standing up and hugging the opponent with the thumb tubercles, pushing, toppling, and apparently embracing, and pulling the adversary towards the breast tubercles. Engystomops seems to prefer road puddles and fouled water for breeding. Its voice is similar to that of a puppy, and a group of this species will stop calling simultaneously, and for relatively long intervals, when approached by an observer. Phyllomedusa was extremely abundant in Dieffenbachia and other broad- leaved plants. Egg clusters were found on leaf surfaces only. It appears that when leaves of the right size are not found, Phyllomedusa does not need to make the leaf nests (funnels) described for many of the species. Besides its normal mating call, the male has a particular release call. It is possible that spent females produce a similar sound. Phrynohyas venulosa were occupying approximately equidistant positions in a shallow pool. Both this species and Hyla minuta, which were extremely common during the first night of observation, had completely disappeared the following night. 1 Instituto Venezolano de Investigaciones Cientificas. BR] mora No. 321 INTRODUCTION On June 25. 1968, the authors made a trip to the region of Guiria. l:.studo Sucre, in the Peninsula de Paria of Venezuela, and collected extensivel) in the vicinity of the village of Rio Salado and on Pica del Mango at a slightb higher elevation (100 m ) . The conditions near Guiria are xcrophytic to mesophytic. but higher up on Pica del Mango the vegetation is quite luxuriant and there is a great abundance oi heliconias. Araeeae and Marantaceae. Most oi the observations reported here were made at Pica del Mango. The excursion to Pica del Mango was made at night after a heavy rain during the afternoon. As a result oi the rain, a depres- sion in the forest had been filled with water to a depth of a few inches. The pool, of about ten by three meters, was undoubtedly temporary, but it ma) have been in existence prior to the rainy period oi that afternoon. The margins, three to live meters beyond the edge of the water, were more or less clear o\ an\ vegetation cx- eept at one end. where there were a number of broad-leaved Mar- antaceae and Dieffenbachia growing in a thicket. I he forest, with its heaw undergrowth of heliconias, CostUS and Araeeae. com- menced at about five meters from the edge of the pool. I he pool was. however, shaded b\ two or three large trees that grew near us margins. I he roar produced In frogs calling from the pool could be heard at a distance o\ about 200 meters. Upon approaching, the noise increase. 1 in intensit) and became almost deafening. Calling from the leaves o\ plants were ll\la miniita. sometimes three, four, or five H' each leaf oi Dieffenbachia. It was often possible to get three specimens with one sweeping movement of the hand over the leal surface. When flashlights were pointed at the water. Phrynohyas venulosa were seen to be occupying the pool, spaced in a pattern at a dis- tance ii\ about three to four feet from one another. As there was no vegetation inside the pool, this "'formation'" could be easil\ observed. Also contributing to the uproar coming from the pool were Leptodactylus pentadactylm and Phyllomedusa trinitatis in great abundance. Engystomops pustulosis in lesser abundance, and a few specimens of Leptodactylus />. petersii. LEPTODACTYLUS PENTADACTYLUS Most attention was given to /.. pentadactylus, since it was hoped that the following night could be dedicated to the other species 1969 BREEDING BEHAVIOR breeding in the pool. Large specimens of L. pentadactylus were everywhere; occasionally one would even step on them. In one place, eight breeding pairs were counted in a space about three meters by one meter, a condition reminiscent of the breeding rookeries of some shore birds. Nests were sometimes eight to ten inches apart, but more commonly they were about one foot from one another. As the eyes of L. pentadactylus shine red at night, it was not difficult to survey the area and locate and count speci- mens. Remaining stationary and moving the flashlight around the pool, we counted 60 individuals, but, if it is considered that nests were more abundant in the Dieffenbachia thicket, that females in nests were probably not seen, and that many specimens were pro- tected from view by other obstructions, the total number may easily have been double that. More specimens, especially females, were pouring in from the forest all the time. Most specimens were males, however. Apart from the mated specimens, which were usually quiet in their nests, unattached in- dividuals were moving all the time, the jumps being short and fre- quently changing in direction. When the butt end of a flashlight was presented to a free male, the animal would move away, and no attempt was made at amplexus ( in contrast to what a Bufo would 8 Q o Id CO a. 4- 2- 0 «►* T T .2 TIME 4 .4 .6 IN SECONDS .8 Fig. 1. — Audiospectrogram of the breeding call of Leptodactylus pei tadactylus, Pica del Mango. Estado Sucre. Venezuela, 25 June 1968. 4 imi mora No. 321 do). Males wore often soon fighting, the two animals rising on their roar logs and attempting to puncture the adversary with their thumb spines b) inward embracing movements of their powerful forelimbs. The movements were brisk, and the light did not usually last more than a few seconds, one of the frogs moving away, or being pushed away, after the first "round." On other occasions, the light lastod for five or ten minutes, and there was much push- ing, toppling (from the raised position), and embracing before the fighl was over.1 Hiese fights created considerable turmoil, and the noise could be heard at a distance of 15 or 20 feet without diffi- culty. In the plant thicket there was much movement o\' the plants as the wrestling animals pushed against them. I he male attached to a female showed agonistic behavior to- wards anything of the right si/e it ,i frog of a smaller species was brought close to the male by the observer, the male would not usualK release its hold but would try to push the animal away with one or both arms. It the "attack" wore more persistent, or it a bigger animal were involved, u would release its hold and fight the intruder in the usual manner, trying to puncture it with the Strong inward movements of the forearms. M the intruder wont away, or if it was moved away, the male it it had boon displaced — would give a low short hops near or around the nest, call, and then come back, either to the original female or to any other neighboring fe- male that was uostmg and was not pre-empted by another male. II the male had not boon displaced, he would just sit above the female, turn around and call repeatedly. It appears that a male does not have an\ strong attachment to an) particular female, and. it it happens to stop on something hav- ing the proper consistency, it gives a tew short hops, turns around several times, and then attempts amplexus. During this time the female is completely passive. During the period of observation, it was the authors' impression that the female does not respond, locally, to an individual male's call. Females were coming to the pool, apparently attracted by the males' calls, but once there they did not seem to go to calling males and. when the) were moving freely, the males did not pay any at- tention to them. In practically all eases of matings observed, the females were Hat on a depression apparently made by them, and 'Similar lights have been described for dendrobatids (Test, 1954; Sexton I960; Duellman, is>: Agar. 1909; von Ihering, 1866) or for this species in Trinidad ( Kenny, 1966), where a funnel made from one or more k\i\cs is used as" a nest. A. and B. but/ ( 1939) wrote that a nest o\ /'. appendiculata was found in a hollow o\ a wet block of stone covered bv a fallen tree trunk in a mountain brook. It appears that when small leaves are not available, Phyllomedusa can adapt lo other conditions. How Phyllomedusa made itself heard under the roaring sound o\ I', venulosa and /.. pentadactylm is difficult to understand Perhaps it comes earlier to the pool; or perhaps the scarcity o\ females can be attribute,! to this factor, kemn (1966:16) reports that the females of Phyllomedusa are also vocal and that males have a mating call (■"huh. huh. huh") and a non-mating call which the) produce throughout the night I he authors onl) heard two calls, one the regular mating call, and the other, a release call (and vibra- tions) produced bv males when other males louch them or attempt amplexus with them. I his call is also produced when a male is grabbed by a human or when several specimens are placed together ui a bag. It is possible that spent females produce a similar sound, although this was not recorded. PHRYNOHYAS VENULOSA I ew observations were made on Phrynohym venulosa, in the hope o\ paying more attention to this species the next day. How- ever, it was observed that they occupy equidistant positions in the pool, that the) retain amplexus alter capture, that one was seen to grab a /.. pentadactylm in amplexus and ride it until they both dis- appeared, and that even when not calling, the two vocal sacs seem to act as floaters, looking as if the animal were wearing water-wing life preservers. The voice of Phrynohym venulosa has been described by a num- ber of authors. The call can perhaps best be described as like the bleating of a goat, but when many specimens are calling together. the noise produced can be compared to that of a low-flying heli- copter or to that produced bv automobiles with holes in their mufflers. 969 BREEDING BEHAVIOR 11 The following afternoon, the surface of the pool was seen to be almost completely covered with what, at first sight, appeared to be the pupae of giant mosquitoes; these were the eggs of P. venulosa. That same night both P. venulosa and Hyla minuta had disappeared completely. And they were neither seen nor heard in another pool a few kilometers away, where they were heard abundantly the night before. 8 © o Ui W id ^2 .4 1.0 1.2 1.4 TIME IN SECONDS 1.6 1.8 2.0 Fig. 3 — Audiospectrograms of breeding choruses. Left, Leptodactylus podicipinus petersii, El Pilar, Estado Sucre, Venezuela, 27 June 1968. Center, Pseudis paradoxus, Rio Salado, Estado Sucre, Venezuela, 25 June 1968. Right, Phrynohyas venulosa, Pica del Mango, Estado Sucre, Vene- zuela, 25 June 1968. It is not known when the forest pool described above was filled with water. If it was during the afternoon rain preceding our visit, then the breeding; of Phrynohyas venulosa lasts only one night. During the periods of observation, no ordinary frog predators were seen. Leptodactylus can easily swallow Engystomops, but it was probably too active, breeding and fighting, to bother about eating. On the first night, two enormous pigs were observed grabbing frogs and apparently eating them (the actual eating was not seen, as the pigs were immediately scared off by the observers, but their insist- ence on coming back left little doubt about their "intentions"). Three of the P. venulosa seen were dead, the other came out of the water dying. The cause of this is not known, but it is not improb- able that they accidentally bumped into a male Leptodactylus, which can be a dangerous adversary. In one instance, Esteves' hand was accidentally grabbed by a male Leptodactylus and, before he could remove it, the animal tried to puncture it with its thumb tubercles 12 BREVIORA NO. 321 by jerky inward movements and did its best to push the hand against the breast spines. A /'. venulosa would probabl) be in serious trouble it' similarly treated. ACKNOWLEDGMENTS I he observations resulting in this paper were made possible b\ the assistance given b\ the Instituto Venezolano de [nvestigaciones ( lenutieas. the institution b\ which both authors were employed at the time the work was conducted. The) are especially thankful to Dr. GilbertO Rodriguez for his help and encouragement in all phases of their work. Partial support was also given b\ NSF grant GB 6944 to Ernesl 1 Williams. bibliography \ GAR, \\ II I Ki n i 1909 I lie nesting habu of the tree frog Phyllomedusa sauvagii. Proc Zool Sot I ondon 1909 ! pi. 84. Hri hi h. Chari is \t . Jr [925 rhe Darien Jungles Nal Hist 25(4): 325-337, 15 figs. 1946. Amphibians and reptiles of the Rio Chucunaque Drainage, I'an- .1111,1. with notes on theii life histories anil habits. Bull. Amer. Mus Nat Hist Hf.isi J75-436, 25 figs., pis. 42-60. Brodi . Wn i hm t 1959. rerritorialit) in Rana clamitans. Herpetologica 15(3): 1 40. Bi doi 1 1 . I . S 1899. Notes on batrachians of the Paraguayan ( haco, with observa- tions upen the breeding habits and development, especially with regard to Phyllomedusa hypochondrialis ( ope \lso a descrip- tion of a new species Quart, Join Micro Sci >2 305-333, pis 28 J2 D III MAN, Wll 1 [AM 1 1956. Ih; frogs of the hylid genus Phrynohyas I itzinger, 1843. Misc. Publ. Mus Zool. t ni\ Mich. No. 9h 1-47, 10 figs., 4 maps. 6 pis. 1960. \ distributional stud) of the amphibians of the Isthmus of Tehuantepec, Mexico, Univ, Kansas Publ. Mus. Nat. Hist. 13: 19-72. 1966. Aggressive behavior in dendrobatid frogs. Herpetologica 22(3): 217-221. 2 figs 1 I RNANDEZ, K \l I . WD MlOl El I IKS \M>I / 1921. Sobre la biologia v reproduccion de algunos batracios amen tinos. I. Cystignathidae. An. Soc. Cient. Argentina 91: 97-140. 4 figs.. 3 pis. 1969 BREEDING BEHAVIOR 13 Gallardo, Jose M. 1958. Observaciones sobre el comportamiento de algunos batracios argentinos. Ciencia e Investigacion 14(7): 291-302. 1964. Consideraciones sobre Leptodactylus ocellatus (L.) (Amphibia Anura y especies aliadas. Physis 24(68): 374-384. 3 pis. Gans, Carl 1960. Notes on a herpetological collecting trip through the south- eastern lowlands of Bolivia. Ann. Carneg. Mus. 35, Art. 13: 283-314. Hemming, Francis (ed.) 1958. Opinion 520. Suppression under the Plenary Powers of the specific name tibiatrix Laurenti, 1768. as published in the com- bination Hyla tibiatrix, and of the generic name Acrodytcs Fit- zinger, 1843, and interpretation of the same Powers of the nominal species Rana venulosa Laurenti, 1768 (Class Am- phibia). Opin. Declar. Intl. Comm. Zool. Nomen. 19: 169-200. Hudson, William H. 1892. The Naturalist in La Plata. London, 394 pp. Ihering, Hermann von 1886. On the oviposition in Phyllomedusa iheringii. Ann. Mag. Nat. Hist. 17: 461-463, 1 fig. Jimenez, de la Espada, Marcos 1875. Vertebrados del viaje al Pacifico, Batracios. Madrid, 208 pp., 6 pis. Kenny, Julian S. 1966. Nest building in Phyllomedusa trinitatis Mertens. Carib. J. Sci. 6(1-2): 15-22, 6 figs. Lutz, Adolpho, and Bertha Lutz 1939. Notes on the genus Phyllomedusa Wagler. Ann. Acad. Bras. Sciencias 11(3): 219-263, 8 pis. Lutz, Bertha 1960. Fighting behavior and an incipient notion of territory in male tree frogs. Copeia 1960 ( 1 ) : 61-63, 4 figs. Lynch, W. Gardner 1959. Some reptiles and amphibians from Trinidad. Herpetologica 15(3): 1 13-117. Mayorga, Horacio 1967. Informe sobre una coleccion de anfibios (Salientia) procedentes de Nicaragua. Carib. J. Sci. 7( 1-2) : 69-77. Netting, Graham M. 1930. The systematic status and breeding habits of Eupemphix trini- tatis Boulenger. Ann. Carnegie Mus. 19(4): 249-254. Noble, Gladwin Kingsley 1927. The value of life history data in the study of the evolution of the Amphibia. Ann. N. Y. Acad. Sci. 30: 31-128. 30 figs., pi. 9. 14 BREVIORA NO. 321 PORTI R. kl NNF.TH K 1962 Mating calls and noteworthy collections of some Mexican amphibians. Herpetologica 18(3): 165-171. SI \ MIS, ()\\ I N I960. Sonic aspects of the behavior and of the territory of a dendro- batid frog. Prostherapis trinitatis. Ecology Hi It: 107-115. 6 figs. 1462. Apparent territorialism in Leptodactylus insularum Barbour. Herpetologica 18(3): 212-214. Sl I .INI (.1 K. 1 I tiNHVKD 1402. An annotated list of batrachians and reptiles collected in the vicinity of La Guaira, Venezuela, \«. nh descriptions of two new species of snakes I'roc. U. S. Nat. Mus. 21: 179-192, figs i J3 Iim. I Kl ni RICK H. I*>54. Social .mgiessixencss in .in amphibian. Science 120(3 108): 140-141. ZWEIFEL, Ki< HVKD ( i 1964 Life hisioi\ of Phrynohyas venulosa (Salientia: Hylidae) in Panama* ( opeia |,>',,l ' »' 201-208. (Received 5 March 1464.) BREVIORA Museum of Commpsir&itlve Zoology Cambridge, Mass. 10 June, 1969 Number 322 THE FOSSIL RECORD OF AMPHIUMID SALAMANDERS' Richard Estes Abstract. The first pre-Pleistocene records of amphiumid salamanders are described here. Proamphiwna cretacea, n. gen., n. sp., from the late Cretaceous Hell Creek Formation of Montana, differs from Amphiuma in having less well-developed vertebral muscle crests, basapophyses, and neural spines. The species is represented only by vertebrae. Amphiuma jepseni, n. sp., from the late Paleocene Polecat Bench Formation, Wyoming, is the earliest record of the modern genus. Its vertebrae do not differ from those of Recent Amphiuma, but its skull lacks the facial elongation char- acteristic of the modern species. It is represented by vertebrae and by a partial skull. Widespread transgressive seas of the Cretaceous, by providing extensive coastal plains and abundant shallow-water riparian habitat, may have been a major factor in the evolution of amphiumids and other elongated, aquatic salamanders. INTRODUCTION No previous fossil record for the salamander family Amphiu- midae has been described, except for two Pleistocene records, one reliable, the other spurious. Auffenberg and Goin (1959) and Brattstrom (1953) have correctly ascribed to Amphiuma speci- mens from Florida Pleistocene localities. An unusual zoogeographic record was described for the Pleistocene of Germany by Brunner (1956), as a new species, Amphiuma norica. This specimen may be the parasphenoid of a teleost; it bears no resemblance to that of Amphiuma, and there is no basis for the identification (Fig. 3g-h). Study of a North American Cretaceous floodplain herpetofauna (Estes, 1964) has disclosed ancestors of some lower vertebrates now living only on the Gulf Coastal Plain of North America. The 1 Fossil vertebrates from the late Cretaceous Hell Creek Formation, Montana: Contribution No. 3. 2 BR1VIORA No. 322 fauna of the latter region today differs in main ways from the ancient floodplain fauna, hut hoth share the presence of the fishes Amia, Lepisosteus, Acipenser, a polyodontid, sirenid salamanders. crocodiles, the turtle Trionyx, and alligators. These animals toda\ are all relicts of forms once widely distributed on fioodplains oi late Cretaceous epicontinental seas, the relict drainage of which is the Mississippi River system. The description here of another Gulf Coast element in the fossil fauna <^ the late C "retaceous of Mon- tana and Paleocene of Wyoming is further indication of the relict nature and former wide distribution of the (iulf Coast forms noted above. ACKNOWLEDGMENTS I am grateful to Dr. Francis Rose for helpful suggestions. Dr. Glenn Jepsen has provided the Silver Coulee specimens, which are described here in advance o\ a collaborative paper on Silver Coulee lower vertebrates, and 1 take pleasure in naming the new species for him. Collection of the Princeton material was supported by the Scott Fund. Princeton University. This research has been sup- ported in part b) National Science Foundation Grant GB-7176. I he figures are In Mr. l.as/lo Meszoly. FAMILY AMPHIUMIDAE Proamphiuma cretacea, n. gen., n. sp. (Figs. 1, 2) Holotype. M( / 3504, vertebra. Paratypes. MCZ 3506-3509, 3627-3636, vertebrae: 3505. 3637, atlantes. Locality. Bug Creek Anthills ( BCA), NO feet below the Tullock Formation in the west half oi Section 9, T 22 N, R 43 F, McCone County, Montana; collected bv A. D. Lewis and party. Horizon. Hell Creek Formation. Age. Fate Cretaceous. Diagnosis. An amphiumid with neural arch of atlas horizontal rather than dorsally-pointing posteriorly; vertebrae relatively nar- rower, more constricted medially, and with muscle crests, basa- pophyses, and neural spines less prominent than in Recent Am- phiuma of comparable size; shares with Amphiwna described below the relatively more constricted centrum and neural arch. Description. Centra amphicoelous, cotyles teardrop-shaped with thin internal coating of calcified cartilage; subcentral keel promi- nent or low, with or without channels for the segmental blood 1969 AMPHIUMID SALAMANDERS vessels; prominent or flattened anterior basapophyses usually pro- jecting beyond anterior centrum margin; indications of two closely- appresscd rib-bearers appearing on the four anterior vertebrae, the 5mm Fig. 1. Proamphiuma cretacea, n. gen., n. sp.: a, left lateral, b, an- terior, c, posterior, d, dorsal, and e, ventral views of holotype vertebra, MCZ 3504; /, anterior, g, posterior, /;, dorsal, i, ventral, and /', right lateral views of atlas, MCZ 3505. HRl VIORA No. 322 others with only one; ventral lamina of rib-bearer present but ap- parently not well developed anteriorly; zygapophyses anteroposteri- orly elongate, prominent, widely separated; posterior zygapophyses with keeled crests dorsally, extending forward to level of root of transverse process, these crests more medial in presumed anterior vertebrae and more lateral in presumed posterior vertebrae; neural spme relatively low, but prominent, thin and keeled, squared-off in lateral view; posterior border of spine slightly forked, often in three- pronged fashion. Atlantes with rounded anterior cotyles and prom- inent intercotylar process; centrum short; neural arch relatively high and blunt, with dorsal surface horizontal; neural spine only a faint ridge. Discussion. The vertebrae are distinctively amphiumid in the combination o\ anterior basapophyses. subecntral keel, and pos- terior zygapophyses with prominent distinctive crests on their dorsal Fig. 2. Proamphiuma cretacea, n. gen., n. sp.: (/. MCZ 3507. right lateral view of trunk vertebra; /'. MCZ 3508. righf lateral view of anterior trunk vertebra; c, MCZ 3506, dorsal view of trunk vertebra; 2. (3?)-l-l. John Packer per A. Loveridge. Pheretima morrisi ( Beddard, 1892) Three oi the tour specimens of this species previously available (Sims. 1964) were from garden soil. OC \l RODRI1 ID \l Green Mountain. 3d August, 1962, 2-0-0. John Parker per \ I overidge. Setae, closelv paired. Gizzards, probabl) lacking. Calciferous glands, long, seemingl) extending through ix-\ deeply constricted h\ 9 10. opening into the gut posteriorly in \, the anterior half more firm. Prostates, one pair. \erv long, external apertures unrecog- nizable. Spermathecae, non<: found. rhese worms (cfi. 1mm thick) were so softened that further information was unobtainable. No Other ocncroJnlc is known to have glands of the kind indicated above. Although reference to any known genus is impossible, the worms obviousl) were ocnero- drile. Remarks. The pheretimas are widely distributed anthropochores. I hey almost certainly were brought from the Orient to the island, directly or indirectly, by man. Any lumbricids on the island probably were brought, also by man. from England. Evidence to be presented elsewhere suggests that the pheretimas may be better adapted than the lumbricids to the climates o\' tropical islands such as Ascension. /'. californica and hawayana have been collected on St. Helena, the nearest land. /'. morrisi was once recorded from St. Helena but was not present in any of the various collections which were made on that island b\ Loveridge and others during the last twenty scars. 1969 EARTHWORMS OF TWO OCEANIC ISLANDS 3 REFERENCE Sims, R. W. 1964. Oligochaeta from Ascension Island and Sierra Leone including records of Pheretima and new species of Dichogaster. Ann. Mag. Nat. Hist.. (13) 7: 107-113. THE JUAN FERNANDEZ ISLANDS The Juan Fernandez islands, all small, are three: Mas a tierra (36 square miles), Mas Afuera (33 square miles), and Santa Clara. The first is inhabited. The islands were discovered by the man for whom they are named in 1563. The first settlement to become permanent was in 1877. Prior to Dr. Y. Kondo's visit, four species had been recorded from Juan Fernandez: Allolobophora caliginosa (Savigny, 1826), Dendrobaena rubida (Savigny, 1826), Eiseniella tetraedra (Savigny, 1826), Eukerria saltensis (Beddard, 1895). In the ab- sence of any indication to the contrary, all presumably were found on Mas a tierra. Collections made by Dr. Kondo on Mas a tierra, enable the listing of three more species. LUMBRIC1DAE Allolobophora Eisen, 1874 A llolobophora tuberculata Eisen, 1 894 Quebrada, Portozuela East, 200m, 2.xi.l965, 4 (+25?) -5-2. No. 31, 11. xi. 1966, 5-0-1. A. tuberculata is one of at least four species long retained in a classical congeries known as A. caliginosa. Worms previously re- corded from Juan Fernandez as A . caliginosa may well have been A. tuberculata. Dendrobaena Eisen, 1874 Dendrobaena octaedra (Savigny, 1826) No. 31, 11. xi. 1966, 1-0-2. Dendrobaena sp. Asondalan, 29. x. 1966, 1-0-0. This small juvenile is not of D. octaedra, but its condition was too poor to permit a positive identification, and it may be of D. rubida (Savigny, 1826), previously recorded from the island. breviora No. 323 Octolasion Oerley, 1 885 Octolasion tyrtaeum (Savigny, 1826) No. 31, 11. xi. 1966. 2-0-1. MON1LIGASTRIDAE Drawida Michaelsen, 19(H) Drawida bahamensis (Beddard, 1892) No. 31. 11. xi. 1966. 0-1-1. The left copulatory chamber, of one of the posterior amputees. is everted and the penis is protruded. Apertures of the other three chambers are tightly closed. Gizzards, in \ii-\iv ( 1 ). in xiii-xiv ( I ). Ovisacs, confined to xii. DISCUSSION Mas a tierra is only two square miles smaller than Ascension. Both islands are in the Southern Hemisphere. The Atlantic island has a known megadrile fauna of live species, and the Pacific island has seven: each island has one ocnerodrile. E. \altcn\i\ originated in southern South America, but in which part is unknown. The Ascension ocnerodrile could have evolved m South America or even in Africa. Ascension pheretimas. like the San Juan monili- v:astrid, originalK were from the Orient, l.umbricids. of course, came from Europe. Each species of the two islands is more or less widely distributed throughout the world. Each very probably was brought to the islands accidentally by man. Lumbricids. so far as can be indi- cated b\ the samples, seem to be dominant on Mas a tierra. Megascolecids. according to a smaller sample, seem to be dominant on Ascension. The Juan Fernandez Islands are well south of the tropics. Ascension well within them. The dilTerence in dominance may. then, be due to climatic factors. Nevertheless, the pheretimas. like the lumbricids. probably are from the Temperate /one of the Northern Hemisphere. ( Received 4 March 1969.) BREVIORA Muscimol of Comparative Zoology Cambridge, Mass. 15 September, 1969 Number 324 POLYMORPHISM AND EVOLUTION OF THE HISPANIOLAN SNAKE GENUS UROMACER (COLUBRIDAE) Henry S. Horn1 Abstract. Four species of the endemic genus Uromacer are recognized: catesbyi, fremitus, oxyrhynchus, and dorsalis. All save dorsalis show a re- markably discrete di- or trimorphism in a color pattern that is apparently cryptic. The adaptive significance of this polymorphism is discussed using an intuitive model that draws on the theory of Levins. The species are linearly ordered on the basis of body shape. Surprisingly, the stomachs of the stoutest species contain arboreal lizards and frogs, while the slimmer species take progressively more terrestrial lizards. The linear order suggests a pattern of geographic speciation occurring at the time when the northern and southern portions of Hispaniola were separate islands. Thus the pattern is consistent with patterns of evolution and migration proposed for other Hispaniolan reptiles by Williams and others. Speculations about the origin of dorsalis, endemic to He de la Gonave, may explain why the fauna of that islet is a curious mixture of species characteristic of the northern and southern portions of Hispaniola. INTRODUCTION The snakes of the genus Uromacer are endemic to Hispaniola and the surrounding islets. Some years ago A. Stanley Rand dis- covered a color polymorphism in several local populations of Uromacer oxyrhynchus. Examination of the other species has shown that several of them are also polymorphic. Since the same morphs of different species are superficially more similar than are different morphs of the same species, this polymorphism sometimes results in the misidentification of museum specimens. Conse- quently, the polymorphism has obscured the pattern of distribution 'Department of Biology, Princeton University, Princeton, New Jersey. 2 BREVIORA No. 324 of the Forms of I 'romacer, making it difficult to relate their distribu- tion to that o\' other Hispaniolan reptiles. The recent concentration on the herpctologv of Hispaniola at the Museum of Comparative Zoolog) (MCZ) has provided large, synchronous collections of U romacer from several localities. The geographical coverage of even these large collections is not yet extensive enough to warrant a definitive discussion o\' intraspeciflc variation. However, the available collections allow me to assess the validity oi the described species of I romacer, to suggest the pattern of their relationships, to document their striking poly- morphism in some detail, and to comment on some interesting evolutionary problems posed by the genus. Though I have studied only preser\ed material. I have made inferences about the eeolog\ oi the species from their stomach contents. However, I must leave several questions unanswered because I lack field knowledge of Hispaniola. particularly o\ seasonal changes during the year. Where \1( / material was insufficient, I have examined speci- mens from the American Museum ol Natural Histor\ ( AMNH), the British Museum (Natural Histor\ ) (BMNH), the Field Mu- seum o\ Natural Histor\ (I \1NII). the Yale Peabod) Museum (YPM), the Albert Schwartz Field Series ( ASFS). and the United States National Museum (USNM). I am grateful to the curators o\ these collections for allowing me to examine specimens in their charge. I thank A. Stanle\ Kand and Ernest E. Williams tor discussion and criticism throughout this stud\. Dr. Rand introduced me to {J romacer, and his unpublished notes on their polymorphism were the skirting point for m\ work. Dr. Williams sponsored part of this stud) as m\ undergraduate diesis at Harvard College in 1962. \lv ideas ^w polymorphism developed from discussions with Richard Levins and Robert II Mac Arthur. For criticisms o\' several manuscripts. I am indebted to I nomas H. Irazzetta. Eliza- beth G. Horn. Thomas W. Schoener, Albert Schwartz, and George (i. Simpson. Dr. Schwartz kindlv compiled the fauna! list for lie dc la Gonave. MATERIAL AND METHODS Cochran (1941 ) recognized six species of V romacer: catesbyi, scandax, frenatus, wetmorei, dorsalis, and oxyrhynchus. There is reason to doubt the specific distinctness o\ scandal and wetmorei, which Cochran recognized as respectively allied to catesbyi and frenatus. For reasons given in the Appendix. I consider scandax 1969 EVOLUTION OF UROMACER o <\J g i CD N »<]oo o < o ■9 . < o to & E ^ £ 4 E ^ -* o o S o S to c o CD a> c o > .2 2 .S> o Q_ O ,'.•:■ »< o u a o _o c o O 'c C CO 'VvJ i'_v' 5 4 bki \ [OR \ No. 324 to be identical with catesbyi, and wetmorei to be a subspecies of frenatus. The remaining species are easily separated by the diagnostic characters ot Cochran's key: snout length relative to eye diameter, ventral scale count, and midbody scale rows. The ranges of snout length and ventral scale count overlap between species, but these characters arc usually sufficient tor tentative identification ot" single specimens. I he number of scale rows at midbody is constant in each species, but is diagnostic only for oxyrhynchus. The subcaudal scale count for each species has a laree range, and though there may be significant differences be- tween the means oi different populations, it is seldom useful in identifying a single specimen. Similarly, the presence or absence of an anterior loreal ma\ be used to infer relationships among populations, but it is useless as a diagnostic character. Stomach contents of the MCZ and YPM specimens were re- moved and identified. I he snakes had fed primarily 00 Anolis whose ecolog) is rclati\el\ well known, so inferences about the ecolog) of Uromacer ma) be made on the basis of these stomach contents. Schwartz (in litt. ) and Merteiis (1939) have observed that freshly caught Uromacer often swallow the cohabitants of a collecting bag. but even those stomach contents that were not eaten by free snakes will retleet the syntopic fauna in which the snakes were collected. To minimize this difficulty, those stomach contents that were undigested were ignored in analysis. Where different color morphs are described for a single species of Uromacer, these morphs are entirely distinct, with no inter- mediates except between the gra\ and the striped gray forms. There was never any hesitation in assigning any well-preserved specimen to one o\ three alternative morphs: green, green above and below, lighter below; gray-green, green above, mottled gray below; or gra) . gray above and below. A detailed description of each morph and its variations in each species is given in the Appendix. RESULTS AND INTERPRETATIONS The ranges of the species of Uromacer are mapped in Figure 1 and listed in Table 1 . External metric characters are listed in Table 2, and there are- photographs of the several species in Figure 2. The species may be linearly ordered, as in the table, on the basis of characters that are correlated with slimming and lengthening the body. U. catesbyi is typically colubrid in shape, with a short, blunt snout, low 969 EVOLUTION OF UROMACER TABLE 1 Ranges of the species of Uromacer Species Southwest and Barahona Peninsulas Port-au-Prince and vicinity N . and E. lie de la Gondve Dominican Republic cateshyi frenatus oxyrhynchus dorsalis TABLE 2 Metric morphology of the species of Uromacer Snout length relative Species Number to eye Ventral Subcaudal examined diameter scales scales Scale rows at Anterior midbody loreal catesbyi 254 2.0 167 ±0.7 183 ±1.0 17 — frenatus 102 2.6 187 ±0.5 204 ±1.9 oxyrhynchus 129 3.0 200 ±0.5 199 ±1.1 17 19 present in 25 dorsalis 84 2.9 200 ±1.2 210 ±2.2 17 + Entries are the means for each character ± the standard error of the mean. The standard error of the mean snout length (±0.01) was, in all cases, less than the reading error of each measurement (±0.2). Numbers separated by * are significantly different at the .05 level by a one-tailed t test (Snedecor, 1956), for snout length, ventral count, and subcaudal count, or a binomial test (Siegel, 1956), for scale rows at midbody and anterior loreal. HRI \I()K\ No. 324 II ilt-HIIII iiiiifui ]2 IMIIIlit !3 IMillllt llll 4 I 1 969 EVOLUTION OF UROMACER Figure 2. a, Uromacer catesbyi MCZ 57764 nr. Carpintero, Dominican Republic; b, U. frenatus MCZ 65530 Perrine nr. Jeremie, Haiti; c, U. oxyr- hynchus MCZ 62713 Bombardopolis, Haiti; d, V. dorsalis MCZ 80815 Pointe-a-Raquettes, He de la Gonave, Haiti. 3 imi \ I or \ No. 324 ventral and subcaudal counts, and a relatively thick bod\ covered with rhomboidal scales that arc nearl) square. U. oxyrhynchui and dorsalis arc shaped more like typical tree and vine snakes. with long, pointed snouts, high ventral and subcaudal counts, and more slender bodies covered with obliquely rhomboidal scales. U. frenatus is intermediate between these extremes. The stomach contents of I romacer are listed in Tables 3 and 3a. In Table 3 the stomach contents of the snakes are pooled within TABLE 3 Habitat of food taken by the species of I romacer Habitat catesbyi fremitus oxyrhynchtu dorsalis Trees I ree trunks and bushes >» ( n.iss and open areas ( i round and rocks 3 1 2 5 II 4 ) 4 II 4 4 10 TABLE 3a Stomach c intents ol the species of i 'romacer Habitat and species eaten <\; frenatus oxyrhynchus ilor\tili\ rREES ll\ln puh hrilineata 1 do/ninh ensis I I noli\ i hlorocyanui 5 1 2 ( oelestinus 1 3 1 Rl K TRUNKS \ Bl sill s A nulls cy botes 4 4 8 3 hrc\ iros'i s 1 distil hits 5 1 3 (.K \Ss,V Ol'l N \RI \S A nulis semilineatus i 3 olssoni 1 11 (.ROUND& ROCKS Ameiva taeniura 1 : 1 chrysolaema ft Leiocephalus melanochlorus i vinculum 3 schreibersii : Sphaerodactylus sp. 1 1969 EVOLUTION OF UROMACER 9 common habitats of the food species; the habitat groupings were made by referring to Mertens (1939), Williams and Rand (1961), and Rand (1962). Since the groupings are only generalizations, and the samples are small and heterogeneous, the data do not fully characterize the diets of the species of Uromacer. However, the differences among the diets are so striking, when compared to the uniformity of diets within each species, that crude relative characterizations can be made with confidence. U . catesbyi takes arboreal food almost exclusively, and dorsalis feeds primarily on terrestrial food, whereas /. jrenatus and oxyrhyn- chus are intermediate and more similar to each other than either is to catesbyi. The tendency for the colubrid-like catesbyi to take arboreal prey, and for the longer-snouted species to take pro- gressively greater proportions of terrestrial prey, is the opposite of expectations based on the morphology of the species (e.g. Mertens, 1939, who suggest that oxyrhynchus is the most arboreal species of Uromacer) . Polymorphism. Of the four species of Uromacer, only the gray dorsalis is not demonstrably polymorphic in at least one locality (see Table 4). Green U. catesbyi occur throughout the range of the species, but with a pattern unlike the corresponding morph TABLE 4 Distribution of the color morphs of the species of Uromacer Species Green Gray-green Gray Gray (striped) catesbyi ENTIRE RANGE Miragoane /. jrenatus He a Vache mainland He a Vache /. wetmorei Barahona Isla Beata oxyrhynchus Haiti & Haiti ENTIRE RANGE Eastern D.R. dorsalis ^ENTIRE RANGE of any other species; catesbyi is polymorphic, with a gray form, near Miragoane, in the middle of the southwest peninsula of Haiti. U. f. jrenatus is monomorphic gray on the mainland, and poly- morphic gray and gray-green on lie a Vache. The gray form on lie a Vache resembles that of dorsalis more than that of mainland jrenatus or oxyrhynchus. U . j. wetmorei on Barahona may be polymorphic, but the few poorly-preserved specimens presently available from scattered localities are insufficient evidence. 10 BR] V ioka No. 324 V. oxyrhynchus is apparently monomorphic green through much of the Dominican Republic, but it is dimorphic green and gray- green throughout the Haitian portion of its range. In scattered Haitian localities it is trimorphic. green, gray-green, and gray. U. oxyrhynchus may in fact be trimorphic throughout the Haitian portion of its range, perhaps in parts of the Dominican Republic as well. The gray morphs are rare enough to be found only in large, synchronous collections from Haiti; no such collections have been made in the Dominican Republic. The discreteness ol the morphs in all the polymorphic species, and the lack of intermediates, suggests that the mechanism con- trolling the polymorphism is an allelic switch at a single genetic locus. In the trimorphic oxyrhynchus, die distribution of the morphs is consistent with the interpretation that the green and the gray tonus are homozygotes and that the commoner gray-green form is the heterozygote; i.e. in all the localities from which both green and gra\ specimens have been taken. gra\ -green specimens have been taken as well. I he distribution ol the color forms among the localities of the polymorphic species and among the species of the genus suggests that the various tonus are cryptic tor different surroundings. U. catesbyi is green throughout its range, darker m the western por- tions ol Hispaniola than in the eastern, with some gray individuals on the southwest peninsula in the Miragoanc area. U. frenatUS is gray 0:1 the southwest peninsula and polymorphic on lie a \ ache, while its ecological counterpart, oxyrhynchus, has green, gray- green, and gra\ morphs in Haiti, but a preponderance o\ green morphs in the Dominican Republic (see I able 5 ). I hese distribu- tions correspond roughh to the pattern of humidit) and vegetation on Hispaniola. as described b\ Wetmorc and Swales ( 1931 : 2-7). TABLE 5 Geographical distribution of the color morphs of frenatUS and oxyrhynchus S/>< i ;, \ 1 rea Gret n ( ,r,r -gn • 1, Gray /. fremitus lie .1 \ ache 15 22 f. frenatiis Haitian mainland 2 56 f. wetmorei Barahona 4 1 oxyrhynchus Haiti 47 48 4 oxyrhynchus Dominican Republic 17 4 1969 EVOLUTION OF UROMACER 1 1 The grayer and darker (i.e. less bright green) forms are associated with the drier, western portion of the island. The presence of the gray-green morph of fremitus on He a Vache and the brighter green of the catesbyi from the islet are correlated with the presence of denser woodland on the islet than on the adjacent mainland. This correlation of the distribution of the color morphs with foliage density suggests cryptic coloration as a possible function of the different morphs. The two most strictly monomorphic species, catesbyi and dorsalis, are characterized by the most restricted diets. The green catesbyi takes only arboreal prey, while the gray dorsalis takes predominantly terrestrial prey. This additional correlation of color pattern with ecology suggests again that the coloration of Vromacer is cryptic, concealing the snake as it stalks agile prey. Polymorphism in a cryptic pattern of coloration poses an interest- ing evolutionary question. If a specific color pattern is indeed cryptic, why is more than one pattern present at all? The theory of polymorphism provides some likely answers and suggests the kinds of data that should be sought to choose between alternatives. Levins (1962-6; Levins and MacArthur, 1966) has examined the adaptive significance of polymorphism, using an abstract mathe- matical model of extreme generality. I shall develop a much simpler model to examine polymorphism in the special case of cryptic coloration, but the intellectual basis and results are the same as those of Levins. This model is almost absurdly simple, but the concepts illustrated are quite general and are not easily apparent without the model. The environment is a checkerboard with black and white squares and the cryptic animals are checkers. Later I shall intro- duce reproduction and "fitness" into the model, but first let us look at crypticity alone. If we randomly place black or white checkers on the squares of the checkerboard, roughly half the black checkers will fall on black squares and half the white checkers, on white. Thus half the population of either black or white checkers will be cryptic. However, if we throw gray checkers on the checkerboard, none of them will be cryptic, unless, of course, the checkers are very large and the squares of the checker- board are so tiny that it looks like the gray of a halftone illustra- tion. Thus if two alternative morphs are to be cryptic, the environ- ment must be a mixture of large patches of different kinds, each with an extreme morph that is more cryptic than an intermediate morph. 12 BREN IORA No. 324 I assume that "adaptation" is achieved through natural selection, which favors that individual whose offspring form the greatest proportion o\' breeders in the next generation. Correspondingly the "fitness" of an individual is defined as the Dumber o\' his offspring that breed in the next generation, and the individual with the greatest fitness in a given environment is the "'best adapted*' to that environment. We inns allow each checker to reproduce an average of X offspring in each generation; in each reproductive period we throw checkers onto the board and give each checker X+d oil- spring ii it is cryptic Inn only X d offspring if it is not cryptic. Black checkers have black offspring; white checkers have white offspring. We define "polymorphic" checkers to have either black or white offspring with a 50-50 chance, whether the phcnolvpc of the "polymorphic" checker is black or white. I he genetic or developmental basis of this polymorphism need not be specified, so I shall call it '"sibling polymorphism," since it requires that the offspring of a given morph maj include both morphs. If the checkerboard has an equal number of black and white squares, then the offspring o\ black, while, or poKmorphic checkers have an equal chance of being crvptic or non-cr\ptic. and the fitness dt each type is an average of X • d and \ d. or simpK X. (You can easil) show that it the environment is an uneven mixture of black and white squares, the morph that is cryptic in the commoner square has a higher fitness than the other morph or the poly- morph. ) I hus, it the environment is a spatial mixture of large patches of two types, a monomorphic population is better adapted than a poKmorphic population, or at least as well adapted. If, instead o\ throwing the checkers onto the board randomly. we allow each ot the checkers to choose and reproduce in its appropriate habitat, then the white squares will be occupied pre- dominant!) b\ white checkers and the black squares by black checkers. On sampling a large and heterogeneous area, we then obtain both morphs. and the population appears to be polymorphic. However, the two morphs could represent two species, rather than a poKmorphic species. In tact, the average fitness of individuals of the two separate species would be higher than that of a poly- morphic species, unless the polymorphism continually adjusted the ratio of the morphs to match the proportions of the corre- sponding patches in the environment. Since this polymorphism ma\ result from sampling a heterogeneous environment and does 1969 EVOLUTION OF UROMACER 13 not require that a given individual be capable of having poly- morphic offspring, I shall call it "sampling polymorphism'1 as dis- tinct from "sibling polymorphism." (Under certain restricted con- ditions the two kinds of polymorphism are convergent, and sam- pling polymorphism may evolve a genetic basis. See Levins, 1965, for a discussion.) Now instead of throwing the checkers onto a checkerboard, we throw them alternately onto a white board or a black board for each generation. That is, the environment is a temporal mixture of different habitats, rather than a spatial mixture. The black and white checkers are alternately cryptic and non-cryptic. Each individual in a cryptic generation has X+d offspring, each of whom has X— d offspring in the next generation; thus the average fitness of black or white checkers over two generations is (X-fd)(X— d) = X-— d-. On the other hand, the "polymorphic" checkers have an equal number of cryptic and non-cryptic offspring in each generation and their average fitness is V2 (X+d) + '/2 (X— d) =X in each generation; then the average fitness of polymorphic check- ers over two generations is X2, which is always greater than X2— d-. If the black and white environments alternate several times within each generation, fitness is averaged over the two environments for each generation, and the results are the same as in the case of spatial heterogeneity. However, sibling polymorph- ism yields a higher average fitness than monomorphism when the environment varies drastically with a period of the same order of magnitude as the time between generations. What does the checkerboard model predict about Uromacer! If the polymorphism of Uromacer is simply a sampling poly- morphism, then the different morphs should be associated with distinctly different habitats or microhabitats. The habitats might be different types of vegetation, in each of which one of the morphs is most cryptic. If the polymorphism of Uromacer is a sibling polymorphism, the alternative morphs should still be cryptic in distinctly different habitats, but the habitats will alternate in an inescapable temporal pattern, e.g. wet-dry season. Therefore, if the polymorphism is a genetic or developmental adaptation to a temporally heterogeneous environment, the different morphs will often be found in habitats in which they are not cryptic. Documenting the adaptive significance of polymorphism in Uromacer is a difficult, long-term project. It is first of all a field, not a museum project. It is necessary to define the habitats in 14 hkiaiora No. 324 which each of the morphs is most cryptic. Then the alternatives ot genetic and sampling polymorphism must he separated by deter- mining the distribution of the morphs among the several habitats. over a time equal to several generations oii Uromacer. As interest- ing as the problem is. it will probabl) not be quickly solved, per- haps not even quickly undertaken, since the required study is longer than the lifetime of the average research grant. Relationships within the genus. I here has been no previous dis- cussion oi the relationships oi the species oi Uromacer. It is at once tempting to suggest that the morphological and dietarv series described corresponds to an evolutionary series as well; that catesbyi is the most primitive of the Uromacer, oxyrhyn- chus and dorsalis the most recently derived, and that frenatus represents an intermediate step. When the present ranges o^i the species, the physiographic historj of Hispaniola, and the mechan- isms ol geographic speeiation are taken into account, this sugges- tion is strengthened. During the Pleistocene. Hispaniola was divided by an arm of the sea into two large islands, a northern island. Haiti north o\ the Cul-de-SaC Plain and the Dominican Republic exclusive of the Barahona peninsula, and a southern island, the southwest peninsula o\ Haiti and the Barahona peninsula ( Schuchcrt. 1935; Weyl, L966; Williams. 1961). I he ancestral colubrid ( Maglio, in prep., suueests something similar to Alsophis) had given rise to a catesbyi-like form, to be called "proto-catesbyi," and to "proto-frenatus." autochthons of the northern and southern is- lands respectively. Proto-frenatus invaded the northern island, and as it came into contact with the well-established proto-catesbyi. the characteristics that distinguished the tWO species were inten- sified (character divergence o\ \la\r. 1963. after Darwin). By its increased divergence from proto-catesbyi. the invading proto- Erenatus gave rise to 'proto-oxyrhynchus.'" ancestor oi oxyrhyn- chlU and dorsalis. I he proposed sequence accounts for the presence in dorsalis of characters similar to both frenatus and o.wrhxnehus. The pro- genitor of dorsalis on Gonavc was presumably proto-oxyrhynchus. which would have retained similarities to proto-frenatus and thus to frenatus. while differentiating in the direction of o.xyrhynchus. Alternatively, dorsalis could have been derived directly from proto-frenatus. and its similarity to o.xyrhynchus could represent parallel evolution of the species. Lack of evidence relevant to these alternatives is in itself sufficient reason for provisionally call- ing dorsalis a separate species. 1969 EVOLUTION OF UROMACER 15 This evolutionary sequence is only an interpretive hypothesis and cannot be tested, but it embodies processes that have been suggested to account for the present distribution of other His- paniolan reptiles. The postulated distinction of proto-catesbyi and proto-frenatus, as the respective autochthons of the northern and southern islands, is paralleled by the distinction of Anolis coeles- tinus and Anolis chlorocyanus, of Anolis aliniger and Anolis singularis, and perhaps of the Anolis of the monticola group and the subspecies of Anolis ricordii (Thomas and Schwartz, 1967; Williams 1965a, 1965b). The subsequent invasion of the northern island by proto-frenatus is paralleled by the Anolis of the setni- lineatus group. Williams (1961) suggests that Anolis olssoni and A. semilineatus are indigenous to the northern and southern islands respectively and that their present distributions are the result of the relatively recent invasion of the northern land mass by semi- lineatus. Similarly, Schwartz ( 1967) suggests that Ameiva taeniura originated on the southern island and has since invaded the northern land mass. There are probably other patterns of radiation that are con- sistent with the present ranges of the species of Uromacer, and almost as simple in the changes of morphology that they would require. However, several conclusions will remain unchanged given such alternative patterns. U. catesbyi is clearly more differ- ent, in morphology and diet, from the other species than they are from one another. It is then significant that only catesbyi is broadly sympatric with the other species of the genus. U. frenatus, oxyrhynchus , and dorsalis are similar in morphology and may provisionally be considered allopatric ecological replacements. Explaining the origin of dorsalis on Gonave as more than a historical accident requires bold speculation in the absence of critical data. This speculation is justified, however, as it also suggests a possible derivation for the problematical fauna of Gonave. Present knowledge of the fauna of Gonave is scanty, but it is already apparent that the fauna is an odd mixture of species characteristic of the northern and southern islands of the Pleisto- cene (See Table 6). During part of the Pleistocene, Gonave was reduced to about one fourth of its present area (Butterlin, 1960). When the northern and southern islands were rejoined, Gonave may have been connected to the mainland by a spit at least 3 km wide to the north and 7 km wide to the south (Fig. 1; also Wood- ring et al., 1924), which would have allowed the fauna of both regions to invade Gonave. When Gonave became insular again, 16 KKl \ [ORA No. 324 those species whose presence had depended on continuing invasion would he eliminated, leaving only those best adapted to the restricted habitats of Gonave and those potential competitors that were most divergent. The geographic origin of the invading species would be irrelevant to their subsequent persistence. TABLE 6 Reptiles of Gonave and their areas of origin \/'ci /< \ Origin Gonatodi i albogularis Sphaerodat tylus < inereus fteliigt r i ci hranae Audits brevirostris (end. subsp i ( hlorot yanus i \ bott I olsoni singularis ( m luni ( hi nnlti 1 1 ocephalus vinculum (end subsp i North North South Reference Williams. 1965 a Williams. 1961 Williams. 1965a Ameiva < (try solai ma ( end. taeniura sub* ».) North South2 Schwai t/ & Klinikowski, 1966 v wartz, 1967 Diploglossus curtissi Amphisbaena gonavensis i end siihsp. ) South ' Ihomas. 1965 / yphlops capitulatus (end. pusillus sub sp.) North Richmond. 1964 Eph rates angulifer fordi I ropidophis haetianus Dromicus parvifrom Hypsirhynchus ferox laltris dorsalii Uromacer i atesbyi dorsalii < endemic I North - North 1 North and south under this heading refer to the northern and southern islands into which Hispaniola was periodical divided during the Pleistocene. See Fig. 1 . -' Island of origin of parent species, though invasion may have been from a population on the other island. 1969 EVOLUTION OF UROMACER 17 In particular, if proto-catesbyi, proto-frenatus, and proto- oxyrhynchus all reached peninsular Gonave, the ultimate per- sistence of the most divergent pair, and the elimination of the intermediate proto-frenatus is predictable. The dietary divergence of dorsalis and catesbyi confirms the role of interspecific competi- tion in the origin of dorsalis. U. dorsalis clearly differs from catesbyi in foraging habitat, if not in foraging behavior, to a greater extent than jrenatus and oxyrhynchus differ from catesbyi (Table 3). Interspecific competition may necessitate this increased divergence of dorsalis and catesbyi on Gonave, whereas less diver- gence insures the coexistence of two sympatric Uromacer on the more extensive and heterogeneous Haitian mainland. If the history above is wrong, Gonave deserves intensive study as an extraordinary example of an island colonized from two com- parable and equidistant sources. If the suggested history is cor- rect, then Gonave is a model of the faunal transformations of a continental island that when peninsular had the full complement of species of the adjacent mainland: while several closely related species could persist when immigration from the mainland was easy, fewer were able to persist after their populations became insular. Those that did persist on the island had to be strongly divergent ecologically. According to such a model, an interpreta- tion of the fauna of a continental island must emphasize the eco- logical compatibilities of the potentially resident species, rather than their relative vagilities and routes of invasion from the ad- jacent mainland. LITERATURE CITED BUTTERLIN, J. 1960. Geologie generate et regionale de la Republique d'Haiti. Travaux et Memoires de ITnstitut des Hautes Etudes de l'Amerique Latine, Paris, 6, 194 pp. Cochran, D. 1931. New reptiles from Beata Island, Dominican Republic. Proc. Biol. Soc. Washington. 44: 89-92. 1941. The herpetology of Hispaniola. Bull. U.S. Nat. Mus.. 177: 1-398. Dunn, E. R. 1 920. On the Haitian snakes of the genera Leimadophis and Uromacer. Proc. New England Zool. Club, 7: 37-44. Levins, R. 1962. Theory of fitness in a heterogeneous environment. I. The fitness set and adaptive function. Amer. Nat., 96: 361-373. IS hki \ IORA No. 324 1963 rheorj ol fitness in .1 heterogeneous environment. II Develop menial flexibility and niche selection, \nu-i Nat., *>7: 75-40. 1964. rheon of fitness in .1 heterogeneous environment. 111. ["he re- sponse to selection. J. rheoret. Biol . 7 224 240. I9h5 ITieory of litness in a heterogeneous environment l\ l he adaptive significance ol gene H iw I volution, IK: 635 638 1966. I he strategy of model building in population biolog) \mei Sci . •"> I: 421-431. I I \ INS. K . \M> R II M \< \KI III K I9hh. The maintenance of genetic polymorphism in .1 spatially hetero- geneous environment: variations on a theme by Howard 1 evene \nu-i Nal . 100 585 5J Mayr, I 1963 XnniKil Species and Evolution Cambridge, Mass., Belknap Press, Harvard I Diversity Press, 797 pp. Mertens, R 193 ' Herpetologische I rgebnisse einei reise nach der Insel Hispaniola, Westindien. \bh. Senckenberg N.iiuit Ges., 149: 1 84 Rand, \ S Il>(i2 Notes Dii Hispaniolan herpetology 5. I Ik- natural history ol three sympatric species of Anolis, Breviora, Mus ( omp. Zool . No 154 1-15 Rll HMOND, N H 1964 I he bliod snakes 1 / yphlopa 1 of Haiti with descriptions of three new species. Bre\ 0 a Mus I omp Zool., No. 202: 1-12. S< HI l III Kl . ( 1935 Historical Geology ol the ^ntillean-Caribbean Region. New > oik. John w iley & Sons. 81 1 pp S( HW VKI/. \ 1965 I Ik- Leiocephalus il acertilia, Iguanidae) of Hispaniola. I Leio- cephalus meianochlorus < ope. lourn. Ohio Herp, Soc., 5: 39-48. 1967. The Ameiva (Lacertilia, reiidae) ol Hispaniola. Ill Ameiva taeniura Cope. Bull. Mus. (omp Zool., 135 $45-375. Si HW \RI/. \ . \N|) K 1 Kl INIKOWSK1 1966 I Ik- Ameiva il acertilia, reiidae) of Hispaniola. II. Geographic variation in Ameiva chrysolaema ( ope Bull. Mus (omp. Zool., 133: 427-4$ Siegi 1 . S 1956 Nonparametric Statistics for the Behavioral Sciences. New York, McGraw-Hill, 312 pp. SM Dl ( ciR. (1 \V. 1956. Statistical Methods \pplieil to I uperiments in Agriculture and Biology. Vines, Iowa, Iowa state Univ. Press. 534 pp. Thomas. R. 1965. Two ne\s subspecies oj Amphisbaena (Amphisbaenia, Reptilia) from the Barahona Peninsula of Hispaniola Breviora, Mus Comp. Zool.. No. 215: 1-14 1969 EVOLUTION OF UROMACER 19 Thomas, R., and A. Schwartz 1967. The monticola group of the lizard genus Anolis in Hispaniola. Breviora, Mus. Comp. Zool.. No. 261: 1-27. Wetmore, A., and B. H. Swales 1931. The birds of Haiti and the Dominican Republic. Bull. U.S. Nat. Mus., 155: 1-483. Weyl, R. 1966. Geologie der Antillien. Beitrage zur Regionalen Geologie der Erde, Vol. 4, Berlin, Gebruder Borntraeger, 410 pp. Williams, E. E. 1961. Notes on Hispaniolan herpetology. 3. The evolution and rela- tionships of the Anolis semilineatus group. Breviora, Mus. Comp. Zool., No. 136: 1-8. 1965a. The species of Hispaniolan green anoles (Sauria, Iguanidae). Breviora, Mus. Comp. Zool., No. 227: 1-16. 1965b. Hispaniolan giant anoles (Sauria, Iguanidae): New data and a new subspecies. Breviora, Mus. Comp. Zool., No. 232: 1-7. Williams, E. E., and A. S. Rand 1961. Notes on Hispaniolan herpetology. 2. A review of the Anolis semilineatus group with a description of Anolis cochranoe, new species. Breviora, Mus. Comp. Zool., No. 135: 1-11. Woodring, W. P., J. S. Brown, and W. S. Burbank 1924. Geology of the Republic of Haiti. Rep. of Haiti Dept. of Public Works, Port-au-Prince, 631 pp. (Received 20 January 1969.) Appendix SYSTEMATICS The synonomy of each species cites only the original descrip- tion and additions to the synonomy of Cochran ( 1941 ). The data for the cited geographical range are given in Figure 1 . The extreme measurements of the snakes examined are given for each metric character; the standard error for each measurement is listed in Table 2. A list of all specimens examined, their localities and color morphs, and the stomach contents listed in Table 3 are filed with the specimens at the Museum of Comparative Zoology. Uromacer catesbyi (Schlegel) Dendrophis catesbyi Schlegel, 1837, Essai sur la physionomie des serpens, 2: 226 (type loc. "L'ile de St. Dominque"). Uromacer scandax Dunn, 1920, Proc. New England Zool. Club, 7: 43 (type loc. Tortuga Island). Jo BRKVIORA No. 324 Range. Hispaniola: the coastal area aorth of the Cul-de-Sac Plain, the southwest peninsula of Haiti, upland valleys of the in- terior of the island, lie a Yaehe. He de la Gonave, an J lie de la Tortue. It is sympatric with ever) other species in the genus. Diagnosis. Snout 2.0 (range: 1.7-2.7) times eye diameter; scale rows at midbody 17; ventrals lb7 (range: 160-180); subcaudals 183 (range: 163-205); anterior loreal absent. 254 specimens were examined: 14 W1NH. 7l> ASFS, 6 BMNil. 21 I'MNH, 126 MCZ. 8 USNM. Color pattern. Green morph: dark green above, light yellow- green below, the colors meeting on the 3rd scale row; head dark green above, cream below, a light line above the eye and a dark line from the snout to the end o\' the head i alter M< / 64800). A few specimens have a light lateral line on the 2nd and or 3rd scale row rhe dorsal dark green varies geographically, from a dark. brownish green in western Haiti to a \er\ light green in the eastern Dominican Republic. Specimens from lie a \ ache are somewhat lighter than those from the adjacent mainland. Graj morph: gray above, cream below, the colors meeting on the 3rd scale row; head gray-green above, cream below, a light line above the eve (after \i( / 66350). I his morph has been found in 5 specimens from the Miragoane area. Nomenclature. Dunn (1920) defines scandal as "like Uro- macer catesbyi but with 19-11 scale rows and 181 ventrals." Examining the holotype (USNM 59438), I count 17 scale rows at midbody and 180 ventrals, with \(> scales around the neck persisting to within 5 em o\ midbody. A count oi \() scale rows persists for some distance down the body in six o{ the 254 catesbyi examined: there are also three typical catesbyi from scattered localities with ventral counts of 177. 178. and 179. Two catesbyi typical in scale characters (NIC/. 37607, USNM 59920) have been taken from He de la I ortue. but larger collec- tions ma\ reveal an endemic subspecies oi catesbyi for which the name scandal would be appropriate. Uro\i\( ik prenatus (Gunther) Ahaetulla frenala Gunther, 1X65. Ann. Mag. Nat. Hist.. Scr. 3. 15: 94 (type 'oc. "no: known" ). Uromacer wetmorei Co:hran. 1931. Proc. Biol. Soc. Washington. 44: 91 ( type lot Isla Beata i Range. Southwest peninsula of Haiti east and north to the Cul-de-Sac Plain, lie a \ ache. Isla Beata. and the eastern coast 1969 EVOLUTION OF UROMACER 21 of Barahona Peninsula. Records outside this area have been based on misidentifications (Cochran, 1941). It is sympatric with catesbyi throughout its range and with oxyrhynchus at the base of the southwest peninsula. Diagnosis. Snout 2.6 (range: 2.1-3.0) times eye diameter; scale rows at midbody 17; ventrals 187 (range: 177-196); sub- caudals 204 (range: 185-212; but AMNH 51416 has 135 and its tail appears intact); anterior loreal absent. 102 specimens were examined: 4 AMNH, 39 ASFS, 2 BMNH, 46 MCZ, 11 USNM. Color pattern. Gray-green morph: Gray-green frenatus from He a Vache are dark blue (probably green in life) above, mottled gray to gray-brown below, the colors separated by a white line on the 2nd and 3rd scale rows from four head lengths down the neck to the vent; head blue (probably green in life) above, cream below, a light line above the eye, dorsal edge of upper labials black (after MCZ 37630). The posterior extent of the white line is variable; on some specimens it fades as far anteriorly as midbody. Gray- green /. wetmorei from Barahona are as above, but the white line is on the 4th and 5th scale rows from the snout to one head length behind the vent (after AMNH 51421). One specimen (MCZ 43806) is not mottled ventrally and may represent a green morph, but it has been so blackened by the preservative that it is impossible to infer its color in life. Gray morph: Mainland /. frenatus are finely mottled gray above and below; a trace of green on the top of the head, light green on upper labials, chin, and ventral neck (after MCZ 65530). Gray frenatus from He a Vache and the type specimen of /. wetmorei (USNM 83891, Isla Beata) are the same as above but with the 4th and 5th scale rows very light, thus giving the appearance of a wide, dark gray dorsal stripe (after MCZ 37629). Nomenclature. Cochran (1931) described wetmorei on the basis of a single specimen (USNM 83891 ) from Isla Beata, noting its close relationship to frenatus. I have re-identified 5 specimens from the Barahona mainland (MCZ 43806, AMNH 51416, 51419-21) as wetmorei. These additional specimens narrow the morphological distinction between frenatus and wetmorei. U. f. wetmorei has a significantly lower ventral count (180) than /. frenatus (189) (t=5.7, 45d.f.), although there is extensive over- lap. The color patterns of /. wetmorei are unlike that of mainland /. frenatus, but are quite similar to those of /. frenatus from He a Vache. Until enough specimens are available to characterize the Barahona population, I prefer conserving the name wetmorei to designate a subspecies of frenatus. 22 BR] \ IORA No. 324 Specimens of frenatus from Diquini show some characteristics that might be attributed to introgression hv oxyrhynchus. BMNH 1913.7.17.29 has a ventral count of l%;'USNM'l 17282 has the anterior loreal scale; MCZ 8683, USNM 117282. and 117346 differ from typical f. frenatus in coloration, being gray with a broad stripe oi gray-green down the back. Each oi these speci- mens can be detiniteK identified as an aberrant /. fremitus, but it seems strange that all the frenatus from Diquini are aberrant. Recent collections from the Diquini area include no frenatus, but typical frenatus and oxyrhynchus are found together in the Jacmel and Miragoane areas, and in Port-au-Prince, adjacent to Diquini. With DO evidence of interbreeding. Without a larger sample, fur- ther speculation i^n the significance o\ the Diquini population seems pointless URcm \< i k ox1! Kin m in s Dumeril and Bibron Uromacei oxyrhynchtu Dumeril and Bibron, 1854. Erp6tologie Generate, 7: ""2 (type loc. "Senegal"). I\an^i\ Hispaniola: coastal area north of the Cul-de-Sac Plain, southwest peninsula ol Haiti west to Miragoane. lie de la Tortue. and Isla Saona. It is sympatric with catesbyi throughout its range. and with frenatus at the base of the southwest peninsula. V. oxyrhynchus is not known to occur with f. wetmorei, though Mertens I 1939: 7l>) reports seeing an oxyrhynchus and a frenatus in the same bush at "Camberlen" (near San Cristobal. D.R.); he did not catch the frenatus, however, so that il is not possible to be sure that the snake was not a gi"a\ oxyrhynchus. One specimen of oxyrhynchus (USNM 75927), collected by J.S.C. Boswell in 192S. is reporte.l as taken from Anse a Calets. Cionave. I \- ceptional localities o\ other specimens in Boswell's 11>2S collection have been attributed to labeling errors (Schwartz, 1965). Collec- tions from lie de la Gonave in the last ten years contain many catesbyi and dor sous, but no oxyrhynchus. Diagnosis. Snout 3.0 (range 2.5-3.7) times eye diameter; scale rows at midbody 19; ventrals 200 (range: 189-212); subcaudals 199 (range: 183-211); anterior loreal present in 25 of the 129 specimens examined: 6 AMMI. 8 BMNH. 2 FMNH. 97 MCZ, 15 USNM. 1 YPM. Color palter)}. Green morph: dark green above, light yellow- green below, the colors separated by a white lateral line on the 1st and 2nd scale rows from the snout to two hcadlengths beyond the vent, ventral edge of the scales in the 1st row black; head 1969 EVOLUTION OF UROMACER 23 green above, cream below, a light line above the eye, dorsal edge of upper labials black (after MCZ 62701). There are variations in color intensity. The lateral line may be edged by black, yellow, both, or neither. In a few specimens, as much as the anterior half of the lateral line is indistinct or lacking. Gray-green morph: like the green morph, but with the ventral surface finely mottled gray to gray-brown from the neck to the tip of the tail (after MCZ 62703). This morph has been found in Haiti and in the eastern Dominican Republic. Variations are as in the green morph. Gray morph: finely mottled gray above, more coarsely mottled gray below, 2nd, 3rd, and 4th scale rows lighter gray, dorsal edge of scales in the 5th row black from midbody to vent; head as in the green morph (after MCZ 62702). Four specimens of this morph have been taken from scattered localities in Haiti. Uromacer dorsalis Dunn Uromacer dorsalis Dunn. 1920, Proc. New England Zool. Club, 7: 43 (type loc. Gonave Island). Range. lie de la Gonave. It is sympatric with catesbyi. Diagnosis. Snout 2.9 (range: 2.5-3.7) times eye diameter; scale rows at midbody 17 (19 in MCZ 93146); ventrals 200 (range: 192-210); subcaudals 210 (range: 197-228); anterior loreal present (absent in MCZ 93139). 84 specimens were ex- amined: 7 ASFS, 65 MCZ, 12 YPM. Color pattern. The species is monomorphic: finely mottled gray above and below, the dorsal region between the 5th scale rows darker; head green above, light green below (after MCZ 61043). In very light specimens the dorsal stripe is indicated only by black spots on the dorsal edge of scales in the 4th row. Nomenclature. U. dorsalis is more similar to oxyrhynchus than to frenatus in general body form, snout length, ventral count, and the presence of the anterior loreal scale, but more similar to frenatus in midbody scale count, subcaudal count, and color pat- tern. Since dorsalis is unique in its diet and morphologically inter- mediate between two distinct mainland species, I retain it as a distinct species. Although I admit a close relationship between dorsalis and oxyrhynchus, the evidence is not compelling enough to warrant calling dorsalis a race of oxyrhynchus. BREVIORA Museum of Connparsitive Zoology Cambridge, Mass. 15 September, 1969 Number 325 THE GENUS PHENACOSAURUS (SAURIA: IGUANIDAE) James D. Lazell, Jr.1 The members of this order are the animals ihe Lord of the Hebrews did not create to grace his Eden; they are not among the products of the six days' labor. These are the sports, the offthrows, of the universe. . . . These are the weird children of the lust of the spheres. Charles Finney Abstract: A complete revision of the Andean genus Phenacosaurus, us- ing all available museum specimens, is presented. The genus is separated from Anolis on the bases of pelvic osteology and digital pad structure. Of four previously described species, two are retained: heterodermus (including "richteri" and "paramoensis") and nicefori. A new species, P. orcesi, is described from Ecuador. Among the strange and varied productions of the high Andes is a small assemblage of grotesque, big-headed, short-legged, pre- hensile-tailed lizards: the genus Phenacosaurus. Since Dunn's (1944) work on this genus in Colombia, it has not been re-exam- ined as a whole. Through the efforts of E. E. Williams, Museum of Comparative Zoology (MCZ), it has now been possible to amass at one time and place the majority of specimens of all the species of this genus and thus attempt a complete review. Without the cooperation of the curators of many institutions, of course, this would not have been possible. I express here my gratitude to the curators of the following institutions : Instituto de La Salle, Bogota (ILS), British Museum (Natural History) (BMNH), Museum of Vertebrate Zoology (MVZ), University of Michigan Museum of Zoology (UMMZ), Stanford University Museum of Natural History (SU), Field Museum of Natural History (FMNH), Phila- delphia Academy of Natural Sciences (PANS), Fundacion La Salle 1 Massachusetts Audubon Society, Lincoln, Mass., 01773 2 breviora No. 325 de Ciencias Naturales, Caracas (FCN), United States National Museum (USNM), American Museum of Natural History (AMNH). Vienna Museum (WEEN), Munich Museum (MUN), Senckenberg Museum (SMF). Berlin Museum (ZMB), Museum National d'Histoire Naturelle de Paris (MHNP). and Universidad National de Colombia (UNC). Professor Gustavo Orces-V, Esquela Politecnica Nacional, Quito. Ecuador, lias very kindly donated the first phenacosaur recorded from Ecuador, which I take great pleasure in designating as the type of a new species named in his honor (see below). Dr. Fred Medem, Cartagena. Colombia, has made available an excellent collection of Colombian phenacosaurs, including the type and paratopes of P. richteri Dunn. Those specimens actually collected by him are fresh alcoholics and combine color reten- tion with very accurate data — including elevations. Dr. George Gorman, Museum of Vertebrate Zoology. Berkeley, has recently collected series in the Bogota area. 1 he Orsono-Mesas ( 1946) have provided an informative treat- ment o{ several aspects of phenacosaur ecology and behavior. I lor photographs of living phenacosaurs taken around Bogota have been provided by Fred Medem and Jose Muho/, Hans Zenke, and Victor I lutchison. In addition to the problems posed by the species-group level taxonomy of Phenacosaurus, it seems necessary to re-examine the status and validity of the genus. Anolis heterodermus Dumeril (1851) was included by Boulenger (1885) in his concept of Xiphocercus Fitzinger (1843). Xiphocercus1' was then sep- arate,! from Anolis (and ChamaeleoUs) on the character com- bination of tricuspid lateral teeth plus a prehensile tail, and from "Norops" on the character of a raised terminal phalanx. Barbour (1920) separated heterodermus from the West Indian form {"Xi- phocercus" [~- Anolis] valencienni) on the basis of body squa- mation. dorsonuchal crest, elevated rim of head plates (= cir- cumoccipital swelling or casque of Dunn. 1944), digital dilations, and tail structure. "Xiphocercus" heterodermus thus became the type of the new genus Phenacosaurus Barbour ( 1920). However, Williams ( 1962) has shown "Xiphocercus" to be a synonym of Anolis; the validity of Phenacosaurus may be questioned also. Cer- tainly neither body squamation nor dorsonuchal crest is useful in separating Phenacosaurus from Anolis. Caudal vertebrae that lack autotomy septa are found in such distantly related Anolis as dar- lingtoni and latijrons; though no Anolis has yet been demonstrated 1969 PHENACOSAURUS 3 definitively to use its tail in a prehensile manner, it seems reason- able to infer from structure that some, like darlingtoni, do. The "circumoccipital swellings," or "casque," referred to is indeed a striking feature of adult male heterodermus. Actually posterolateral^ directed flanges of the parietals, this "casque" is but weakly developed in juveniles and females of heterodermus, or in most specimens of P. nicefori, regardless of sex, and is re- duced in the new form from Ecuador. In addition, some species of Anolis (e.g. richardi of the Lesser Antilles) develop similar parietal flanges; there is, admittedly, still a definite degree of differ- ence between the condition in some Phenacosaurus and that shown even by adult male Anolis richardi, but one would hesitate to ac- cept a genus wholly based on such a degree of difference. The digital dilations of Phenacosaurus are distinctive. The toes are comparatively shorter than the toes of Anolis of similar size; the digital pad is expanded not only beneath the second and third phalanges of the fourth toe, but beneath the first phalanx as well; proximal continuation of the expanded digital pad is the rule with all the toes, and produces a rather gecko-like appearance of the foot. Although this condition is not matched by any species of Anolis, the variation in structure and dilation of the digital pad in Anolis is sufficiently great (see Williams, 1963) so that any degree of difference shown by the phenacosaur condition is surely of di- minished importance. After examination of skeletal material of more than 70 species of Anolis in the MCZ, I have discovered that not only is the struc- ture of the ilium remarkably constant in that genus, but that it is strikingly different from the condition seen in all forms of Phena- cosaurus. I therefore propose a redefinition of the genus Phena- cosaurus and provide the following definition of Anolis for com- parative purposes: Anolis is a genus of iguanid lizards without femoral pores; sub- digital lamellae bear cuticular hairs; pre- and post-zygopopheses of trunk vertebrae not expanded to form great lateral flanges; angular absent; lateral teeth cusped; ilial shaft subcylindrical, dorsal border forming an obtuse angle with the acute or nodular, elongate, anterior iliac process (see Figure 6). Phenacosaurus Barbour Anolis (part), Dumeril, 1851, Cat. Meih. Rept., p. 59. Xiphocercus (part), Boulenger, 1885, Cat. Lizards Brit. Mus. II, p. 10. Phenacosaurus Barbour, 1920, Proc. New England Zool. Club 7, p. 62. Type species. Anolis heterodermus Dumeril, 1851. 4 breviora No. 325 Definition. [guanid lizards without femoral pores; subdigital lamellae bear cuticular hairs; pre- and post-zygopopheses of trunk vertebrae not expanded to form great lateral flanges; angular ab- sent; lateral teeth cusped; ilia! shaft compressed and bladelike, in- clining smoothly into a blunt, subrectangular anterior iliae process ( see I ig. 6). The condition of the ilium can be readily checked without un- due damage to the specimen by merely lifting the skin above one hind limb and separating the muscle attachments from the bone. This has been done for all oi /'. nicefori and the new species, as well as many P. heterodermus. The only dry skeletons of Phena- cosourus examined are Il.S 55d and MCZ 17111. both lietero- dermus. Polychrus have femoral pores present in males and lack an an- terior iliac process. Anisolepis and Aptycholaemus (nearly or quite synonymous, tnle Etheridge, pers. eonun.) have an ilium basically similar to that of Phertacosaurus, but possess a "fairly large'* angular, fide Ether- idge (in Hit.). Anolis and Phertacosaurus lack an angular bone. 1 he remaining anoline genera isensn Etheridge. 1959) have an ilium like that oi Anolis. In addition, Tropidodactylus is unique in lacking cuticular hairs on the lamellae; Chamaeleolis has sim- ple, peglike lateral teeth and a small angular; ChomaeUnoropS has trunk vertebrae bi/arrely modified by expansion and amalgama- tion of the pre- and postzygopophases to form great lateral llanges. The usefulness of separating Phenacosourus from Anolis may be questioned, and Dr. R. A. Etheridge (in lilt.) has questioned the separation in principle. This, certainly, is the place to make a case for the genus, especially as I regard it as valid. Etheridge notes that phenaeosaurs are an early offshoot of "alpha Anolis stuck in South America that have developed some peculiar be- havioral specializations and assorted anatomical specializations." In that, we concur; what alarms Etheridge. however, is that "Phen- acosourus heterodermus (for example) is much more closely re- lated to Anolis jacare than Anolis jacare is to Anolis nebulosus (for example).'* That is also true, but it in no way invalidates the genus Phenacosourus. The difficulty, and the principle at stake, is the old bugbear of relationship. It is perfectly true that birds are. in a very real sense, more closely related to crocodilians than crocodilians are to lizards, and far more closely than crocodilians are to therapsids (for example). Yet. birds are not included within the Reptilia; they belong to a distinct class of their own. Simpson 1969 PHENACOSAURUS 5 (1961, pp. 129-132 and 140-145) has given detailed con- sideration to this problem. Its solution lies in defining taxa in higher categories in a way that best expresses their evolutionary position and adaptive similarities. In the present case, Anolis jacare and A. nebulosus are members of a presumably monophy- letic group that have so much in common, from an adaptive and evolutionary point of view, that they are best regarded as con- generic despite obvious differences. Also, a plethora of other Anolis species spans the gap between them, and thus helps to unite them. The species of Phenacosaurus, on the other hand, are sharply dis- tinct, in important adaptations, from any Anolis; the gap is not spanned by living intermediates. Of course, intermediates must once have lived, and this leads us to recognize a compelling truth: Any taxon at the level of a higher category (genus or above) must — either now or in the past — contain some member more closely related to members of some other taxon than that member may be to some members of its own taxon. I draw a line by defining the genus Anolis: the species of Phenacosaurus are outside that line. It is only my opinion that Phenacosaurus — an apparently nat- ural group of species — qualifies for the rank of genus; no defi- nition of the category genus exists to tell me whether Phenacosaurus is one or not. I rest my case, therefore, on the evolutionary and adaptive significance of the characters that separate the groups: the structure of the ilium and the digital dilations. These seem to me to be quite as important as the distinctions, noted above, which separate the other anoline genera. Phenacosaurus heterodermus (Dumeril) Anolis heterodermus Dumeril, 1851, Cat. Meth. Rept., p. 59. Syntypes: MHNP 1664, A1664, B1664, 6798, A6798. Xiphocercus heterodermus, — Boulenger, 1885, Cat. Lizards Brit. Mus. II, p. 10. Phenacosaurus heterodermus, — Barbour, 1920, Proc. New England Zool. Club 7, 61-63. Phenacosaurus richteri Dunn, 1944, Caldasia 3, p. 60. Type: MCZ 69120. Phenacosaurus paramoensis Hellmich, 1949, Dtsch. Aquar.-u. Terrar.-Ztschr. 2, p. 91. Type: MUN 118/37. Diagnosis. A Phenacosaurus with large dorsal scales: 8-19 dor- sals contained in the standard distance1 (not counting interstitial i Defined as the distance from the center of the eye to the tip of the snout. 6 rrf.yiora No. 325 granules); head plates largo: interparietal sometimes as long as dis- tance aeross head between orbits, but often shorter; subdigital lamel- lae under second and third phalanges oi fourth toe 18-24. Lectotype. MHNP 1664, an adult male with 14 dorsals in the standard distance and 19 subdigital lamellae. Type locality. "Nouvelle Grenade" (= Colombia), here re- stricted to the vicinity of Bogota. Colombia. Discussion. The extreme variability of this species has led to taxonomic difficulties and confusions, and to the naming, there- fore, of two additional species richteri Dunn ( 1944) and paramoensu Hellmich (1949). Dunn, in his 1944 review, used the following characters to separate "richteri" and heterodernuis: Dorsal crest squamation. Dunn separated "richteri" (and iiicc- fori) from heterodermus on the basis o\' single, rather than double, dorsal crest scale row. In attempting to make sense out of the remarkable variation I found in this character. I assigned numbers, from one to ten. to the various sorts of crest scale arrangements noted. In Table 1 each specimen is graded according to the sort of crest scale variant it possesses. Where specimens showed more than one sort of arrangement, the two or more tirades arc indi- cated. The following ten of variation were noted: 1. Undifferentiated middorsal scales; i.e., no dorsal crest at all, except sometimes a median scries of single, enlarged scales on the nape or tail. 2. Some enlarged scales along the middorsum, but not arranged in a particular pattern. 3. Enlarged, tectiform to carinate scales along the middorsum, but separated from each other by undifferentiated smaller scales. 4. Single row of enlarged, tectiform to carinate scales in contact with each other along the middorsum. 5. Single row of spike-like scales in contact with each other along the middorsum. 6. Staggered series of tectiform to carinate scales, every other one separated by undifferentiated smaller dorsals, but each usually contacting the scale immediately adjacent at the dorsal midline. 7. Alternating series of tectiform to carinate scales, continu- ously in contact with each other. 8. Double row of spike-like scales on anterior body combined with alternating tectiform to carinate scales posteriorly. 9. Alternating series of spike-like scales. 10. Double row of spike-like scales. PHENACOSAURUS L» CT> o .9 — ■ 03 c 03 "a, x X 00 CO oo s o «J a c -s; a, c o a3 CSJ 03 — O •a •a 1 a o .2 03 > 3 oo 8 breviora No. 325 In Figure 1 each of these ten variants is pictured. I do not find the condition of the dorsal crest squamation diagnostic for any of the named forms. Cranial ridges. Dunn (1944) differentiated both of his new species from heterodermus on the basis of the structure of the circumoccipital and postocular ridges. P. heterodermus was sup- posed to have swollen scales making up these ridges, whereas "riehteri" and nicefori were said to have swollen bone structure underlying the scales: the scales themselves were not supposed to be swollen in the latter two forms. Of "riehteri" he states: "nor are they [these scales] enlarged" except slightly in old males (p. 60). In all the specimens I have seen, including the entire tvpe series of "riehteri" and the svntvpes of heterodermus, the scales of the circumoccipital and postocular ridges are greatly enlarged, tecti- form to carinate, and. in addition, lie on bony ridges of the skull. This character, therefore, fails to separate taxa. Chin shields. Dunn (1944) separated "riehteri" from hetero- dermus on the basis of the extent of contact between the enlarged chin shields and the infralabials. Anteriorly, the chin shields and infralabials are always in contact, posteriorly they arc separated by small sublabials. The point at which separation begins is very variable, and the condition supposedl) representative of hetero- dermus is to be found in the type series of "riehteri." I could not correlate variation in this character with any of the others men- tioned by Dunn. Contact of dorsal scales. As with the preceding characters, Dunn recognized two conditions: enlarged dorsal scales either contacting each other, or separated. Both conditions do, ad- mittedly, occur, but very often on the same li/ard. I could not correlate this condition, even when clear-cut. with any of the other characters. Leg length. As noted by Aleman ( 1953). leg length is variable and will not separate taxa unequivocally. In general, however, nicefori do tend to have much shorter legs in the southern part of their range (e.g., Pamplona); heterodermus appear to have longer legs; in the northern part of their range (e.g., Saota, Paz del Rio). Therefore, though leg length cannot be used to diagnose the forms, it may indicate character divergence between heterodermus and nicefori, and is discussed further under the latter species. Leg length cannot be used to divide heterodermus as here defined into taxa, and "riehteri" is. therefore, not separable from heterodermus on any character or combination of characters discernible by me. 1969 PHENACOSAURUS 9 Scale size. As may be noted from the above diagnosis of hetero- dermus, I do regard scale size as useful in separating forms. I have used two indices to scale size: the size of the head plates is here indicated by the length of the interparietal plate compared to the distance between the orbits across the top of the head. As noted, heterodermus has rather large head plates; the interparietal plate may be as long as the distance between the orbits, though it is usually somewhat shorter. The size of the dorsal scales, indi- cated by the number contained in the standard distance just lateral to the midline at midbody, separates nicefori and heterodermus. The new form from Ecuador combines scale size measured by both indices in a way that is radically different from either of the previously named forms. Both nicefori and the new species are discussed fully below. Dorsals in the standard distance just lateral to the midline at midbody are tabulated for heterodermus and nicefori in Table 1. Coloration and pattern. Although not utilized taxonomically by Dunn (1944), coloration and pattern are obviously subject to some striking variation, at least in heterodermus as here defined, and must be considered more fully. Hellmich (1949) described a single specimen from Paramo de Sumapaz, 3750 meters, as a new species, P. paramoensis. The coloration and pattern of this individual are striking (see Figs. 2 and 3). Two other specimens I have seen match this condition: the smaller of ZMB 4256, a female (as is MUN 118/37, the type of "paramoensis"), and AMNH 91752, a male. Numerous other specimens approach this condition. Two specimens, MUN 9/49 and MVZ 68686, from Tambo, Cauca, and San Antonio, Valle, respectively (the most south- western localities for P. heterodermus), are pale ash-grey with very broad, brown bands across the dorsum. Excellent color transparencies taken at Bogota by Hutchison show that the colors noted occur in living individuals at the type locality. Hutchison further notes (pers. comm.) that individuals are capable of chang- ing their patterns as well as their colors. Range. Phenacosaurus heterodermus, as may be seen in Fig- ure 4, occurs in both the Cordillera Oriental and the Cordillera Central. The species occurs at least in the extreme southern Cordillera Occidental near the head of the Cauca River, at San Antonio, Valle. Because the Cauca River, which separates the Cordillera Occidental from the Cordillera Central, is nowhere as great a barrier as is the Magdelena, which separates the latter 10 BRLVIORA No. 325 X =: O =: 5 u Z s c o to ! - I - -=: a. I 1969 PHENACOSAURUS 11 N.STRZK . Figure 3. Phenacosaurus heterodermus, top of head. Specimen MUN 1 18/37, the top of " paramoensis" Hellmich. 12 breviora No. 325 from the Cordillera Oriental, and because all three Cordilleras merge in southern Colombia (e.g., vicinity of Tambo, Cauca), I expect the total distribution of the species, at least in the western range, to be greater than is presently known. Dunn (1944) gives the altitudinai range as from 1800 m to 3500 m. Hellmich (1949) cites the elevation for "paramoensis" (= heterodermus) as 3750 m. This is the highest recorded elevation for the species. Size. The largest specimen measured is MCZ 78531, from Paz del Rio, Boyaca. It is 86 mm snout to vent. Remarkably, it is a female. In the more southern populations of heterodermus males seem to average larger than females, and reach 83 mm snout to vent (male of ZMB 521 1 ) in the Bogota area; females in the Bogota area reach 80 mm (MHNP 1923.55). I therefore expect some males from the northern part of the range to be larger than any specimen here recorded. Si/e is discussed comparatively under the following species: Phlnacosaurus nicefori Dunn Phenacosauriu nicefori Dunn, 1944. Caldaaia 3, p. -s^. Type: ILS 64. Diagnosis. A Phenacosaurus with small dorsal scales: 21-34 contained in the standard distance (not counting interstitial gran- ules): head plates small, interparietal much shorter than the dis- tance across head between orbits; subdigital lamellae 16-20. Type. ILS 64, an adult female with 24 dorsals in the standard distance and 17 subdigital lamellae. Type locality. Pamplona, Norte de Santander, Colombia. Discussion. On the basis of the eight specimens I have seen, this species is quite closely allied to heterodermus. The only defin- itive difference I have been able to discern is dorsal scale size. With respect to this character, it must be noted that for both nice- fori and heterodermus the range of variation within the species, respectively, is greater by far than the gap which separates them. I have indicated in my discussion of heterodermus that I believe there is some evidence for character divergence between these two forms as they approach the area of potential sympatry. Though heterodermus and nicefori have not yet been demonstrated to be sym- or parapatric at any point (see Fig. 4), what evidence for character divergence we do have should be considered in full. Specimens of both forms have been arranged in Table 1 in a north- south sequence from Cerro Tetari (northernmost locality for nice- fori, and the genus) to Tambo (the southernmost locality for 1969 PHENACOSAURUS 13 heterodermus) . Thereafter, in Table 1, the sequence is northward through the more western heterodermus localities in the Cordil- leras Occidental and Central. Characteristics of dorsal scale size, dorsal crest type, and subdigital lamellae are tabulated; these, with the exception of dorsal crest type (which I do not regard as indicative of either taxon or character displacement), are included in the discussion below: Dorsal scale size. Although the numbers of specimens in the samples do not permit any meaningful statistical analysis, nicefori and heterodermus do seem to become more different as they ap- proach one another. The smallest dorsals in heterodermus ( 1 9 in the standard distance) are in FMNH 69673, from San Antonio, Huila, one of the southernmost examples. Similarly, the largest dorsals in nicefori (only 21 in the standard distance) are in FCN 664, from Cerro Tetari, Venezuela — the northernmost repre- sentative of that form. The differences between the more nearly adjacent populations — Pamplona (with one specimen of nicefori with 22) and Saota (heterodermus with 14) — are not spectacu- lar, but with more evidence might prove meaningful. Subdigital lamellae. In this case the situation is not indicative; assuredly, the Cerro Tetari nicefori has the highest number of lamellae found in that species, but the highest counts in hetero- dermus come from the southern part of that range. The more nearly adjacent populations hardly differ with respect to this char- acter. Leg length. As mentioned previously, leg length does seem to provide an example of character divergence. However, females of both forms seem to be shorter-legged than males, and I can find little difference between them. Fortunately, the three southern- most heterodermus localities (San Antonio, Huila; Tambo; and San Antonio, Valle) are represented by males. In all of these the appressed hind limb barely reaches the axilla. In most males from around Bogota, the one male from Paz del Rio, and in the Saota specimen, the appressed hind limb extends beyond the axilla. In Pamplona males of nicefori (2) the hind limb is, as in females, shorter than the distance to the axilla, whereas in the Cerro Tetari male, as noted by Aleman (1953), the hind limb is quite as long as this distance. Even in a case like this, however, where the evidence seems clear cut on the face of it, the number of individuals involved is much too small for any sort of surety. Size. Here again there is but little evidence. The largest hetero- dermus examined, as noted, was from a northern locality, one of 14 breviora No. 325 the two closest to a nicefori population. Similarly, the largest nicefori, FCN 664, is 86 mm snout to vent, and from the northern- most locality, farthest removed from the known range of hetero- dermus. Because all phenacosaurs begin life small, and because I have such a paucity of specimens and information about living animals, I cannot claim that this is necessarily meaningful. Coloration and pattern. Most of the specimens of nicefori are formalin specimens, and show nothing in the way of coloration or pattern. FCN 664. however, is well preserved and appears to have been rather uniform green with a white stripe from the supralabials to the shoulder, as in many heterodermus. There is nothing about any of the other specimens of nicefori that could contradict the assumption that this is how they all look. In the foregoing discussion I have tacitly assumed that FCN 664 actually represents a nicefori. If it does, it is by far the largest representative seen; those from around Pamplona are all less than 65 mm snout to vent Also, it has the largest dorsals, the highest number of siibdigital lamellae, and the longest legs of any nicefori seen. I have suggested that these differences may be attributable to character divergence in two species. On the basis of present infor- mation I see no alternative; if FCN 664 is not a nicefori, then a great deal of additional material will be required to demonstrate its true relationships. Certainly heterodermus and nicefori. whether they are distinct species or geographic representatives of the same species, are closely related. Extreme with respect to major characters, the relationships of the new form from Ecuador are in no way so clear. I describe it as: Phenacosaurus orcesi sp. nov. Type. MCZ 38937, collected iv. 1957 by Jorge Olalla. Type locality. Mt. Sumaco. Napo Pastaza Province, Ecuador. Paratype. USNM 16533, collected by James Peters between L'Alegria and La Bonita, Ecuador. Diagnosis. A Phenacosaurus with very small dorsals, 38-48 contained in the standard distance (no interstitial granules pres- ent); head plates very large, interparietal much longer than the distance across the head between the orbits; subdigital lamellae under second and third phalanges of fourth toe 16-18. Description of the type. MCZ 38937 is a female measuring 58 mm snout to vent, with a 61 mm tail. Both the foot structure and the ilium (dissected on left side) are typically phenacosauran. 1969 PHENACOSAURUS 15 cerJo \ VENEZUELA 'A Tetari -f{ i/\ Pamplona ^ San Pedro ^ ±*T* -~ f\\ . -+- Paz di -> ' -r- Sonson ' del Rio }~j$- Bogota 4- Paramo de Sumapaz San Antonio, Valle-f-/,' "" -|- San Antonio, Huila -j- Tambo COLOMBIA + heterodermus A nicefori (g> orcesi Figure 4. Localities for three species of Phenacosaurus. Names are not cited here for localities in the immediate vicinity of Bogota but are tabulated in north-south order in Table 1. 16 breviora No. 325 There are three large plates across the snout at the level of the second canthals. Two scales, the right one of which appears to be two smaller scales anastomosed, border the rostral. The nasal is entire, in contact with the first supralabial. and separated from the rostral by a single, subrectangular prenasal. The supraorbital semicircles are composed of large plates, three of which are broadly in contact at the midline. The supraocular disks are composed of several large plates, two or three of which border directly on the supraorbital semicircles, that decrease in size laterally. There are no supraeiliarv scales distinguished from the orbital granules. There are two rows of loreals. The interparietal is very large, ca. 1.3 times as long as the distance between the orbits across the top of the head, and is in contact with the supraorbital semicircles. I he circumoccipita] ridges are reduced to the point of absence. The suboculars are in contact with the supralabials. The mental is sutured, bordered posteriorly by the infralabials, sublabials, and four gulai granules — eight scales in all. A single series of sharply leetiform crest scales, separated by undifferentiated dorsal granules, begins on the nape and continues to the rump; this is a '.V' type crest scale pattern. The dorsal scales are granular and of rather uniform si/c; there are 38 con- tained in the standard distance at midbody. The ventrals are smooth, slightly swollen, and subimbricate medially. There are 18 subdigital lamellae under the second and third phalanges of the fourth toe. The digital dilations are very broad and become only gradually narrower proximally. The tail is very slightly compressed; there is a single row of slightly enlarged, tcctiform scales middoisally beginning some- what posterior to the level of the vent. There are no scale whorls or other indications o( autotomy septa. There is apparently not a functional throat fan in this female. This is a formalin specimen; it has been rendered a dark, disa- greeable, mud-brown all over; no trace of coloration or pattern remains. The type is illustrated in Figure 5. The paratype. The only other specimen of P. orcesi currently available is a male 60 mm, snout to vent, with a 67 mm tail. The digital pads of both fourth toes on the hind feet taper more abruptly proximally than is usual in Phenacosaurus, but are broader under the first phalanx than in Anolis of similar size. The ilium (dis- sected on the left) is typical of the genus: blade-like, with a blunt, obtuse anterior prominence. 1969 PHENACOSAURUS 17 Figure 5. Phenacosaurus orcesi sp. nov., type, MCZ 38937, from Mt. Sumaco, Ecuador. 18 breviora No. 325 The paratype is similar to the type in all details described above except the following: There are four large plates across the head between the second canthals. and three scales border the rostral posteriorly. Two plates of the supraorbital semi-circles are in contact at the midline. The circumoccipital ridges, though low, are quite evident. Because one row of gular granules fails to reach the mental, only seven scales border it posteriorly. There are no enlarged crest scales on nape or back; this is a "1" type crest scale pattern. The dorsals are granular but variable in size; counts of 46 to 48 can be made in the standard distance at midbody. There are 16 or 17 lamellae under the second and third phalanges of the fourth toe. The tail bears a scries of enlarged, tectiform scales, separated by smaller granules, beginning just posterior to the level ol the vent; this is a "3" type crest scale pattern. The throat fan appears about as well developed as voung males of P. nicejori or P. heterodermus: not apparently greatly extensible. Dr. James Peters reports (in lilt.): "The specimen was picked up by me personally. ... It was found on the trail between L'Ale- gria and La Bonita. in Napo Province, Ecuador, high above the river that forms the boundary between Ecuador and Colombia, on the eastern slopes, on June 26, 1962. It was found dead on the trail, in a horse's hoofprint. ... It was rather badly beaten up . . . — in the state you see it now. unfortunately. The elevation at which it was collected is approximately 5800 feet." Discussion. While Phenacosaurus heterodermus and P. nicejori seem quite closely allied forms. P. orcesi appears remote. The dor- sal squamation lacks the large scales interspersed with granules that both other species exhibit. The dorsals are smaller than in nicejori, though the head plates are larger than in heterodermus. The general appearance is very much like some Anolis (e.g., soli- tarius); the small scales augment this effect. The pelvic girdle, however, is distinctly phenacosauran; P. orcesi, in fact, differs from Anolis with respect to the ilium in the same ways that hetero- dermus and nicejori do, but to an even greater extent; the ilial prominence is very obtuse and weakly developed in both speci- mens available. All considered, we know so little about the phenacosaurs in general that any attempt at a reasonable assessment of relation- ships is pointless. Until series of specimens can be collected from the many crucial areas presumably included within the range of the genus, and until comparisons can be made utilizing such 1969 PHENACOSAURUS 19 characteristics as coloration in life and behavior in the field — features demonstrably of great importance in the systematics of anoline lizards — our knowledge of Phenacosaurus will remain reprehensibly elementary. ACKNOWLEDGMENTS Figures 2 and 3 were prepared by N. Strekalovsky. Dr. Richard Etheridge has provided data on anoline osteology. This work was supported by National Science Foundation Grants G- 16066 and GB6944, held by Ernest E. Williams. LITERATURE CITED Aleman, C. 1953. Contribucion al estudio de los reptiles y batracios de la Sierra de Perija. Mem. Soc. Cien. Nat. La Salle (Venezuela) 13: 205-225. Barbour, T. 1920. A note on Xiphocercus. Proc. New England Zool. Club 7: 61-63. BOULENGER, G. A. 1885. Catalogue of the Lizards in the British Museum, II. London, xiii -+- 497 pp. Dumeril, A. A. 1851. Catalogue methodique de la collection des reptiles. Museum National d'Histoire Naturelle, Paris, 224 pp. Dunn, E. R. 1944. The lizard genus Phenacosaurus. Caldasia 3 (11 ) : 57-62. Etheridge, R. 1959. The relationships of the anoles (Reptilia: Sauria: Iguanidae): an interpretation based on skeletal morphology. Univ. Micro- films 60-2529, Ann Arbor, Mich., xiii + 236 pp. Fitzinger, L. J. 1843. Systema Reptilium. Vienna, vi -f 106 pp. Hellmich, W. 1949. Auf der Jagd nach der Paramo-echse. Dtsch. Aquar.-u. Terrar.- Ztschr. 2 (5): 89-91. Orsono-Mesa, H., and E. Orsono-Mesa 1946. Anotaciones sobre lagartos del genero Phenacosaurus. Caldasia 4 (17): 123-130. Simpson, G. G. 1961. Principles of Animal Taxonomy. Columbia University Press, xii + 247 pp. 20 BREVIORA No. 325 Williams, E. E. 1962. Notes on the herpetology of Hispaniola. 7. New material of two poorly known ;mo!es: Anolis monticola Shreve and Anolis christophei Williams. BrevioraNo. 164: 1-11. 1963. Studies on South American ancles. Description of Anolis mirus, new species, from Rio San Juan, Colombia, with comment on digital dilation and dewlap as generic and specific characters in the ancles. Bull. Mus. Comp.^Zool. 129 (9): 463-480. (Received 29 April 1969). Figure 6. Ilia of two anoline lizard genera: A. Anolis (specimen MCZ 61027, A. richardi, Tobago, West Indies); B. Phenacosaurus (speci- men MCZ 17111, P. heterodennus, Guasca, Colombia). 1969 PHENACOSAURUS 21 TABLE 1: SPECIMENS EXAMINED Specimens of the three species of Plienacosauriis and three of their characters: dorsal scales counted in the standard distance, type of dorsal crest, and number of subdigital lamellae. See text and map (Fig. 5). The locality "Bogota" is of highly variable precision. PHENACOSAURUS NICEFORI Lamel- Specimen Sex Locality Dorsals Crest lae FCN 644 $ Cerro Tetari 21 3 20 FMNH 5684 9 Paramo de Tana 28 6 17 FMNH 56443 juv. Pamplona 31 3 18 PANS 25865 $ 44 27 3 17 ILS 64 9 44 24 3 17 ILS 64a 9 44 22 3 16 MCZ 6729 $ 44 34 3,4 17 USNM 72745 9 Tapata 33 3 16 PHENACOSAURUS HETERODERMUS ILS 63 $ Saota 14 9 18 UNC 692 9 Paz del Rio 10 2,9 21 UNC 693 $ 44 44 14 5,7 20 MCZ 78520 S Cogua 11 3,5 20 MCZ 78521 $ (( 12 4 20 ILS 62 S Subachoque 13 8 18 ILS 62a S tc 15 9 19 MCZ 78524 £ Cajica 13 5,4 21 MCZ 78525 8 ^ 10 5,4 21 MCZ 78526 9 t< 9 5 20 MCZ 78527 $ u 11 5,3 20 MCZ 78528 $ (4 12 5 20 MCZ 78529 $ 41 14 5,3 19 MCZ 78534 juv. M 12 4 20 UNC 705 9 (4 10 5,4 21 MCZ 104409 $ Chia 10 4 19 MCZ 110133 4 44 10 7 10 MCZ 110134 9 44 12 4 18 MCZ 110135 juv. 44 17 2 20 MCZ 110136 9 44 8 2 22 MCZ 110137 9 44 14 2 22 MCZ 110138 S 44 11 2 20 MCZ 110139 6 )» 11 7 19 MCZ 78522 9 Usaquen 10 8 19 MCZ 78523 S Bosque Popular 16 6 20 22 BREVIORA No. 325 Specimen Sex Locality Lamel- Dorsals Crest lae 12 4,7 19 10 4 20 9 5.7 19 12 4 21 10 5 23 12 8 19 10 8 20 11 8 18 9 5 19 11 5 20 10 5 19 12 3 20 8 5 20 12 5,6 18 12 5 20 10 5,6 19 10 5 20 13 5,6 20 10 5.6 20 10 10 19 1 1 9.5 20 14 4 19 14 l>. 5 21 13 9.6 20 14 9 19 12 9.5 19 10 6 21 12 5 20 1 1 5,6 20 13 9. 10 20 13 9,6,4 21 12 9 20 15 5 20 14 5 19 12 1 21 1 1 8 21 13 1 19 13 5. 6, 3 20 1 1 8,6 20 12 8 21 14 9,5 20 11 8 21 14 8 20 13 8 20 MCZ 78519 $ Ten a MCZ 69120 $ Tabio MCZ 69121 9 " MCZ 69122 <5 .. MCZ 69123 $ .4 PANS 25074 juv. .. PANS 25192 6 .4 PANS 25193 & .. PANS 25194 6 .» PANS 25195 $ II PANS 25196 9 II PANS 25197