bee Aetams Ar tiieg ba) : BULLETIN OF THE MUSEUM OF COMPARATIVE ZOOLOGY AT HARVARD COLLEGE, IN CAMBRIDGE VOL. 121 CAMBRIDGE, MASS., U.S.A. 1959 Tue Cosmos Press, Inc. CampBripGe, Mass., U.S.A. CONTENTS PAGE No. 1.—-DESCRIPTION OF THE SKULL OF POMATODELPHIS IN- AEQUALIS ALLEN. By Remington Kelloge. (6 plates). May, 1959 . : 5 : ‘ ; : ] No. 2.—LAND AND FRESHWATER MOLLUSKS OF GREAT AND LittLE INaagua, BAHAMA ISLANDS. By William J. Clench. (1 plate). May, 1959 . 27 No. 3.—THE SPIDER GENERA ACHAEARANEA, THERIDION AND SPHYROTINUS FROM Mexico, CENTRAL AMERICA AND THE West INDIES. (ARANEAE, THERIDIIDAE). By Herbert W. Levi. July, 1959 . f ? : paler 15), No. 4.—Turee New GENERA AND ONE NEW SPECIES OF THE FAMILY GONOSTOMATIDAE. By Marion Grey. July, 1959 : : ? : : : : A ellos No. 5.—THE ANOLES OF THE EASTERN CARIBBEAN (SAURIA, I@GuUANIDAE). Parts I-III. By Julan S. Kenney, Victor C. Quesnel, Garth Underwood, Ernest E. Williams. (1 plate). July, 1959 . ASS No. 6—On a TAXONOMIC PUZZLE AND THE CLASSIFICATION OF THE EARTHWORMS. By G. E. Gates. August, 1959 227 No. 7.—BATHONIAN FORAMINIFERA OF ENGLAND. By Richard Cifelli. (7 plates). September, 1959 . 263 No. 8.—S micirIeED MippLE ORDOVICIAN TRILOBITES. (REMO- PLEURIDIDAE, TRINUCLEIDAE, RAPHIOPHORIDAE, EN- DYMIONIIDAE). By H. B. Whittington. (26 plates). November, 1959 , s ‘ : : i a69 ae Mi ‘ab ene ir, |) ay ee ih « a. 7 - oy Bulletin of the Museum of Comparative Zoology AUT rH ASR Wave, DiC Oia; Bh Gan Vor da Now! DESCRIPTION OF THE SKULL OF POMATODELPHIS INAEQUALIS ALLEN By REMINGTON KELLOGG WitnH Six PLAtTEs CAMBRIDGE, MASS., U.S.A. Pee NGsD Se One ist Mees Havent May, 1959 PUBLICATIONS ISSUED BY OR IN CONNECTION WITH THE MUSEUM OF COMPARATIVE ZOOLOGY AT HARVARD COLLEGE BULLETIN (octavo) 1863 — The current volume is Vol. 121. BrREVIORA (octavo) 1952 — No. 109 is current. Memoirs (quarto) 1864-1938 -— Publication was terminated with Vol. 55. JOHNSONIA (quarto) 1941 — A publication of the Department of Mollusks. Vol. 3, no. 38 is current. OcCASIONAL PAPERS OF THE DEPARTMENT OF MOoLuUSKS (octavo) 1945 — Vol. 2, no. 22 is current. PROCEEDINGS OF THE NEW ENGLAND ZOOLOGICAL CLUB (octavo) 1899-1948 — Published in connection with the Museum. Publication terminated with Vol. 24. The continuing publications are issued at irregular intervals in num- bers which may be purchased separately. Prices and lists may be obtained on application to the Director of the Museum of Comparative Zoology, Cambridge 38, Massachusetts. Of the Peters ‘‘Check List of Birds of the World,’’ volumes 1-3 are out of print; volumes 4 and 6 may be obtained from the Harvard Uni- versity Press; volumes 5 and 7 are sold by the Museum, and future volumes will be published under Museum auspices. Bulletin of the Museum of Comparative Zoology AT HARVARD COLLEGE Monn 2iG No: 1 DESCRIPTION OF THE SKULL OF POMATODELPHIS INAHKQUALIS ALLEN By RemInatTon KELLOGG Wirth Six PLATES CAMBRIDGE, MASS., U.S.A. PRE NEED HOR, THK MUsSiEhAul Mm May, 1959 No. 1 — Deseription of the Skull of Pomatodelphis inaequalis Allen By REMINGTON KELLOGG Included in the collections of the Museum of Comparative Zoology, Harvard College, is an essentially complete fossilized skull of a porpoise which Dr. Alfred S. Romer, Director, has kindly placed at my disposal for study and description. This specimen was found in sandy clay in the pit of the Homeland Mine, Homeland, Polk County, Fla., during 1957, by the field party of A. Lewis. Before describing the skull, it seems desirable to review briefly the published occurrences of specimens referred to the genus Pomatodelphis. The portion of the right maxillary discovered in the Helvetian ‘caleaire erossier”’ of France and subsequently described and figured by Cuvier (Recherches sur les Ossemens Fossiles, 1823, ede 2: vol. 5, pt. 1, p. 31%, pl 23, fig. 38) didinot receivesthe veneric and specific name Delphinus stenorhynchus until 1829 (Holl, 1829, p. 70). Although this basal rostral fragment barely exceeded 12 inches in leneth, it was regarded as belonging to a lone-snouted porpoise at least as early as 1834 (Keferstein, 1834, vol. 2, p. 203). In the second edition of his ‘* Recherches, ”’ Cuvier (1823, vol. 5, pt. 1, p. 317) states that the type specimen came from the **département de 1’}Orne.’” but in the 4th edition (Cuvier, 1836, vol. 8, pt. 2, p. 168) the locality is given as “département de Maine-et-Loire.”” M. Renou, who presented this specimen to Cuvier, was professor of natural history at Angers, and this fact affords reasonable grounds for the belief that Cuvier was advised subsequent to the 1823 edition that the original locality designation was incorrect. It should be noted also that Cuvier recorded bones of a seal (1836, vol. 8, pt. 1, p. 454) and a sirenian (1836, vol. 8, pt. 2, p. 63) which had been secured by Professor Renou in the department of Maine-et-Loire. Presumably the name Delphinus renovi was bestowed on this same specimen by Laurillard (1844, vol. 4. p. 634) in acknowledg- ment of the collecting activities of Professor Renou. Dr. G. M. Allen (1921, p. 153) referred the Helvetian (Middle Miocene ) stenorhynchus to the genus Pomatodelphis and concluded that 4 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY the large and somewhat closely crowded proximal alveoli of the French odontocete afforded ample grounds for the specific sep- aration of the newly discovered Florida porpoise. In 1921, G. M. Allen (1921, p. 148) selected as a type of the new species Pomatodelphis inaequalis a fragment of the right maxillary from the mine pit which was being excavated by the Amalgamated Phosphate Co. at Brewster, Polk County, Va. Other cranial fragments from unrecorded localities in Florida, including a portion of the rostrum of one individual as well as the top and the posterior basal region of the cranium of another individual were referred by Allen to this species. Of less certain generic allocation is the incomplete skull from the lower Miocene *‘mollasse burdigalienne’’ quarries of Chama- ret, Department of Drome, France, which Paquier (1894, p. 7) described as Schizodelphis depereti. Allen (1921, p. 153) was almost certain that this Burdigalian specimen was referable to P. stenorhynchus. In some of the pits the bedrock (Cooke, 1945) that unecon- formably underlies the phosphate deposits consists of either vellow sandy limestone (Bartow, Fla.) or heht yellow marl and limestone (Mulberry, Fla.). At Pierce, Fla., laminated blue clays underlie the pebble phosphate. These underlying beds have been referred to the middle Miocene Hawthorn formation. The relatively shallow sediments containing phosphates overlying the above strata vary in thickness and in composition. In view of the occurrence in these overlying layers of long-snouted porpoises found elsewhere only in formations of Miocene age, those advo- cating a Pliocene age for all of these stratified sediments should support such a conclusion by incontestible proof that at least in some localities one or more of these layers are not referable to an older geologic age. It is hardly conceivable that either a complete isolated long-snouted skull or one accompanied by the mandibles could represent specimens derived from a reworking of older Miocene deposits. Two of the genera of long-snouted porpoises, Pomatodelphis and Schizodelphis, represented by specimens obtaimed in the course of commercial mining of the pebble phosphate deposits of Florida, are present in the Helvetian deposits of France, and Schizodelphis also occurs in the Helvetian of Switzerland and KELLOGG: SKULL OF POMATODELPHIS INAEQUALIS ) Germany. In geological time, Schizodelphis makes its first appearance in the Langhian (lower Miocene) deposits of Austria, Switzerland, France, Italy and Egypt. The genus Schizodelphis also occurs in the Calvert formation of Maryland, the Kirkwood formation of New Jersey, and the Bolderian of Belgium, all of which are regarded as early upper Miocene. At the present time, Pomatodelphis is limited, by known occurrences in Europe, to the Helvetian stage. No skeletal remains attributable to Schizo- delphis have been recorded from Pliocene deposits in Europe or elsewhere. From the laminated blue clays underlying the pebble phos- phate at Pierce, Polk County, Fla., a larger long-snouted por- poise (Megalodelphis magnidens Kellogg, 1944) and a second specimen of a smaller porpoise (Goniodelphis hudsoni Kellogg, 1944) have been described. The type specimen of Goniodelphis hudsoni (Allen, 1941) was, however, recorded as having been found in the pebble phosphate at Pierce, Polk County, Fla. The stratigraphic position of the oecurrence of Schizodelphis bobengr (Case, 19384) in the mine at Mulberry, Polk County, Fla., un- fortunately was not recorded. The type (No. 828, Florida Geo- logical Survey) of Schizodelphis depressus (Allen, 1921) came from the pit of the Dominion Phosphate Company five miles south of Bartow (Sellards, 1915). Porpoises with skulls characterized by long, narrow rostra are found in most marine Miocene assemblages. The known geologi- cal record reveals that the rostra of the porpoises in this line of development became progressively elongated during the Miocene period. To illustrate this observation, reference is made to the lower Miocene Patagonian Argyrocetus, whose rostrum was twice as long as the cranium, and to the upper Miocene Calvert Zar- hachis, whose rostrum was more than five-sixths of the total leneth of the skull. The rostrum of both Pomatodelphis and Schizodelphis was equal to or exceeded four-fifths of the total leneth of the skull. The pronounced elongation of the rostrum and other differential structural details, including the reduplica- tion of the teeth in the upper and lower jaws, may have cul- minated in an overspecialization which led ultimately to the extinction of this line of odontocete development prior to the Pliocene period. 6 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY The precise stratigraphic position of the Tertiary cetaceans recovered during pit-mining operations for phosphates in Florida in a number of instances has not been carefully recorded. Conse- quently, some difference of opinion has arisen not only regarding the occurrence but also the geologic age of some of the deseribed marine mammals. Our present knowledge of the geologic oceur- rence of the long-snouted porpoises suggests that a re-evaluation is needed of the criteria on which age determinations of the stratified layers overlying the Hawthorn formation have been based. The field party recorded the section at the site (Florida Geo- logical Survey Station 57-1-28 B) where the skull (MCZ 4433) was uncovered in the pit of the Homeland Mine as follows: Top of spoil bank 0-17 ft. 3% ft. thick Bed 1 316 to 4% ft. thick sed 2 2% ft. thick Bed 3 2 to 2% ft. thick Bed 4 1 to 1% ft. thick Bed 5 3 ft. thick sed 6 Covered to water level. Top of section Spoil Rubble in yellow-gray, sandy clay. Rubble consists of vesicular, sandy clay and dark brown ironstone. Unconformity Phosphorite, white pebbles in sandy clay. Unconformity Thin layers of sandy clay and phosphate. (MCZ 4483; Pomatodelphis inaequalis ). As above but not as stratified and with some H-green clay. Unconformity Stratified rubble, layers and masses of vellow-gray to dark yellow-orange phos- phoritic, sandy clay. Upper surface a pave- ment of dark brown phosphorite. Unconformity Stratified layers of yellow-gray to dark yellow-orange, phosphoritic (fine to pebble, light to dark-brown colored) sandy clay or b clayey sand. Numerous ‘‘manatee’’ bones (MCZ 4481-4432). Z. 8. Altschuler of the U.S. Geological Survey suggests that this field party possibly mistook pronounced hthologic or weather- ing changes for unconformities. KELLOGG: SKULL OF POMATODELPHIS INAEQUALIS ( The rostrum (U.S.N.M. 20738) of the second specimen herein- after deseribed was found in hight tan, medium to coarse, slightly clayey, quartz, phosphate sand approximately 8 feet above the Hawthorn formation in the Noralyn Mine, 3 miles south of 3artow, Polk County, Florida. The generic diagnosis published by Allen (1921, p. 148) is revised as follows: Genus POMATODELPHIS G. M. Allen Pomatodelphis G. M. Allen, Journ. Mammalogy, vol. 2, no. 35, p. 148. August 19, 1921. Genotype. Pomatodelphis imaequalis G. M. Allen. Diagnosis. Rostrum elongated, attenuated, dorso-ventrally compressed, and equivalent to more than four-fifths (84%) of the leneth of the entire skull; mesorostral gutter roofed over for most of its length by premaxillaries; width of premaxillary on dorsal surface of rostrum exceeds that of corresponding maxil- lary; ventral or palatal surface of rostrum formed entirely by maxillaries; premaxillaries seemingly not visible ventrally at or near extremity of restrum; palate flat, except for longitudinal, median, V-shaped groove formed by bevelled inner edges of opposite maxillaries; alveoli closely approximated, shallow, some- what slit-like and more or less contracted at one or both ends, except the anteriormost 13 alveoli which progressively increase in size, both transversely and antero-posteriorly as well as in depth toward extremity of rostrum; a row of shallow depres- sions internal and parallel to alveoli for right tooth row on proximal portion of palate presumably lodged tips of teeth in corresponding mandibular tooth row; vertex of skull not con- spicuously elevated; nasal bones sub-rectangular, wider trans- versely than antero-posteriorly, and smaller than exposed pos- tero-internal portions of frontals on vertex ; supraorbital process of frontal noticeably thickened dorso-ventrally, the antero-exter- nal portion of each process being conspicuously enlarged and projected upward above level of premaxillaries; supraoccipital wider than high, concave from side to side, and bounded laterally by flaring lambdoid crest; zygomatic process robust, and dorso- ventrally expanded ; occipital condyles borne on relatively short necks. 5 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY teferred Specimen | POMATODELPHIS INAEQUALIS G. M. Allen Referred specimen. No. 4433, Museum of Comparative Zool- ogy, Harvard College. Essentially complete skull, but laekine most of right zygomatic process and all the teeth. Collector, A. Lewis, 1957. Horizon and locality. Bone Valley gravels, Bed 3 (2.5 feet of thin sandy clay and phosphate). Southeast corner of Home- land Mine Pit, Homeland, Polk County, Florida. NE 14, NW 14 Sec. 4, T 31 8, R 25 E. Middle Miocene. SKULL Dorsal view. This skull (Pl. 1) differs in a number of strue- tural details from that of the Miocene Schizodelphis although the Polk County form Schizodelphis depressus (Allen, 1921) is known solely from fragments of the rostrum. The most obvious peculiarity of the braincase is the protuberant and dorsally elon- evated antero-external portion of the supraorbital process of the frontal. Distortion resulting from dorso-ventral crushing of the brain- case 1s shown by lengthwise folding of the parietal in each temporal fossa, by the downwardly pushed posterior end of each supraorbital process of the frontal, and by the outward displace- ment of the zygomatic process. Except for the portion underly- ine the premanxillary on the dorsal surface of the cranium and abutting against the crest of the supraoceipital, the remainder of the ascending plate of each maxillary including that which over-rides the supraorbital process of the frontal is destroyed. The general form of the slender elongated rostrum (Fig. 1 and Pl. 5, fig. 1) is similar to that of Schizodelphis, the widest portion being at the level of the antorbital notches. In an interval of 180 mm. the rostrum diminishes in transverse diameter from 106 mm. (at level of antorbital notches) to 55 mm.; the attenua- tion of the remainder of the anterior portion of the rostrum is more gradual. The rather shallow antorbital notches are bounded by the lachrymal, the maxillary and the supraorbital process of the frontal. KELLOGG: SKULL OF POMATODELPHIS INAEQUALIS 3) fl) The premaxillaries are somewhat flattened anterior to the antorbital notches for a distance of approximately 60 mm. and then become progressively more convex for an additional 8300 mm. On the distal portion of the rostrum (about 400 mm.) the premaxillaries are more noticeably dorso-ventrally flatten- ed and progressively narrowed. The in- ner borders of the opposite premaxillar- ies came in contact approximately 170 mm. anterior to the antorbital notches and continued closely approximated to the extremity of the rostrum. Unlike the specimen figured and described by Allen (1921, p. 149, pl. 10, figs. 7-8) this rostrum is more symmetrical and con- sequently there is a less obvious asymm- etrical deformation of the premaxillar- ies on the proximal portion. The pre- maxillaries are seemingly fused at the extremity of the rostrum. Posterior to | the level of the antorbital notches, the premaxillaries exhibit the maximum transverse expansion behind the nasal passages. Opposite the nasal passage each premaxillary is elevated above the corresponding maxillary and the outer border is rather indistinctly outlined. The posterior end of each premaxillary is somewhat thickened, irregularly trun- cated and extended backward beyond the nasal bones. On the cranial frag- ment described and figured by Allen (1921, p. 148, pl. 11, fig. 11) the pos- terior ends of the premaxillaries are Fig. 1. Pomatodelphis inaequalis. Dorsal view of skull, restored. Abbrevi- ations: ant.n., antorbital maxillary notch; ¢., occipital condyle; /r., frontal; Max., maxillary; Na., nasal; n.p., nasal passage; Pmz., premaxillary; Sq., squamosal; S.oc., supraoccipital; s. or. pr., supraorbital process of frontal; 2Yg., zY¥gomatie process. 10 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY similarly widened and on both of these specimens a large foramen opens backward under the posterior end of each premaxillary. The premaxillary foramina are large and are located anterior to the level of the antorbital notches. A broad shallow groove extends obliquely backward and inward from each foramen to the internal edge of the premaxillary. With the exception of the proximal 170+ mm., the mesorostral gutter was completely roofed over by the close approximation of the opposite inner borders of the premaxillary bones. The floor and sides of the mesorostral gutter on at least the proximal 150 mim. of the rostrum are constituted by the vomer. In front of the nasal passages the vomer is pinched in to conform with the shape of the presphenoid whose lateral surfaces are sheathed at least in part by this bone between these passages. The rather porous presphenoid is largely destroyed but obviously forms a_ plug at the hinder end of the mesorostral gutter. The preservation of this region does not permit a precise determination of the extent and relations of the mesethmoid. Inasmuch as the anterior wall of the braincase is destroyed in the region where the open- ines for olfactory nerves persist 1m some genera, it can not be assumed that the fused ethmoid bones form a continuous sheet of bone overspreading the posterior walls of the nasal passages dorsally. A V-shaped anterior border of the combined nasal passages is formed by the approximation of the internal edges of the opposite premaxillaries. The lower portion of the posterior wall of each nasal passage is located anterior to the antero-dorsal margin of the nasals, and the dorsal portion of the posterior wall of each passage unfortunately is destroyed. The maxillaries and premaxillaries constitute the major por- tion of the dorsal surface of the skull. For a distance of 85+ mm. in front of the antorbital notches, the outer border of the maxil- lary 1s somewhat compressed dorso-ventrally. In front of this somewhat flattened region, the maxillary from a side view in- creases In dorso-ventral diameter for a short distance, and then the dorsal surface becomes more convex and the outward slope from the premaxillary-maxillary suture steeper. In conformity with the attenuation of the rostrum, the maxillary decreases in breadth and depth anteriorly. On the major portion of the KELLOGG: SKULL OF POMATODELPHIS INAEQUALIS 1] length of the rostrum the maxillary is rather narrow as com- pared with the width of the premaxillaries. Posterior to the antorbital notch the maxillary on most porpoise skulls expands horizontally and overspreads the frontal bones. This thin ascend- ine maxillary plate presumably did not completely overspread the supraorbital process of the frontal; the protuberant antero- external angle of the latter certainly projected beyond it. Around the antorbital notch the maxillary overspreads the underlying lachrymal bone. This flattened lachrymal bone was wedged in between the preorbital portion of the supraorbital process and the overlying ascending plate of the maxillary. The maximum width of the maxillary seems to have been attained opposite the vertex, but this can not be ascertained with certainty im view of the damaged condition of both ascending plates of the maxil- lary above the temporal fossae. Each maxillary is deeply de- pressed posteriorly, the hinder and inner borders curving upward to the crest of the supraoceipital and to the level of the exposed postero-internal portion of the frontal on the vertex. The postero-internal portions of the frontals are broadly ex- posed on the vertex, elevated posteriorly and abut against the transverse crest of the supraoccipital. Allen (1921, p. 148, pl. 11, fig. 11) recognizes an interparietal wedged in transversely be- tween the supraocecipital and the frontals on the cranial fragment referred to this species. The interparietal is not readily recog- nizable as a distinct bone on this Homeland skull. If present, the boundaries of this bone have been obliterated by ankylosis with the frontals. The thin ascending plates of the maxillaries and the corresponding underlying lateral extensions of the frontals may not have completely roofed over the temporal fossae (Fig. 1). Dorsally, on the braincase, each frontal is largely con- cealed by the overspreading ascending plate of the maxillary, the premaxillary, and the nasal, and is suturally united pos- teriorly with the dorsal border of the supraoccipital. The pos- tero-internal portions of the opposite frontals meet edge to edge medially and constitute the vertex which is wider posteriorly than anteriorly and slopes forward from the hinder margin. The thin lateral plate-like extension of each frontal (underlying the corresponding ascending plate of the maxillary) is thickened anteriorly to form the supraorbital process which forms the 12 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY complete osseous roof of the orbit. The antero-external portion of each supraorbital process is conspicuously enlarged and projects strongly upward above the level of the narial openings. The nasals are rather small, somewhat rectangular in shape, wider transversely than antero-posteriorly, slightly indented anteriorly, and markedly thickened dorso-ventrally. The dorso- ventral diameter of the left nasal anteriorly is 19mm. The nasals constitute the upper portion of the posterior wall of the nasal passages. Lateral view. The skull as viewed from the side (Pl. 3) is characterized by a large dorsally prolonged enlargement of the anterior portion of the supraorbital process of the frontal, small orbit, robust dorso-ventrally expanded zygomatic process, flat- tened vertex, and dorso-ventrally compressed rostrum. At the base of the rostrum the depth (60 mm.) is more than half the corresponding width (106+ mm.). For a distance of approxi- mately 120 mm. in front of the antorbital notch the concave external contour of the maxillary is noticeably curved upward. The outward and downward slope of this bone from the pre- maxillary-maxillary suture becomes steeper toward the level of the hindmost alveolus. Near the extremity of the rostrum the external face of the maxillary is almost vertical. Except for the portion posterior to the level of the hindmost alveolus, the maxil- lary is relatively thin throughout the length of the rostrum. Although the terminal end of the rostrum is damaged, the portion preserved suggests that the maxillary extended the full length of the rostrum. The alveoli are not visible from a lateral view. The longitudinal groove which follows the line of contact between the maxillary and premaxillary, commencing at the level of the ninth alveolus and extending almost to the anterior end of the rostrum, is deep and well defined. The dorsal surface of the exposed postero-external portion of the frontal is the highest point on the dorsal profile. The dorsal surface of the vertex slopes forward and downward from the crest of the supraoccipital to the anterior borders of the nasal bones. The premaxillaries slope less strongly downward and the depressed ascending plate of the maxillary slopes more gradually from the supraoccipital toward the antorbital notch. On its external border, the supraorbital process of the frontal is conspicuously dorso-ventrally thickened. The antero-external KELLOGG: SKULL OF POMATODELPHIS INAEQUALIS 13 angle or preorbital portion of the supraorbital process is mark- edly enlarged, and projected upward, terminating in a blunt, rounded, erest-like anterior edge. The posterior angle or post- orbital portion of the supraorbital process is dorso-ventrally compressed, attenuated, and, prior to crushing, overlapping the anterior extremity of the zygomatic process. The maximum antero-posterior diameter of the left supraorbital process is 85 mm. and the corresponding diameter of the orbit 25 mm. Both lachrymal bones were dislodged and lost; a portion of the de- pressed area in which the lachrymal was lodged is preserved on the left side of the skull. The cranium is somewhat dorso-ventrally crushed, resulting in an antero-posterior pressure ridge in the region of the parie- tal-squamosal sutural contact in the left temporal fossa. On the right side the effect of this crushing is more pronounced. Origi- nally, the temporal fossa may have been almost as long as high. Dorsally the temporal fossa is roofed over at least partially by the thin plate-like lateral extension of the frontal that underlies the ascending plate of the corresponding maxillary, and pos- teriorly is bounded by the lambdoid crest that follows the lateral margin of the supraoccipital. In the temporal fossa, the parietal is suturally united anteriorly and dorsally with the frontal, posteriorly with the supraoccipital and ventrally with the squa- mosal. The parietal does not contribute to the formation of the vertex. The laterally flattened zygomatie process is somewhat thicker dorsally than ventrally, obtusely pointed anteriorly, and strongly coneave internally ; the postglenoid portion is relatively short. The greatest length of the left zygomatic process is 74 mm., and the greatest depth anteriorly 42.5 mm. As a result of crushing both supraorbital processes have been pushed downward posteriorly from their original alignment and consequently the postorbital portion of each is depressed below the anterior end of the corresponding zygomatic process and the antero-external portion is compensatorily upturned. Posterior view. The back of the brainease (Pl. 4) displays less evidence of crushing than the lateral walls in the temporal fossae. The supraoccipital is wider than high, concave from side to side, and bounded laterally and dorsally by a flaring lambdoid erest. Externally and ventrally the lambdoid erest follows the contour 14 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY of the posterior border of the squamosal in the temporal fossa and overhangs the exoccipital on each side. The exoccipitals slope backward and outward, and are prolonged ventrally to form the paroccipital process. Each exoccipital is in contact laterally and anteriorly with the squamosal, coalesced dorsally with the supraoccipital and ankylosed internally with the basioccipital. The deep jugular incisure separates the exoccipital from the faleate process of the basioccipital. From a posterior view the left zygomatic process is relatively rather broad, projecting laterally noticeably beyond the extrem- ity of the exoccipital. The foramen magnum was slightly wider than high. The semielliptical occipital condyles are strongly convex dorso- ventrally, less convex transversely and are set off laterally from the exoccipital by a short neck. The external and internal borders of each condyle are sharp edged. Ventral view. The ventral or palatal surface of the attenuated rostrum (PI. 5, fig. 2) is formed entirely by the maxillaries. No indication of sutures to mark the existence of the premaxillaries on the ventral surface of the rostrum was noted. The palate is quite flat except for the shallow median V-shaped groove, formed by the bevelled inner edges of the opposite maxillaries, which commences at the anterior end of the exposed keel of the vomer (approximately 45 mm. anterior to the level of the hindmost alveolus) and continues forward to within 140 mm. of the anterior extremity. Commencing at the level of the anterior end of the exposed keel of the vomer the maxillary widens transversely as the antorbital notches are approached. As seen from the ventral aspect of the rostrum (Pl. 2), the external contour of the basal portion of the maxillary is somewhat convex, and agrees in this respect with the type of Pomatodelphis stenorhynchus. The tooth row, also, terminates posteriorly at approximately the same relative posi- tion as on the type of stenorhynchus, although the hindmost alveoli are somewhat smaller. All of the teeth were lodged in the maxillaries. Near the proximal end of the rostrum the maxillaries are separated by the keel of the vomer which is visible on the ventral surface for a distance of 125 mm. Posteriorly, the keel of the vomer seems to have been concealed, at least partially, by the anterior ends of the palatine bones. KELLOGG: SKULL OF POMATODELPHIS INAEQUALIS 15 Both lachrymal and jugal bones are missing on this skull. The eroove or depression in which the lachrymal was lodged is pre- served, however, on the left side, and this indicates that this bone was wedged in between the anterior border of the supra- orbital process of the frontal and the overlying ascending plate of the maxillary. On this skull the course of the optie nerve from its point of exit from the cranial cavity to the base of the supraorbital process of the frontal is concealed by the external reduplication of the pterygoid. The groove for this optic nerve on the ventral surface of the supraorbital process becomes indistinct near the middle of its outward course. Although the palatine bones (Pl. 2) are incomplete in their present state of preservation the dorsal portion of the left pala- tine is sufficiently well preserved to determine its relations. Each palatine bone is suturally united with the ventral surface of the corresponding maxillary and with the external reduplication of the pterygoid. On skulls of Zarhachis (Kellogg, 1926, pl. 5) and Hurhino- delphis (Kellogg, 1925, pl. 1, p. 16), the internal plate of the pterygoid contributes the lower outer wall of the corresponding nasal passage and to a lesser extent the lower anterior wall. Posteriorly the internal plate of the pterygoid overlaps the basisphenoid laterally and is suturally united with the vomer internally. This relationship seems to have been present on this Florida skull although only a remnant of the internal plate of the pterygoid on the outer wall of the right nasal passage is preserved; the portion of the internal plate of each pterygoid which overlaps the basisphenoid is, however, present. These rather fragile pterygoid bones presumably deteriorated during the period when the skull was being buried by sediments. The plate-like internal portion and the external reduplication were obviously separated by a narrow deep sinus apparently not -ex- ceeding 5 mm. in width. Posteriorly, the ventral surface of each maxillary is sheathed by the external reduplication of the pterygoid laterally and by the palatine ventrally. Neither external reduplication of the pterygoid projects forward to the level of the antorbital notches, and each is suturally united with the corresponding palatine 16 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY anteriorly, the frontal dorsally and the squamosal posteriorly. The anteriorly attenuated deep cavity in front of each nasal passage is bounded medially by the keel of the vomer and lat- erally by the palatine and the external reduplication of the pterygoid. The vomer is horizontally widened posteriorly, sheathing the basisphenoid and meeting the vaginal portions of the internal plates of the pterygoids along the vomer’s lateral margins. Be- tween the nasal passages the vomer is compressed laterally and forms the trough in which the presphenoid rests. The vomer contributes the lower portion of the posterior and internal walls of each nasal passage and, extending dorsally, presumably meets the descending portions of the ethmoids. On this skull the line of union between the basisphenoid and basioccipital can not be ascertained with certainty. On each side of the broad median longitudinal elevation the ventral surface of the basioccipital is irregularly depressed internal to its broad convexly curved descending plates or falcate processes. Each of these plates is directed downward and outward, and anteriorly is united with the vaginal portion of the internal plate of the pterygoid which overlaps the basisphenoid. The deep jugular incisure is located between the internal margin of the exoccipital and the posterior marein of the faleate process of the basioceipi- tal. As pointed out by Allen (1921, p. 150, pl. 11, fig. 12), the occipital condyles of this porpoise are quite large and lack the well developed neck shown on skulls of Schizodelphis sulcatus (Abel, 1899, pl. v1, fie) 13 Daly Piaz, 1903); ple 28). The squamosal is relatively large and its lateral projection forms the forward projecting zygomatic process. The left zygo- matic process is nearly complete; its outer border is curved down- ward and inward, and the ventral articular surface is strongly concave. It should be noted, however, that the hinder external border of this left process is eroded immediately in front of the rather slender ventral postglenoid projection. A relatively nar- row groove for the external auditory meatus, originating at the external margin of the irregular cavity or fossa in which the periotic is lodged, curves outward behind this postglenoid projec- tion. The posterior face of the squamosal abuts against the KELLOGG: SKULL OF POMATODELPHIS INAEQUALIS 17 exoccipital. Behind the above-mentioned groove for the external auditory meatus a short paroccipital process is developed on the ventral border of the exoecipital. The facet on the paroecipital process for articulation with the stylohyal is relatively small and narrow. No teeth are preserved in the alveoli on either side of the rostrum, (Pl. 5, fig. 2). There are, however, 52 vacant alveoli in the left maxillary. The alveoli of the 13 anteriormost teeth, 40th to 52nd counting forward from the hindmost, progressively increase in size, both transversely and antero-posteriorly toward the extremity of the rostrum. These 13 alveoli occupy an interval of 205+ mm. The largest of these anterior alveoli measures 16 mm. antero-posteriorly and 6.5 mm. transversely. Near the middle of the right tooth row, the alveoh vary from 3.5 to 6.5 mm. antero-posteriorly and from 2.5 to 3.5 mm. transversely. The 20 hindmost alveoli in the right maxillary are more closely approximated than in the middle portion of the rostrum. In the right tooth row, counting forward from the hindmost, there are 28 alveoli in 236 mm., and 40 alveoli in 421 mm. Counting for- ward from the 20th alveolus (143 mm. anterior to hindmost), there are 20 alveoli in 278 mm. These middle alveoli are shallow, somewhat slit-like, and more or less contracted at one or both ends. Most of the teeth in the rostrum were seemingly relatively small, with short, laterally compressed roots held in place by the gums. Allen (1921, p. 150), however, observed that the teeth in the upper jaw must have been very small and non-functional if such existed. Internal to the right tooth row is a parallel row of shallow depressions on the proximal portion of the palate which obviously are attributed to the tips of the mandibular teeth. No similar depressions are present on the left side of the rostrum. Three similarly located depressions are present on the right maxillary of the type of Delphinus renovi (= stenorhyn- chus; Van Beneden and Gervais, 1874-80, pl. 57, fig. 9). The rostrum (No. 20738, U.S.N.M.) of a somewhat larger specimen from Polk County, Fla., likewise displays this row of depres- sions on the right side of the rostrum (PI. 5, fig. 3). These two Florida specimens, however, do not confirm the observation of Allen (1921, p. 150) that the width across the lower or mandibu- lar tooth rows was less than that across the upper tooth rows. 18 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY No satisfactory explanation of the presence of these depressions on only one side of the palate is apparent or can be offered until a mandible of this fossil porpoise is found. Measurements of the skull (in millimeters) Greatest length (occipital condyles to extremity of rostrum as preserved) .... BED i De alin rrates Bi: 925 Apex of supraoccipital to extremity of Hee at as peeneeved 875 Posterior end of ascending plate of right maxillary to extremity Of STOStRUM! ASMpPLESCLVCOinsrestet = Gets ce ete ota. tee ee eee 885 Length of rostrum (level of maxillary notches to extremity of THOS THUNONY US OMASAVECL)) oa lb pnwes Suomen eG 40 bald hee 775 Breadth of rostrum at base (between maxillary antorbital notehes) 106 Breadth of rostrum at level of hindmost alveoli ......... 67 Breadth of rostrum at level of alveoli of 20th pair of teeth Count: ING LORWALG strom hin GIN OS tie testes eno k saya eres tere ete 43 Breadth of rostrum at level of alveoli of 40th pair of teeth eee ing TOrWwarG dromuhindmOst) spe ts ee fun 3220 Zygomatic width, estimated ........... 210 Length of left zygomatic process Coevtgienoia’ process 4g erent ) 74 Height of skull, basioccipital to vertex (frontals) .............. 110 Greatest breadth of skull across supraorbital processes, estimated 199+ Dorsal margin of apex of supraoeccipital to anterior margin of left MAS ally ol enol ey eis Peers Gebers sinh ot re ue te Ee ee ae 46 Posterior end of ascending plate of left maxillary to level of hinderwallyof masallppassagesia-.4" 74) 54-24-1445) 2: 53 Distance between outside margins of premaxillaries at lev el of anterior margins of nasal passages eee ere 80 Greatest breadth of left premaxillary at level of antorbital note ; 33 Maximum distance between inner margins of maxillaries at vertex 36 Greatest antero-posterior diameter of right supraorbital process |. 87 Greatest length of exposed portion of frontal on vertex __. 25 Greatest antero-posterior diameter of right nasal : 15 Transverse diameter of right nasal ............ CHS (oe: 14.5 Greatest transverse diameter of the two nasals ............... 31.5 Distance from dorsal margin of supraoccipital to dorealle margin of OUAMEN, TM ACTMM Re ae ye cies Aces ties, see AACR RTC Cae ec : 91 Height Oi toramenyumea fmm Gao. eee] eee ae err eee 21 Wadthvot oramenmaenum ay. 2... ae eee ee ee eee ae 3.5 Greatest distance between outside margins of occipital sonavics 88 Greatest height of right occipital condyle ...... aS ee 48 Greatest breadth of right occipital condyle ....... 39 KELLOGG: SKULL OF POMATODELPHIS INAEQUALIS 19 Distance between inner margins of alveoli of hindmost teeth in ‘opposifemmascillaries!: -piarcec i. eto eeeas ster) eRe es hee 47 Distance between inner margin of 20th alveolus in right wieisilleesy and 20th alveolus in left maxillary (counting forward from HIN GMOSt)) _ ..k..4 3.5 UR ar CEES ane CI bao: SETS A Se 24.5 Distance from left antorbital notch to posterior margin of hind- most alveolus in left maxillary ...... er 154 Rostrum, left side, distance between margins of aiveol, counting backward from anterior extremity. Between 4thiandy othe. Hee eee. 8 mm. Between Sthrands6thi= 4. 4.0.4... : 9mm. Between’ 6th and’ 7th) 55 5...5.227 i. 9mm. Rostrum, left side, antero-posterior diameter of alveoli, counting backward from anterior extremity. bth alveolushes a aes at AL: 8.5 mm. Gthevalveolus@h cw soul ae eee 10.5 mm. Tbh wALVIGOUS, 2 sn Bae cick yee Slee ees 8.7 mm. POMATODELPHIS STENORHYNCHUS For reference purposes the following information is here re- corded regarding the type specimen of Pomatodelphis stenorhyn- chus (Holl). Type. No. 2228, Laboratoire de Paléontologie, Museum Na- tional d’Histoire Naturelle, Paris. Rostral portion of right maxillary and right premaxillary. Département de 1’Orne (=Maine-et-Loire), France. Caleaire grossier. Measurements of type of Pomatodelphis stenorhynchus (in millimeters). Maximum total length as preserved (essentially as figured by Cuvier, 1823, pl. 23, fig. 38, at 4% reduction) ......... Be Transverse diameter of palatal surface of maxillary at proximal end 66.5 Transverse diameter of palatal surface of maxillary at distal end just posterior to broken end ............. 26.7 Dorso-ventral diameter of maxillary and premaxillary orice at distal end just posterior to broken end .... 29.8 Dorso-ventral diameter of maxillary and premaxillary combined at PLOXAm AL BENS o..cg5 acme Le Beton See é . 84.7 Transverse diameter of premaxillary at proximal end . Pere PON Transverse diameter of premaxillary at distal end : 13.5 20 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY Transverse diameter of maxillary at level of posterior alveolus ... 40 Hindmost five alveoli occupy an interval of ............../°.....) 39 Hight anterior alveoli oceupy an interval 6f eee eau, 83.5 Antero-posterior diameter of hindmost five alveoli, about: 7.2 Transverse diameter of hindmost five alveoli, about . A Bel 6.5 Originally 20 alveoli were present in this section of the palatal surface of the right maxillary. Referred Specimen II Referred specimen. No. 20738, Division of Vertebrate Paleon- tology, United States National Museum. Incomplete rostrum. Collected by Z. S. Altschuler and E. T. Young, U. 8. Geological Survey, Florida Land Pebble Phosphate Area. February 23, 1954. Horizon and Locality. Three miles south of Bartow, Polk County, Florida. Noralyn Mine, center NE 14 Sec. 31, T 30 §, R 25 E. In hght tan, medium to coarse, slightly Canes. quartz, phosphate sand; in situ in face of active mine pit approximately 8 feet above Hawthorn Ls. This incomplete rostrum (PI. 5, fig. 3) lacks the terminal por- tion and the outer border on both sides between the level of the hindmost alveolus and the antorbital notch. It represents a some- what larger individual than the skull from the Homeland Mine, the transverse diameter at the level of the hindmost alveoli exceeding the latter by more than 10 per cent. Attenuation of the rostrum may have been rather gradual. The dorso-ventral compression of the rostrum is, however, comparable to that of the specimen from the Homeland Mine. The palatal surface of both maxillaries is flattened except for the narrow longitudinal median groove formed by their bevelled inner borders. In contrast to the Homeland skull, the keel of the vomer is not visible between the maxillaries on the palatal surface of the rostrum. Thirty-one alveoli are present in the right maxillary, (Pl. 5 fig. 3). For purposes of comparison it should be noted that the 30 posterior alveoli on this rostrum occupy an interval of 422 mm., and on the Homeland Mine specimen these same 30 alveoli are located in an interval of 255 mm. The alveoli for the ten hindmost teeth are placed obliquely to the axis of the major KELLOGG: SKULL OF POMATODELPHIS INAEQUALIS 21 portion of the tooth row and are also more closely spaced than the alveoli anterior to them. Toward the anterior end of this rostral fragment, the alveoli (although more or less slit-like and relatively shallow) progressively increase shghtly in antero- posterior diameter. The interval between the alveoli increases irregularly from a minimum of 5 mm. to a maximum of 12mm. At the posterior end of the palate the opposite tooth rows are separated by an interval of 68 mm. and by 29 mm. at the anterior end of the rostral fragment. Commencing at the level of the 5th alveolus on the right side and terminating at the 17th, there are 19 depressions, variable in depth and more or less parallel to the tooth row, which are presumed to represent pits for the tips of the teeth in the corresponding portion of the mandible. Both premaxillaries were fractured and detached from the proximal portion of the rostrum when the specimen was un- covered in the mine pit. One section of the right premaxillary measures 420 mm. in length and a shorter more anterior piece 154 mm. A section of the left premaxillary 194 mm. in length was also recovered. These premaxillary bones, although larger, agree in general conformation and attenuation with the corresponding bones of the skull (MCZ 4433) from the Homeland Mine. The internal face of each of these premaxillaries is flattened, the ventral face longitudinally grooved and the dorsal surface con- vexly curved. Measurements (in millimeters) of rostrum No. 207388 U.S.N.M. Total length of rostrum as preserved : 579 Breadth of rostrum at level of hindmost alveoli 75 Breadth of rostrum at level of alveoli of 20th pair of feath count- ing forward from hindmost . “f, 52 Breadth of rostrum at level of alveoli of 30th - pair of teeth count- ing forward from hindmost ........ Bag Uke ya 45 Anterior end of palatine to level of fandmost aly COlie ea eeet: 68 Hindmost 5 alveoli in right maxillary in an interval of . 38 Hindmost 10 alveoli in right maxillary in an interval of 88 Hindmost 20 alveoli in right maxillary in an interval of . 230 Distance between inner margin of 20th alveolus in right maxillary and 20th alveolus in left maxillary, counting forward from himdmostiees ee ce Hoes t Boro Antero-posterior diametes of sth Al ening’ fete maxillary aby se 5. Co 1 22 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY Transverse diameter of 5th alveolus, right maxillary ............. 3.5 Antero-posterior diameter of 19th alveolus, right maxillary a: 6 Transverse diameter of 19th alveolus, right maxillary ........... 225 Referred Specimen III Referred specomen. No. 6688, Division of Vertebrate Paleon- tology, United States National Museum, anterior extremity of rostrum ; and No. 6684, anterior extremity of symphysis of man- dibles. Received from H. B. Lunn, August 3, 1910. Horizon and locality. Christina, about 644 miles south of Lakeland, Polk County, Florida. Land Pebble Phosphate Area. The anterior ends of the rostrum (No. 6683, U.S.N.M.) and of the symphyseal portion of the mandibles (No. 6684, U.S.N.M.) of a larger individual show more clearly the details of the large anterior alveol. Although the right side of the rostrum of the previously deseribed skull (No. 4483, M.C.Z.) is damaged exter- nally, the indentations suggest that originally eight large alveoli were present and that these anterior larger alveoli progressively decrease in size in the antero-posterior direction at least to the 15th and 16th alveoli (counting backward from the anterior extremity ). Six essentially complete large alveoli as well as a portion of the seventh alveolus are present on the left side of this anterior end of the rostrum (PI. 6, fig. 1). The terminal portions of the roots of the teeth are present in the first and fourth alveoli on the right side (counting backward from the anterior end) and also in the second alveolus on the left side. The ventral surface of the dorso-ventrally compressed rostrum is more or less flat- tened. No distinct line of separation between the right and left sides of the rostrum is discernible on the ventral surface. The outer face of the rostrum is slightly indented between the alveoli. All of the alveoli are large and are separated by slightly de- pressed wide intervals. The attenuated root of each tooth extends backward below this interval, and the lateral compression of the basal portion of the alveolus indicates that the root was somewhat flattened transversely. The teeth in the anteriormost pair of alveoli may have projected more forward than downward. The dorsal surface of this anterior portion of the rostrum (PI. 6, fig. 3) is somewhat convex. The narrow premaxillaries are anky- KELLOGG: SKULL OF POMATODELPHIS INAEQUALIS 23 losed, although a thin longitudinal groove marks the median line of contact. Externally a longitudinal groove which tends to become indistinct some 35 mm. posterior to the anterior extremity delimits each premaxillary. Whether or not the premaxillaries expanded and contributed the anterior end of the rostrum cannot be determined in view of the coalescence of the bones at the extremity. On the dorsal face of the left maxillary four shallow grooves (originating in the above described longitudinal groove), following the lateral border of the corresponding premaxillary, curve downward to the alveolar margin. The dorsal surface of this rostrum is also characterized by the presence of numerous thin curved grooves which suggest that the extremity at least was covered by a tight fitting skin. he maximum length of this rostral fragment is 125 mm.; width anteriorly 28.5 mm.; and width posteriorly, 37 mm.; dorso-ventral diameter anteriorly, 15 mm.; and dorso-ventral diameter posteriorly, 17.5 mm. The anterior extremity of the symphyseal portion of the mandibles is dorso-ventrally flattened, rounded at the end, and very gradually attenuated. Although the mandibles are firmly ankylosed along the symphysis (Pl. 6, fig. 2), a shallow longi- tudinal groove marks the contact of the opposite rami dorsally. On the ventral surface the line of contact is less clearly de- mareated. Irregular grooves of varying width and depth and directed for the most part anteriorly are present on the ventral surface (Pl. 6, fig. 4). Two nutrient foramina are recognizable on the ventral surface, and some of the numerous narrower grooves apparently anastomose. The dorsal surface of the sym- physis is essentially flat transversely. Seven large alveoli are present on the left side of this symphyseal fragment. All of the teeth with the exception of the 5th and 6th were dislodged from the left ramus and portions of the roots of four teeth are lodged in alveoli in the right ramus. The teeth were implanted opposite one another, and were separated by relatively wide intervals from preceding and succeeding teeth. The root of each tooth extended backward below this interval. The anterior- most alveolus at the extremity of both rami is shghtly smaller than the preceding alveolus. These large alveoli slope more back- ward than inward. The distance between opposite alveoli at the second pair counting backward from the anterior extremity is 10.5 mm. and at the fifth pair 19 mm. In the left ramus five 24 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY alveoh were located in an interval of 95.5 mm. The roots of the two teeth in situ are conspicuously enlarged below the crown (which apparently was relatively small), transversely com- pressed, bent backward, and attenuated at the extremity. The outer face of each ramus is indented between the alveoli, sug- gesting that the opposing teeth on the terminal portion of the rostrum bite between these terminal mandibular teeth. The maximum length of this symphyseal fragment is 133 mm.; width anteriorly, 26 mm.; width posteriorly, 37.5 mm.; dorso-ventral diameter anteriorly, 15 mm.; dorso-ventral diameter posteriorly, 17.5 mm. Measurements of rostrum, no. 6683, U.S.N.M. Rostrum, distance between margins of alveoli, counting backward from anterior extremity. Right side Left side Between Ist and 2nd alveoli 6 6.5 mm. Between 2nd and 3rd alveoli 9 10) mm. Between 3rd and 4th alveoli 9.5 8.5 mm. Between 4th and 5th alveoli 9.5 8.2 mm. Between 5th and 6th alveoli 9 9.5 mm. Between 6th and 7th alveoli x 10 mm. Rostrum, antero-posterior diameter of alveoli, counting backward from anterior extremity. 1st 2nd 3rd 4th Sth 6th alveolus alveolus alveolus alveolus alveolus alveolus Right side Left side x 13.4 mm. 11 mm. 10.3 mm. 10 mm. 9 mm. Measurements of mandibular svmphysis, no 6684, U.S.N.M. Anterior end of mandibular symphysis, distanee between margins of alveoli, counting backward from anterior extremity. Between Ist and 2nd alveoli Between 2nd and 3rd alveoli Right 1.5 6 Left 2 mm, 6 mm. KELLOGG: SKULL OF POMATODELPHIS INAEQUALIS 25 Right Side Left Side Between 3rd and 4th alveoli 4a 8.2 8.3 mm. Between 4th and 5th alveoli iA 10 7.7 mm. Between 5th and 6th alveoli . Sy 9 6.5 mm. Between 6th and 7th alveoli - x 8.7 mm. Anterior end of mandibular symphysis, antero-posterior diameter of alveoli, counting backward from anterior extremity. Right Left Nistamalvi@OlUsna es to.cewaci escent 2 Gronks Seale ae Tal 8.8 mm. 2nd alveolus _. it Aer etal sine Sena: Geet ia lap 11.8 mm. Srdevalveolus’ ss eee cee oe aR ee 12 12.4 mm. AthmalveOluis™ sepa aac vere ce ae Mets, ee es 12.5 mm. thy valivieOlus: sass. ce ae PE RTT SON aN 11 11.5 mm. Gthalveolus}=4.500 000.050: Ralston ee hs xx 10.5 mm. LITERATURE CITED ABEL, O. 1899. Untersuchungen iiber fossilen Platanistiden des Wiener Beckens. Denkschr. kais. Akad. Wiss. math. nat. Cl. Wien, vol. 68, pp. 839-874, 4 pls. ALLEN, G. M. 1921. Fossil cetaceans from the Florida phosphate beds. Journ. Mam- malogy, vol. 2, no. 3, pp. 144-159, pls. 9-12. 1941. A fossil river dolphin from Florida. Bull. Mus. Comp. Zool., Harvard College, vol. 89, no. 1, pp. 3-8, 3 pls. Cask, E. C. 1934. third,-05 mm-:; fourth; 0:23° mm: The only known specimen of this species is in very poor con- dition; it had been determined as T. biolleyi by Banks. Diagnosis. This species differs from 7. everum in the sub- tegulum of the palpus lacking the median notch and in the embolus having an ectal projection (Fig. 137). THERIDION EVEXUM [eyserline Fieures 124-135; Map 5 Theridion evexrum Weyserling, 1884, Die Spinnen Amerikas, vol. 2, pt. 1, p. 65, pl. 3, fig. 39, 9 (9 type from N. Granada, in the British Mu- seum, Natural History). Roewer, 1942, Katalog der Araneae, vol. 1, p. 492. Phyllonethis electa O. P. Cambridge, 1895, Biologia Centrali Americana, Araneidea, vol. 1, p. 165, pl. 20, fig. 2, 9, ?fig. 3 9 (2 type from 98 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY Omilteme, Guerrero, in the British Museum, Natural History). Theridion electum, F. P. Cambridge, 1902, ibid., vol. 2, p. 390, pl. 36, fig. lly Se Theridion cabriolatum Franganillo, 1930, Mas Aracnidos Nuevos de Cuba, p. 12, fig. 4,9 (Q types from Sierra Maestra, Cuba). Roewer, 1942, Katalog der Araneae, vol. 1, p. 490. NEW SYNONYMY. Comment. This large brightly colored species has a dorsal abdominal pattern like 7’. pictipes. The venter of the abdomen has a white spot framed by black stripes similar to the female Map 5. Distribution of Theridion everum Keyserling. believed to be 7. omiltemi (Fig. 174). The long lees have black rings like Chrysso cumbridgei. It differs from the latter species in having an abdomen of typical shape. The epigynum is a de- pression (Figs. 125, 127) of variable shape. There is consider- able variation in the shape of several palpal sclerites; however, the embolus is similar in all males examined, and the subtegulum has a characteristic notch (Figs. 129, 131, 133, 1385). Records. Michoacan: Jan-Itaro, 2 (Hoogstraal). Hidalgo: Chapulhuacan, ¢ (W. J. Gertsch). Puebla: Villa Juarez, ¢ (H. M. Wagner). Veracruz: La Planta de Moctezuma nr. For- tin, @ (C. and M. Goodnight). Guerrero: Omiltemi (QO. P. Cambridge, 1895). Guatemala: Los Amates, 2, 6 (Kellerman). El Salvador: San Salvador (Kraus, 1955). Panama: El Vol- LEVI: ACHAEARANEA, THERIDION AND SPHYROTINUS 99 ean, 2, 6 (A. M. Chickering; W. J. Gertsch). Cuba: Mts. N. of Imias, 3-4000 ft., @ ; Trinidad Mts., 2500-3500 ft., 6 ; S. side of Pico Turquino, 2°, ¢ ; Sierra del Cobre, 3000-3800 ft., @ ; 5 mi. EB. of Soledad, Oriente, ? (all P. J. Darlington) ; Mina Carlota, Trinidad Mts., 2, 4 (Salt and Myers); Sierra del Cobre, 2600-3325 ft. (S. C. Bruner). Colombia: N. Granada (Keyserling, 1884) ; Boyaca Prov.: Rio Upia, 850-950 m, ¢. Venezuela: Rancho Grande, 2 (W. Beebe). Brazil: Est. Rio de Janeiro: Petrépolis, 850-1000 m; @ (H. Sick) ; Cidade Sao Paulo, @ (H. Sick). THERIDION RICENSE, new species Figures 138-141 Type. Male holotype from El Yunque, Puerto Rico, Feb. 27- 28, 1955 (A. M. Nadler), in the American Museum of Natural History. Description. Carapace yellow-white with a wide black band in female, a dusky band in male. Sternum, legs yellow-white. Legs with indistinct dark marks. Abdomen yellowish, dorsum spotted with white pigment. Anterior median eyes slightly smaller than others, a little more than a diameter apart, their radius from laterals. Posterior eyes slightly less than their diameter apart. Total length of female 3.8 mm. Carapace 1.4 mm. long, 1.1 mm. wide. First femur, 3.4 mm.; patella and tibia, 3.5 mm.; metatarsus, 3.6 mm.; tarsus, 0.9 mm. Second patella and tibia, 2.1 mm.; third, 1.2 mm.; fourth, 2.4 mm. Total length of male 2.9 mm. Carapace 1.2 mm. long, 1.0 mm. wide. First femur, 3.5 mm.; patella and tibia, 3.6 mm.; meta- tarsus, 3.6 mm.; tarsus, 1.0 mm. Second patella and tibia, 2.1 mm.; third, 1.1 mm.; fourth, 2.2 mm. Diagnosis. The epigynum which has a small oval depression, wider than long (Fig. 139), and the median apophysis and tegu- lum of the palpus (Figs. 140, 141) separate this species from others of the 7. frondeum group. THERIDION LEONES, hew species Figures 148-151 Type. Male type from Desierto de los Leones, Distrito Fed- eral, Mexico, May 26, 1946 (J. C. and D. L. Pallister), in the American Museum of Natural History. 100 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY Description. Carapace yellow-white with a dusky median stripe which becomes as wide anterior as posterior median eyes; border dusky. Sternum, legs yellow-white ; legs with dusky rings on middle and ends of segments. Abdomen with white median dorsal band whose sides are wavy ; sides of dorsum black to gray ; sides, venter white. Anterior median eyes one diameter apart, two-thirds from laterals. Posterior median eyes one diameter apart, one and one-third from laterals. Total length of a female, 3.38 mm. Carapace 1.2 mm. long, 1.1 mm. wide. First femur, 1.9 mm.; patella and tibia, 2.0 mm.; metatarsus, 1.7 mm.; tarsus, 0.6 mm. Second patella and tibia, 1.8 mm.; third, 0.8 mm.; fourth, 1.4mm. Total length of a male 2.2 mm. Carapace 1.1 mm. long, 0.9 mm. wide. Seeond patella and tibia, 1.8 mm.; third, 0.8 mm.; fourth, 1.8 mm. Diagnosis. The epigynum has a circular to wider than long depression whose lateral walls are heavily sclerotized (Fig. 149). The shorter legs, lack of black mark on the sternum of the female and the shape of the tegulum of the male palpus (Fig. 151) separate this species from 7. contreras. Records. Distrito Federal. Desierto de los Leones, 2, 4 paratypes (J. C. and D. L. Pallister; C. Goodnight, F. Bonet and Bolivar). THERIDION CONTRERAS, new species Figures 144-147 Type. Male type from Contreras, Distrito Federal, Mexico, 2600 m., July 23, 1947 (H. Wagner), in the American Museum of Natural History. Description. Carapace, sternum, lees whitish in female, darker yellowish in male. Carapace with head dusky. Sternum with a median black line on posterior portion. Legs with distal ends of segments yellow. Abdomen yellow-white, with a median dor- sal white band. Male with some black pigment on abdomen. Anterior eyes slightly larger than others in female, subequal in male, their diameter apart, two-thirds from laterals. Posterior median eyes one diameter apart, one and one-half from laterals. Total length of female 3.4 mm. Carapace 1.8 mm. long, 1.1 mm. wide. First femur, 2.7 mm.; patella and tibia, 2.9 mm.; meta- tarsus, 2.6 mm.; tarsus 0.7 mm. Second patella and tibia, 1.8 LEVI: ACHAEARANEA, THERIDION AND SPHYROTINUS 101 mm.; third, 1.0 mm.; fourth, 1.8 mm. Total length of male 2.7 mm. Carapace 1.3 mm. long, 1.2 mm. wide. First femur, 3.1 mm. ; patella and tibia, 3.5 mm.; metatarsus, 3.0 mm.; tarsus, 0.9 mm. Second patella and tibia, 2.2 mm.; third, 0.9 mm.; fourth, 1.8 mm. Diagnosis. The shape of the tegulum of the male palpus (Fig. 147), the longer legs, dusky mark on the sternum and wider eplgynum (Fig. 145) differentiate this species from 7. leones. Records. Durango: 6 mi. NE. of El Salto (W. J. Gertsch). Hidalgo: Apuleo (H. Wagner). Distrito Federal: Santa Rosa, 9 paratypes (H. Wagner) THERIDION METABOLUM Chamberlin and Ivie Figures 152-158 Theridion niveum, Banks, 1929, Bull. Mus. Comp. Zool., vol. 69, p. 84 (err. det.). Theridion metabolum Chamberlin and Ivie, 1936, Bull. Univ. Utah, biol. ser., vol. 3, no. 5, p. 33, pl. 10, fig. 79, 9 (92 type from Barro Colorado Island, Panama, in the American Museum of Natural History). Description. Carapace white. Sternum black. Legs white with black rings. Abdomen white with dorsal white and black spots (Fig. 154), venter gray except for black genital area. An- terior median eyes one diameter apart, two-thirds from lat- erals. Posterior eyes one diameter apart. Total length of females 2.3-3.8 mm. A female measured total length 2.9 mm. Carapace 1.1 mm. long, 1.0 mm. wide. First femur 3.1 mm.; patella and tibia, 3.2 mm.; metatarsus, 3.0 mm.; tarsus, 1.0 mm. Second patella and tibia, 1.7 mm.; third, 0.9 mm.; fourth, 1.6 mm. A male measured total lenreunt 2.1 mm. Carapace 1.1 mm. long, 0.8 mm. wide. First femur, 3.2 mm.; patella and tibia, 3.2 mm.; metatarsus, 3.0 mm.; tarsus, 1.0 mm. Second patella and tibia, 1.7 mm.; third, 0.9 mm.; fourth, 1.5 mm. Records. Panama: El Valle (A. M. Chickering). Canal Zone: Experimental Gardens; Summit; Forest Reserve; Madden Dam; Ft. Sherman; Barro Colorado Isl., very common (all A. M. Chickering). THERIDION NIVEUM QO. P. Cambridge Figures 162-167 Theridion niveum O. P. Cambridge, 1898, Biologia Centrali Americana, Araneidea, vol. 1, p. 255, pl. 34, fig. 11, 9 (@ type from Bugaba, 102 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY Panama, in the British Museum, Natural History). F. P. Cambridge, 1902, ibid., vol. 2, p. 390, pl. 37, figs. 1, 2, 29, 6. Roewer, 1942, Katalog der Araneae vol. 1, p. 496. (Not 7. nivewn, Petrunkevitch, 1930, Trans. Connecticut Acad. Scei., vol. 30, p. 204, figs. 51, 52, err. det.) Description. Carapace, sternum, legs yellow. Abdomen yel- lowish white, dorsum with white pigment, sometimes with an anterior median black spot or black spots on sides. Anterior median eyes one and one-half diameters apart, one from laterals. Posterior median eyes one diameter apart. Chelicerae with two teeth on anterior margin. Epigynum (Figs. 163, 164) a pro- jeecting knob having an opening on each side. Measurements of female total length, 2.5 mm. Carapace, 1.0 mm. long. First femur, 2.0 mm.; patella and tibia, 2.1 mm.; metatarsus, 1.9 mm. ; tarsus, 0.8 mm. Second patella and tibia, 1.1 mm.; third, 0.7 mm.; fourth, 1.3 mm. Total length of male 1.6 mm. Carapace, 0.8 mm. long. First femur, 2.4 mm.; patella and tibia, 2.5 mm.; metatarsus, 2.5 mm.; tarsus, 0.7 mm. Second patella and tibia, 126.mm.- third..0:8 mm.: fourth, 1.3 mm: These descriptions are from males and females from Teapa, Mexico, determined by F. P. Cambridge. The specimens were in poor condition, having probably once been dry. A male of T. trepidum determined by F. P. Cambridge proved to be the same species. Natural History. This species has been collected on bromeliads in Veracruz by Bordas. Records. San Luis Potost: 23 mi. S. of Ciudad de Valles (A. M. and L. I. Davis); Xilitla (R. Dreisbach). Veracruz: Martinez de la Torre, 45 m. (H. Wagner) ; Cérdoba (J. C. and D. L. Pallister) ; Tlacotalpan (H. Wagner) ; La Buena Ventura (A. Petrunkevitch) ; Tuxtilla (H. Wagner); Mantla; Yanaa (Bordas). Guerrero: Amula, Omiltemi (O. P. Cambridge). Oaxaca: Papaloapan (H. Wagner). Chiapas: El Real (C. and M. Goodnight); Puerto Madero (N. L. H. Kraus). THERIDION EXCAVATUM FEF. P. Cambridge Figures 206-207 Theridion excavatum F. P. Cambridge, 1902, Biologia Centrali Americana, Araneidea, vol. 2, pl. 37, fig. 5, 9 (9 type from Guatemala, in the British Museum, Natural History). Roewer, 1942, Katalog der Ara- neae, vol. 1, p. 492. LEVI: ACHAEARANEA, THERIDION AND SPHYROTINUS 103 THERIDION UNCATUM F. P. Cambridge Figures 203-204 Theridion uncatum F. P. Cambridge, 1902, Biologia Centrali Americana, Araneidea, vol. 2, p. 390, pl. 37, fig. 3, ¢ (¢ type from Omilteme, Guerrero, in the British Museum, Natural History). Roewer, 1942, Katalog der Araneae, vol. 1, p. 500. THERIDION STYLIGERUM FE’. P. Cambridge Figures 168-171 Theridion styligerum F. P. Cambridge, 1902, Biologia Centrali Americana, Araneidea, vol. 2, p. 389, pl. 36, fig. 24, ¢ (é type from Guatemala, in the British Museum, Natural History). Description. Carapace yellow-white with a median dusky band which splits in two anteriorly, each branch touching a posterior median eye. Legs yellow-white, banded in male. Abdomen yel- low-white, dorsum white with two black spots above spinnerets ; often a pair of black spots on venter. Anterior median eyes one and one-half diameters apart, one and three-quarters from lat- erals. Posterior median eyes one diameter apart, one and three- quarters from laterals. Total length of female 3.2 mm. Carapace 1.3 mm. long, 1.2 mm. wide. First femur, 2.6 mm.; patella and tibia, 2.7 mm.; metatarsus, 2.7 mm.; tarsus, 0.9 mm. Second patella and tibia, 1.6 mm.; third, 1.1 mm.; fourth, 1.9 mm. The epigynum has an oval depression (Fig. 169); the embolus of the palpus is enlarged (Fig. 171). Natural History. This species has been found in oak forest at 7200 ft. in Veracruz. Records. Durango: 30 mi. NE., 10 mi. E., 6 mi. NE. of El Salto (W. J. Gertsch). Veracruz: nr. Acultzingo (C. Good- night). Hidalgo: Apulco (H. Wagner) ; 10-20 mi. 8. of Jacala (V. Roth, W. J. Gertsch). Distrito Federal: Santa Rosa (H. Wagner); Contreras, 2600 m. (H. Wagner); Xochimileo (C. Goodnight; H. Wagner); Coyoacan (H. Wagener); Tlalpam, 2300 m. (H. Wagner). Puebla: Teciutlan (C, and P. Vaurie). Morelos: nr. Cuernavaca, 8700 ft. (R. Dreisbach). Oaxaca: (B. Malkin). Guatemala: Chichicastenango (C. and P. Vaurie) ; Nebaj (C. and P. Vaurie). 104 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY ‘THERIDION TREPIDUM QO. P. Cambridge Figures 177-180 Theridion trepidum O. P. Cambridge, 1898, Biologia Centrali Americana, Araneidea, vol. 1, p. 256, pl. 34, fig. 10, ¢ (4 type from Montafia de Chilase6, Guatemala, in the British Museum, Natural History). F. O. P. Cambridge, ibid., 1902, vol. 2, p. 391, pl. 37, fig. 4, ¢. Roewer, 1942, Katalog der Araneae, vol. 1, p. 499. Phyllonethis trepidum O. P. Cambridge, 1902, Biologia Centrali Americana, Araneidea, vol. 1, p. 307, pl. 39, fig. 9, @. Description. Carapace, sternum, legs yellow white; carapace with a median dusky double line. Legs of male with dark rings. Dorsum of abdomen white usually with a pair of black bands, irregular in outline, about their width apart. Anterior median eves one and three-quarters diameters apart, one diameter from laterals. Posterior eyes one and one-quarter diameters apart. Anterior median eyes shghtly smaller than others. Total length of female 2.4 mm. Carapace 0.85 mm. long, 0.75 mm. wide. First femur, 2.16 mm.; patella and tibia, 2.18 mm.; metatarsus, 2.10 mm.; tarsus, 0.69 mm. Second patella and tibia, 1.28 mm.; third, 0.75 mm.; fourth, 1.37 mm. The epigynum has an oval depression (Fig. 178). The male ean be identified by its palpus (Figs. 179, 180). Records. Hidalgo: Rancho Viejo nr. Jaecala (C. and M. Good- night). Distrito Federal: Santa Rosa (H. Wagner) ; Desierto de los Leones (C. Goodnight). Puebla: ?Tehiutaitlan (H. Wag- ner). Oaxaca: Oaxaca. Chiapas: Tenejapa (C. Goodnight) ; Las Casas (C. and M. Goodnight and L. Stannard) : San Crist6- bal de las Casas (C. and M. Goodnight; H. Wagner). Panama: El Volean (W. J. Gertsch). THERIDION MARTINI, hew species Figures 181-182 Type. Male type from Volcan San Martin, near San Andrés, Veracruz, 5000 ft., July 14, 1948 (C. J. Goodnight), in the American Museum of Natural History. Description. Carapace, sternum, legs bright yellow. Abdomen light gray with four small dark spots on dorsum. Anterior me- dian eyes slightly smaller than others, less than one diameter apart, one-third from laterals. Posterior median eyes one diame- LEVI: ACHAEARANEA, THERIDION AND SPHYROTINUS 105 ter apart, two-thirds from laterals. Total length of male 2.5 mm. Carapace 1.3 mm. long, 1.1 mm. wide. First femur, 2.3 mm.; patella and tibia, 2.7 mm.; metatarsus, 2.1 mm.; tarsus, 0.9 mm. Second patella and tibia, 1.8 mm.; third, 1.3 mm.; fourth, 1.7 mm. Diagnosis. The heavy median apophysis of the palpus (Figs. 181, 182) separates this species from others of the 7. frondewm croup. THERIDION OMILTEMI, new species ? Figures 172-176 Theridion evexum, F. P. Cambridge, 1902, Biologia Centrali Americana, Araneidea, vol. 2, p. 389, pl. 36, fig. 25, 6, 2 (err. det.). Type. Male type from Omiltemi, Guerrero, in the British Museum (Natural History). Description. Carapace brownish yellow. Chelicerae brown. Maxillae, sternum dark brown. Legs yellow with dark spots on distal ends of segments. Abdomen white with an anterior median dorsal black patch and a pair of black patches on posterior which are continuous with black venter (Fig. 174); venter of male dusky yellow. Anterior median eyes one diameter apart, slightly more in female, two-thirds their diameter in male. Posterior median eyes almost one diameter apart, one and one-third from laterals. Total length of female 5.8 mm. Carapace 2.2 mm. long, 1.4 mm. wide. First femur, 6.3 mm.; patella and tibia, 5.5 mm.; metatarsus, 6.8 mm.; tarsus, 1.5 mm. Second patella and tibia, 3.6 min.; third, 2.2 mm.; fourth, 4.0 mm. Total length of male 3.6 mm. Carapace 1.7 mm. long, 1.2 mm. wide. First femur, 5.0 mm.; patella and tibia, +.3 mm.; metatarsus, 5.8 mm.; tarsus, 1.3 mm. Second patella and tibia, 2.7 mm.; third, 1.4 mm.; fourth, 2.5 mm. Differences in coloration might suggest that the female does not belong to the male. Diagnosis. The median apophysis of the palpus (Fig. 175) distinguishes this species from 7. evexrum. Record. Guatemala: San Jerénimo (C. and P. Vaurie). THERIDION GRECIA, new species Figures 185-186 Type. Female type from Grecia, Costa Rica, Nov. 22, 1955 (B. Malkin), in the American Museum of Natural History. 106 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY Description. Carapace whitish with a median dusky line. Legs whitish with distal end of first tibiae and middle and distal end of metatarsi black. Abdomen yellow-white with a median dorsal white band and a white band on each side. Anterior me- dian eyes a little more than one diameter apart, one diameter from laterals. Posterior eyes slightly more than one diameter apart. Total length of female, 2.9 mm. Carapace, 1.0 mm. long, 0.9 mm. wide. First femur, 2.2 mm.; patella and tibia, 2.2 mm.; metatarsus, 1.9 mm.; tarsus, 0.7 mm. Second patella and tibia, 1.2 mm.; third, 0.9 mm.; fourth, 1.5 mm. Diagnosis. The epigynum whose openings are on the anterior face of a heavily sclerotized slight elevation (Fig. 186) dis- tinguishes this species from others of the frondeum group. Records. Nayarit: Tepie (C. and M. Goodnight). Chiapas: nr. Rio San Gregorio, betw. Comitan and Ocotal (C. and M. Goodnight) ; 30 km. NE. of Cintalapa (H. Wagner). Costa Rica: Grecia, ° paratype (B. Malkin). Panama: Fl Valle (A. M. Chickering). Venezuela: Botanical Gardens, Caracas (A. M. Nadler). THERIDION BIOLLEYI Banks Figures 195-196 Theridion biolleyi Banks, 1909, Proc. Acad. Nat. Sci. Philadelphia, vol. 61, p. 204, pl. 6, fig. 26, 9 (92 type from Tablazo, Costa Rica, in the Museum of Comparative Zoology). Description. Carapace yellow-white with a dusky median band, as wide as eyes in front with a narrow dusky cross band. Sternum dusky. Lees yellow-white with bands. Abdomen with an indistict white median band, sides mottled gray; a black diagonal mark on each side. Venter with a pair of black spots on the side of a larger white spot. Anterior median eyes one and one-half diameters apart, one diameter from laterals. Posterior median eyes one diameter apart, one and one-half from laterals. Epigynum with a heavily sclerotized groove (Fig. 196). Total length of female, 3.6 mm. Carapace 1.8 mm. long, 1.5 mm. wide. First femur, 3.2 mm.; patella and tibia, 3.6 mm.; metatarsus, 3.0 mm.; tarsus, 1.3 mm. Second patella and tibia, 2.2 mm.; third, 1.5 mm.; fourth, 2.6 mm. ———— LEVI: ACHAEARANEA, THERIDION AND SPHYROTINUS LOT THERIDION VULVUM, new species Figures 197-198 Type. Female holotype from El Valle, Panama, July 1936 (A. M. Chickering), in the Museum of Comparative Zoology. Description. Carapace, sternum, lees yellow. Legs with distal ends of segments brown. Abdomen with a median dorsal white band on dusky background ; sides with several large white spots. Anterior median eyes their diameter apart, two-thirds from laterals. Posterior median eyes one diameter apart, a little more than one from laterals. Total length of female 2.7 mm. Carapace 1.11 mm. long, 0.92 mm. wide. First femur, 1.85 mm.; patella and tibia, 1.96 mm.; metatarsus, 1.46 mm.; tarsus, 0.72 mm. Second patella and tibia, 1.29 mm.; third, 0.87 mm.; fourth, 1.40 mm. Diagnosis. The heavily sclerotized epigynum (Fig. 198) dif- ferentiates this species from other species of the 7. frondeum eroup. Record. Panama: El Valle, 2 paratypes (A. M. Chickering) ; El Volean (W. J. Gertsch). THERIDION HONDURENSE, New species Figures 193-194 Type. Kemale type from Honduras, 1940 (Von Hagen), in the American Museum of Natural History. Description. Carapace, sternum, legs vellow-white. Legs yel- low at distal ends of tibiae one and four. Abdomen with three dorsal white stripes; area in between narrower than stripes; two black spots above spinnerets. Anterior median eves smaller than others, one and one-half diameters apart, one diameter from laterals. Posterior eyes a little more than a diameter apart. Total length, 3.7 mm. Carapace 1.5 mm. long, 1.38 mm. wide. First femur, 3.5 mm.; patella and tibia, 3.5 mm.; metatarsus, 3.6 mm.; tarsus, 0.9 mm. Second patella and tibia, 2.2 mm.; third, 1.8 mm.; fourth, 2.4 mm. Diagnosis. The epigynum having the openings on the anterior surface of a sheht knob (Fig. 194) differentiates this species from others. Record. Honduras: @ paratype (Von Hagen). 108 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY THERIDION BOLIVARI, hew species Figures 189-190 Type. Female type from Desierto de los Leones, Distrito Fed- eral, Mexico, Aug. 5, 1946 (C. Goodnight, Bolivar and Bonet), in the American Museum of Natural History. Description. Carapace golden yellow with a black band as wide as posterior median eyes in front, narrower behind, border with a narrow black line and two black lines across elypeus. Sternum golden yellow. Legs yellow with black bands or lines on venter at middle and distal ends of segments. Dorsum and sides of abdomen covered by small white pigment spots; two lines of black patches on dorsum converging towards spinnerets. Venter with a black semicircular band anterior to pedicel, a black band across epigynum and a black spot behind genital furrow. Anterior median eyes slightly larger than others, three- quarters diameters apart, one-quarter from laterals. Posterior medians three-quarters diameter apart, one diameter from lat- erals. Total leneth of female 3.7 mm. Carapace 1.5 mm. long, 1.4 mm. wide. First femur, 2.2 mm.; patella and tibia, 2.5 mm.; metatarsus, 1.8 mm.; tarsus, 0.8 mm. Second patella and tibia, 1.8 mm. ; third; 1.3)mm.; fourth, 1-9 2mm: Diagnosis. The structure of the epigynum (Fig. 190) dif- ferentiates this species from other Theridion. THERIDION CHIHUAHUA, new species Figures 191-192 Type. Female type from Santa Barbara, Chihuahua, July 18, 1947 (W. J. Gertsch), in the American Museum of Natural History. Description. Carapace, sternum, legs yellow-white. Carapace with two fine parallel median lines. Abdomen white. Anterior median eyes their diameter apart, two-thirds diameter from lat- erals. Posterior median eyes one diameter apart, a little more than one from laterals. Total length of female 3.0 mm. Carapace 1.3 mm. long, 1.1 mm. wide. Second patella and tibia, 1.5 mm.; third, 1.1 mm.; fourth, 1.8 mm. Diagnosis. The epigynum of this species (Fig. 192) resembles that of T. arizonense, but it differs in having two dark posterior areas. The internal genitalia (Fig. 191) are very different and indicate that this species belongs to the 7. frondewm group. LEVI: ACHAEARANEA, THERIDION AND SPHYROTINUS 109 THERIDION APULCO, new species Figures 187-188 Type. Female type from Apuleo, Hidalgo, Oct. 6, 1947 (TH. Wagner), in the American Museum of Natural History. Description. Carapace whitish with a broad median black band. Clypeus dusky. Sternum, legs whitish with dark rings on distal ends of tibiae. Dorsum of abdomen with an irregular white band on each side; a white stripe above spinnerets which continues mottled and darker anteriorly. Black in between me- dian and lateral stripes. Sides and venter yellowish with some indistinct dusky markings. Anterior median eyes slightly smaller than others one diameter apart, one-third of a diameter from laterals. Posterior median eyes three-quarters diameter apart, one diameter from laterals. Total length of female 2.2 mm. Carapace 1.04 mm. long; 0.89 mm. wide. First femur, 1.96 mm.; patella and tibia, 2.09 mm.; metatarsus, 1.62 mm.; tarsus, 0.75 mm. Second patella and tibia, 1.27 mm.; third, 0.70 mm. ; fourth, 1.88 mm. Diagnosis. This species, which has a median depression on the epigynum (Fig. 188), differs from 7. everum by its smaller size, coloration of the venter and internal genitalia (Fie. 187). Record. Veracruz: Wuatuseco (H. M. Wegener). THERIDION TURRIALBA, new species Figures 183-184 Type. Female type from Turrialba, Costa Rica, April 10-17, 1944 (F. Schrader), in the American Museum of Natural His- tory. Description. Carapace, sternum, legs yellow-white. Sternum with a dusky median stripe. Legs with black rings on distal ends of first tibiae. Abdomen whitish, with a white median dorsal band, which becomes narrower toward spinnerets; three white patches on each side. Eyes small. Anterior median eyes slightly smaller than others one and one-half diameters apart, one from laterals. Posterior eyes shghtly more than their diameter apart. Total length of female 3.0 mm. Carapace 0.97 mm. long, 0.88 mm. wide. First femur, 2.34 mm.; patella and tibia, 2.34 mm.; metatarsus, 2.18 mm.; tarsus, 0.81 mm. Second patella and tibia, 1.830 mm.; third, 0.78 mm.; fourth, 1.40 mm. 110 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY Diagnosis. The epigynum is less sclerotized (Fig. 184) than that of T. metabolum. Record. Costa Rica: Turrialba, May 1944 (F. Schrader). TTHERIDION FRIO, New species Figures 199-200 Type. Female type from two miles west of Rio Frio, 3200 m., Distrito Federal, Mexico, July 24, 1956. (W. J. Gertsch and V. Roth), in the American Museum of Natural History. Description. Carapace, sternum, legs whitish. Abdomen white except for two black stripes above and radiating from spinnerets and a pair of black spots on venter. Anterior median eyes one and one-quarter diameters apart, one diameter from laterals. Posterior median eyes one diameter apart, one and one-half from laterals. Total length of female 4.0 mm. Carapace, 1.4 mm. long, 1.3 mm. wide. First femur, 2.6 mm.; patella and tibia, 2.7 mm.; metatarsus, 2.4 mm.; tarsus, 0.8 mm. Second patella and tibia, 1.5 mm.; third 1.1 mm.; fourth, 1.9 mm. This female was collected with a male of T. rothi. Diagnosis. The heavily sclerotized epigynum (Fig. 200) dif- ferentiates this species from others of the 7. frondewm group. THERIDION MORULUM O. P. Cambridge Figures 159-161 Theridion morulum O. P. Cambridge, 1898, Biologia Centrali Americana, Araneidea, vol. 1, p. 254, pl. 35, fig. 6, 9. (2 types from Omilteme, Guerrero, in the British Museum, Natural History.) Roewer, 1942, Katalog der Araneae, vol. 1, p. 495. Steatoda morula, F. P. Cambridge, 1902, Biologia Centrali Americana, Araneidea, vol. 2, p. 384, pl. 36, fig. 14, 9. Theridion jeanae Gertsch and Archer, 1942, Amer, Mus. Novitates, no. 1171, p. 7, figs. 25, 26. (¢ holotype from Santa Catalina Mts., Ari- zona, in the American Museum of Natural History.) Levi, 1957, p. 79. NEW SYNONYMY. Description. Carapace yellow. Sternum, legs, dusky yellow. Abdomen black with some dorsal median light stripes and two pairs of diagonal lines. Venter dark gray. Anterior portion of sides very light. Anterior median eyes one diameter apart, one- quarter diameter from laterals. Posterior median eyes less than LEVI: ACHAEARANEA, THERIDION AND SPHYROTINUS HO one diameter apart, one diameter from laterals. Clypeus pro- jecting below eyes. Abdomen nearly spherical, as wide as long, height less than length. Epigynum (Figs. 160, 161) cone shaped, cone black, the tip slightly lghter. Measurements of female, total length, 3.8 mm. Carapace, 1.30 mm. long, 1.17 mm. wide. First femur, 2.02 mm.; patella and tibia, 2.28 mm.; meta- tarsus, 1.56 mm.; tarsus, 0.78 mm. Second patella and tibia, 1.56 mm. Theridion morulum is much darker than jeanae, and the me- dian apophysis is of slightly different shape. Although this is undoubtedly one species, jeanae might well be used as a sub- species. Records. Morelos: Cuernavaca (W. J. Gertsch, V. Roth) ; Oaxtepee (C. Bolivar). Puebla: Tlacotepee (W. J. Gertsch, V. Roth). Miscellaneous species of Theridion The species related to 7. positivum (istokpoga, cazieri, and sinaloa) are probably close to 7. frondeum; however, the abdo- men tends to be wider than long as in many other species put in this group. Miscellaneous species found in this region and also occurring north of Mexico, illustrated in a previous paper (1957, Bull. Amer. Mus. Nat. Hist., vol. 112) are: THERIDION RUFIPES Lueas Theridion rufipes, Levi, 1957, p. 56, figs. 188-198, 9, 6, Map 18. Distribution. Cosmotropical, but uncommon wherever found in the region under consideration. Additional records. Panama: El Valle; Boquete (A. M. Chickering). Cuba: Soledad, Cienfuegos (Parsons; Salt and Myers) ; Vilcher’s Cave, Soledad (L. G. Worley). Puerto Rico: Aguadilla (R. Turner). Venezwela: Caracas (A. M. Nadler) ; Rancho Grande (W. Beebe). THERIDION ADAMSONI Berland Theridion adamsoni Berland, 1935, Bull. B. P. Bishop Museum, no. 113, p. 102, figs. 6-9, 9 (@ syntypes from Hitiaa, Tahiti, in the B. P. Bishop Museum, Honolulu). WIP BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY Theridion hobbsi Gertsch and Archer, 1942, Amer. Mus. Novitates, no. 1171, p. 95, fig. 6, 2 (2 type from Florida, in the American Museum of Natural History). Levi, 1957, p. 62, figs. 198, 199, 209, 213, 214, 9, 6, map 28. NEW SYNONYMY. Theridion insulicola Bryant, 1947, Psyche, vol. 54, p. 88, fig. 1, 9. (9 type from Mona Island, West Indies, in the Museum of Comparative Zool- ogy). NEW SYNONYMY. Dr. Gertsch first noticed this synonymy. The type of 7’. adam- son’ Was examined. Distribution. Cosmotropieal. Additional records. Canal Zone: Summit (A. M. Chicker- ing). Venezuela: Maracay (A. M. Nadler). THERIDION ATROPUNCTATUM Petrunkevitch Wamba congener O. P. Cambridge, 1896, Biologia Centrali Americana, Araneidea, vol. 1, p. 191, pl. 24, fig. 1, 6 (¢@ type from Chicoyoito, Guatemala, in the British Museum, Natural History). Name preoceu- pied by Theridion congener O. P. Cambridge, 18638. NEW SYN- ONYMY. Theridion atropunctatum Petrunkevitch, 1930, Trans. Connecticut Acad. Sci., vol. 30, p. 210, figs. 59, 60, 2 (2 type from Arecibo, Puerto Rico, in the American Museum of Natural History). Levi, 1957, p. 66, figs. 225-228, 232-234, 9, ¢, map 21. Crustulina congener, Roewer, 1942, Katalog der Araneae, vol. 1, p. 399. NEW SYNONYMY. Theridion brevipalpus Bryant, 1942, Bull. Mus. Comp. Zool., vol. 89, p. 341, figs. 28, 31, 32, 6 (6 type from St. Croix, Virgin Islands, in the Museum of Comparative Zoology). The leg length of some males seems to be shorter than others ; this might be correlated with heavier sclerotization. Males from Central America have a spine on the cymbium as in 7. crispulum but the spine is smaller and on the side of the ceymbium. Females have the epigynum with a more or less circular depression hay- ing a dark spot on each side. The very short connecting ducts separate it from 7’. crispulum. Distribution. Florida, Yucatan, Bimini, Puerto Rico, Virgin Isl and Central America. Additional records. Yucatan: Chichén Itza (C. Goodnight). Panama: Arraijan (A. M. Chickering). Canal Zone: Summit; Barro Colorado Isl. (several records) ; Experimental Gardens ; Balboa; Forest Reserve (A. M. Chickering). Jamaica: Discoy- LEVI: ACHAEARANEA, THERIDION AND SPHYROTINUS 113 ery Bay; St. Ann’s Bay; Holland Bay; 8S. W. of Unity Valley ; St. Catharine; Kinloss; Lyssons (all A. M. Nadler). Hardwar Gap, Blue Mts.; Kingston; St. Thomas Par. Dominican Rep.: Loma Vieja, Cord. Central (P. J. Darlington) ; La Matica, Boca Chica (A. M. Nadler); Ciudad Trujillo (A. M. Nadler). Puerto Rico: Cidra, Treasure I[sl.; Port Grila, San German ; Mayaguez (all A. M. Nadler). Virgin Isl.: St. Croix (Beatty ; A.M. Nadler). Lesser Antilles: Antigua; St. John. Guadeloupe : Pointe-a-Pitre. Martinique: Fond la Hage. Trinidad: Piarco (all A. M. Nadler). Venezuela: Maracay; Caracas (A. M. Nadler). THERIDION CRISPULUM Simon Theridion crispulum Simon, 1895, Ann. Ent. Soe. France, vol. 64, p. 142 (2 type from La Guiara, Venezuela, in the Museum National d’Histoire Naturelle, Paris). Theridion intervallatun Emerton, 1915, Trans. Connecticut Acad. Sci., vol. 20, p. 136, pl. 1, fig. 1, ¢ (4 type from Intervale, New Hampshire, in the Museum of Comparative Zoology). Levi, 1957, p. 64, figs. 222-224, 229-231, 9, 6, map 21. NEW SYNONYMY. The tvpe specimens of 7. crispuluwm examined were marked as coming from San Esteban, Venezuela. The spine on the evmbium of the male palp is at or close to the tip; frequently the eymbium is drawn out to the spine. The connecting duets are much longer than those of 7. atropuncta- tunv; sometimes they open close together, at other times farther apart. There also seem to be differences in the thickness of the duets. Distribution. New Hampshire, southeastern United States, eastern Mexico to Central America, Lesser Antilles. Additional records. Panama: El Volean (A. M. Chickering). Canal Zone: Barro Colorado Isl.; Forest Reserve; Summit; La Boca; Ft. Sherman (A. M. Chickering). Jamaica: Hope Gar- dens; Hermitage Reservoir; Mt. James, St. Andrew Par. (A. M. Chickering). Lesser Antilles: Aruba: Oranjestad, ¢ (A. M. Nadler). Trinidad: nr. Port of Spain (R. Thaxter). Vene- zuela: San Esteban. 114 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY THERIDION ISTOKPOGA Levi Theridion istokpoga Levi, 1957, p. 67, figs. 235, 236, 247, 248, 2, ¢, map 22. The only female found in Central America has the posterior rim of the epigynum wider than Florida specimens; the openings are thus much closer to the posterior end of the depression. The legs are white with black spots on the venter. Distribution. Florida, Chiapas and Canal Zone. Additional record. Canal Zone: Summit, Aug. 1950, ¢ (A. M. Chickering). THERIDION PostTiIvUM Chamberlin Theridion positivum Chamberlin, 1924, Proce. California Acad. Sci., ser. 4, vol. 12, p. 686 (2 type from Pond Island, Gulf of California, in the California Academy of Science). Levi, 1957, p. 68, figs. 237-239, 243- 246, 29, 6, map 22. Theridion cybele Bryant, 1942, Bull. Mus. Comp. Zool., vol. 89, p. 342, fig. 39, 2 (2 type from St. Croix, Virgin Islands, in the Museum of Com- parative Zoology). Specimens examined from Central America have the embolic thread more curved and the posterior border of the depression of the epigynum more straight, than specimens from the north. Distribution. Southern California, southern Texas, Mexico, Central America and Bimini Isl. in the Greater Antilles. Additional records. Canal Zone: “Experimental Gardens; Summit; Chilibre (all A. M. Chickering). Cuba: San Vicente, Pinar del Rio (C. and P. Vaurie). Jamaica: Kingston; Lys- sons; Negrill; Holland Bay (all A. M. Nadler); St. Thomas Par.; St. James Par.; St. Andrew Par. (A. M. Chickering). Puerto Rico: Mayaguez; La Matica, Boca Chica (all A. M. Nad- ler). Lesser Antilles: Antigua: St. John (A. M. Nadler). Venezuela: Maracay (A. M. Nadler). THERIDION ALBULUM QO. P. Cambridge. Figures 213-215 Theridion albulum O. P. Cambridge, 1898, Biologia Centrali Americana, Araneidea, vol. 1, p. 255, pl. 34, fig. 9, 2 (2 type from Bugaba, Panama, lost). Roewer, 1952, Katalog der Araneae, vol. 1, p. 489. In general appearance this species 1s very much hke the two preceding and can easily be confused. It differs, however, in the LEVI: ACHAEARANEA, THERIDION AND SPHYROTINUS A) shape of the base of the palpal embolus (Fig. 215) and in having two dark sclerotized plates in the depression of the epigynum (Fig. 214). The depression of the epigynum is usually filled with a white substance, which has to be removed before exami- nation. Records. Canal Zone: Summit; Forest Reserve; Boquete; Ar- raijan (A. M. Chickering). THERIDION CAZIERI, new species. Figures 216-218 Type. Male type from South Bimini, Bahama Islands, June 1951 (M. A. Cazier), in the American Museum of Natural His- tory. Description. Carapace yellow-white with an irregular dusky median band which is widest in front and has two large yellow white spots behind eyes. Eye region, clypeus, median portions of chelicerae black. Sternum, legs yellow-white. Legs with many brown and black spots. Abdomen white with black spots; a median dorsal white band; several black streaks above spin- nerets. Anterior median eyes their diameter apart, almost touching laterals. Posterior median eyes their radius apart, two- thirds diameters from laterals. Total length of female 3.1 mm. Carapace 1.1 mm. long, 1.0 mm. wide. First femur, 2.2 mm.; patella and tibia, 2.6 mm.; metatarsus, 2.0 mm.; tarsus, 0.8 mm. Second patella and tibia, 1.6 mm.; third, 0.9 mm.; fourth, 1.6 mm. Total length of male 2.7 mm. Carapace 1.3 mm. long, 1.1 mm. wide. First femur, 3.0 mm.; patella and tibia, 3.5 mm.; metatarsus, 3.0 mm.; tarsus, 0.9 mm. Second patella and tibia, 2.38 mm.; third, 1.2 mm.; fourth, 1.8 mm. Diagnosis. The genitaha (Figs. 217, 218) differentiate this species from 7. positivum Chamberlin. Record. Bahama Isl.: South Bimini, @, May 1951 (W. J. Gertsch, M. A. Cazier). THERIDION SINALOA, new species Figure 212 Type. Male type from north section of Mazatlan, Sinaloa, 1956 (A. F. Archer), in the American Museum of Natural His- tory. 116 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY Description. Carapace yellow-white. Sternum white with a dusky border. Legs white with some black spots. Abdomen light covered with white pigment and some dusky spots and two dusky lines converging towards spinnerets on dorsum. Anterior median eyes slightly larger than others, two-thirds diameter apart, touching laterals. Posterior eyes two-thirds diameter apart. Total length of male 1.2 mm. Carapace 0.55 mm. long, 0.50 mm. wide. First femur, 0.91 mm.; patella and tibia, 1.02 mm.; meta- tarsus, 0.70 mm.; tarsus, 0.39 mm. Second patella and tibia, 0.69 mm.; third, 0.44 mm.; fourth, 0.62 mm. Diagnosis. The shape of the embolus (Fig. 212) separates this species from 7. positivum. Record. Sinaloa: Mazatlan, 6 paratype (A. F. Archer). THERIDION ANTILLANUM Simon Figures 223-226; Map 4 Theridion antillanwn Simon, 1894, Proe. Zool. Soe. London, p. 522, fig. 2, 6 (2, 6 syntypes from St. Vincent Isl., Lesser Antilles, in the British Museum, Natural History). Spintharus minutus Petrunkevitch, 1926, Trans. Connecticut Acad. Sci., vol. 28, p. 51, fig. 11, 9 (2 type from St. Thomas, Virgin Isl.). NEW SYNONYMY. Theridion dexteri Petrunkevitch, 1930, ibid., vol. 30, p. 200, figs. 45-50, 9, 6 (6 9,1 4 syntype from University Campus, Rio Piedras, Porto Rico). NEW SYNONYMY. Comments. The embolus of the palpus is on the mesal side (Fig. 225). The radix seems absent; it has probably been lost secondarily as a result of the shifting of other sclerites. The ventral ducts of the epigynum (Figs. 223, 224) which form a rough rectangle or oval are sometimes wider apart, sometimes closer together; the seminal receptacles may be visible through the epigynum at times. The dark areas (coil of the duct), on each side in the epigynum, readily distinguish this species from other small Theridion lacking pigment. Specimens determined by Simon were examined. Natural History. It has been collected from leaves of cocoanut palms (Petrunkevitch, 1930). Records. Bahama Isl.: South Bimini (A. M. Nadler, W. J. Gertsch; M. A. Cazier) ; New Providence Isl. (EK. B. Hayden) ; West Caicos Isl. (EH. B. Hayden). Jamaica: St. Thomas Par.:; LEVI: ACHAKEARANEA, THERIDION AND SPHYROTINUS iy Trelawny Par.; Hanover Par.; St. James Par.; St. Andrew Par. (many records) ; Manchester Par. (A. M. Chickering) ; St. Catherine Par. (A. M. Chickering); Holland Bay (A. M. Nad- ler) ; Lyssons (A. M. Nadler) ; Kinloss (A. M. Nadler) ; Negrill (A. M. Nadler) ; Kingston (A. M. Chickering). Dominican Rep.: Roseau: Valle de Polo. Puerto Rico: (Petrunkeviteh, 1930) ; Maricao Forest, 2500 ft. (P. J. Darlington) ; San Juan; Cidra, Treasure Isl. (A. M. Nadler) ; Mayaguez (A. M. Nadler) ; Virgin Isl.: (Petrunkevitch, 1926) ; St. Croix (H. A. Beatty; C. BE. Wil- son; F. BE. Lutz). Lesser Antilles: Guadeloupe: Point-a-Pitre (A. M. Nadler); Martinique: Fond la Haye (A. M. Nadler) ; St. Vineent Isl.: (Simon, 1894). THERIDION JAMAICENSE, new species Figures 219-222 Type. Male holotype from Dolphin Head Trail, Hanover Par- ish, Jamaica, June 24, 1954, in the Museum of Comparative Zoology. Description. Yellow-white without marks. Reported to be erass-green when alive. Eyes small. Anterior medians of female almost two diameters apart, two-thirds from laterals, of male one diameter apart, one-quarter from laterals. Posterior me- dians two diameters apart, one from laterals. Abdomen some- times wider than long. Total length of female 2.0 mm. Carapace 0.85 mm. long; 0.65 mm. wide. First femur, 1.62 mm.; patella and tibia, 1.56 mm.; metatarsus, 1.43 mm.; tarsus, 0.52 mm. Second patella and tibia, 1.06 mm.; third, 0.59 mm. ; fourth, 1.14 mm. Total length of male 1.7 mm. Carapace, 0.91 mm. long, 0.75 mm. wide. First femur, 1.56 mm.; patella and tibia, 1.56 mm.; metatarsus, 1.30 mm.; tarsus, 0.47 mm. Second patella and tibia, 1.00 mm.; third, 0.65 mm.; fourth, 0.94 mm. Diagnosis. The female of this peculiar species has a scape on the epigynum (Figs. 220, 221) which reminds one of linyphiid spiders. The palpus is quite translucent and the various sclerites are lobed. The dark portion on the right edge of the palpus (Fig. 222) is the embolus; on the left the sclerotized portion is the median apophysis. This specices is believed to be related to T. antillanum. The genitalia distinguish the two speccies. Natural History. This species is found on vegetation. 118 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY Records. Jamaica: Dolphin Head Trail, Hanover Par., 9 paratype; Dolphin Head, 1600 ft. (G. R. Procter) ; Hermitage, St. Andrew Par. (Bengra) ; Grier Mtn., 2300 ft., St. Catherine Par. (G. R. Procter) ; St. Ann Par. (A. M. Nadler). THERIDION NUDUM, new species Figures 227-229 Type. Male type from El Volean, Panama, Aug. 9-14, 1950 (A. M. Chickering), in the Museum of Comparative Zoology. Description. Yellow-white, lacking pigment except for a black spot on thoracic depression. Anterior median eyes slightly smaller than others, two diameters apart, three-quarters from laterals. Posterior median eyes one and three-quarters diameters apart, one from laterals. Total length of females 1.5-2.2 mm. A female measured total length 2.1 mm. Carapace 0.79 mm. long, 0.71 mm. wide. First femur, 1.36 mm.; patella and tibia, 1.36 mm.; metatarsus, 1.06 mm.; tarsus, 0.56 mm. Second patella and tibia, 0.91 mm.; third, 0.56 mm.; fourth 0.88 mm. A male measured total length 1.7 mm. Carapace 0.90 mm. long, 0.72 mm. wide. First femur, 1.20 mm.; patella and tibia, 1.24 mm.; metatarsus, 0.94 mm.; tarsus, 0.45 mm. Second patella and tibia, 0.88 mm.; third, 0.52 mm.; fourth, 0.80 mm. One male from Chiapas has seattered red spots on the abdo- men. Diagnosis. This species has the abdomen wider than long; it can be separated from 7. sexmaculatum and T. rufipunctum by the genitalia (Figs. 227-229). Records. Hidalgo: Chapulhuacan (W. J. Gertsch). Puebla: Villa Juarez (H. Wagner). Chiapas: Tenejapa (C. Goodnight) : nr. Cacahuatan, 38000 m. (C. Goodnight). Panama: El Vol- ean, 2 allotype, 2, ¢ paratypes; El Valle; Boquete (all A. M. Chickering). THERIDION RUFIPUNCTUM, new species Figures 230-233 Type. Male type from El Voleén, Panama, Aug. 9-14, 1950 (A. M. Chickering), in the Museum of Comparative Zoology. Description. Carapace with one or two small median black spots. Sternum white. Legs white with a small black spot pro- LEVI: ACHAEARANEA, THERIDION AND SPHYROTINUS 119 lateral on patellae and on distal ends of tibiae. Abdomen white, dorsum and sides with red spots (Fig. 230). Anterior median eyes one and one-half diameters apart, one from laterals. Pos- terior median eyes one and one-half diameters apart, one from laterals in female; one and one-half in male. Total length of females 1.8-2.2 mm. One female measured total length 2.0 mm. ; carapace 0.91 mm. long, 0.71 mm. wide. First femur, 1.43 mm.; patella and tibia, 1.30 mm.; metatarsus, 1.01 mm.; tarsus, 0.74 mm. Second patella and tibia, 1.01 mm.; third, 0.59 mm. ; fourth, 0.92 mm. Male measured total length, 2.0 mm. Carapace 0.91 mm. long, 0.75 mm. wide. First femur, 1.20 mm.; patella and tibia, 1.47 mm.; metatarsus, 1.00 mm.; tarsus, 0.50 mm. Second patella and tibia, 1.03 mm.; third, 0.65 mm.; fourth, 0.92 mm. Diagnosis. The spotted abdomen (Fig. 230), which is wider than long in the female, and the genitalia (Figs. 231, 233) dis- tinguish this species from 7. sexrmaculatum. Records. Panama: El Voleén, 2 allotype, paratypes (A. M. Chickering; W. J. Gertsch) ; Boquete (A. M. Chickering). THERIDION NADLERI, new species Figures 240-241 Type. Female type from Simla, near Arima, Trinidad, Dee. 28, 1954 (A. M. Nadler), in the American Museum of Natural History. Description. Spider yellow-white in coloration except dorsum of abdomen which has some pigment spots. Diameter of an- terior median eyes one and one-half times that of other eyes. Anterior median eyes their diameter apart, almost touching lat- erals. Posterior median eyes one and one-half diameters apart, two-thirds from laterals. Abdomen wider than long. Total length of female 1.9 mm. Carapace 0.75 mm. long, 0.65 mm. wide. First femur, 1.20 mm.; patella and tibia, 1.20 mm.; metatarsus, 0.81 mm.; tarsus, 0.43 mm. Second patella and tibia, 0.91 mm. ; third, 0.65 mm.; fourth, 0.83 mm. Diagnosis. The large opening of the epigynum (Fig. 241) dif- ferentiates this species from others. THERIDION SEXMACULATUM Keyserling Figures 256-239 Theridion sermaculatum Keyserling, 1884, Die Spinnen Amerikas, vol. 1, no. 1, p. 82, pl. 4, fig. 51, 2 (@ syntypes from Amazonas Province, 120 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY Brazil, in the Hope Department of Entomology, Oxford University) ; 1886, ibid., vol. 2, p. 238, pl. 20, fig. 292, Steatoda sermaculata, F. P. Cambridge, 1902, Biologia Centrali Americana, Araneidea, vol. 2, p. 386, pl. 36, fig. 22, Q. Diagnosis. Color yellow-white except for three pairs of black spots on abdomen (Fig. 238); the spots which are only rarely absent and the genitalia (Figs. 237, 239) distinguish this small species. Records. Guatemala. Canal Zone: Barro Colorado Isl.; Sum- mit; Experimental Gardens; Forest Reserve (all A. M. Chieker- ing); El Volean (W. J. Gertseh). Venezuela: Rancho Grande (A. M. Nadler). THERIDION DOTANUM (Banks), new combination Figures 245-248 Singa dotana Banks, 19138, Proce. Acad. Nat. Sci. Philadelphia, vol. 65, p. 679, pl. 28, fig. 1 (2 type from St. Maria Dota, Costa Riea, in the Museum of Comparative Zoology). Description. Carapace, sternum, legs yellow-white. Legs with small black spots on distal ends of segments. Abdomen wider than long (Fig. 247) covered by white pigment and a black spot on each side; that of male narrower and darker. Eyes without black rings. Anterior median eyes more than a diameter apart, one-third of a diameter from laterals. Posterior eyes a third of a diameter apart. Anterior median eves shghtly smaller than others. Total length of female 2.5 mm. Carapace 0.94 mm. long, 1.11 mm. wide. First femur, 1.89 mm.; patella and tibia, 1.92 mm.; metatarsus, 140 mm.; tarsus, 0.62 mm. Second patella and tibia, 1.56 mm.; third, 1.04 mm.; fourth, 1.30 mm. Total leneth of male 2.0 mm. Carapace 0.98 mm. long, 0.92 mm. wide. Kirst femur, 2.14 mm.; patella and tibia, 2.14 mm.; metatarsus, 1.50 mm.; tarsus, 0.57 mm. Second patella and tibia, 1.70 mm. ; third, 0.92 mm.; fourth, 1.17 mm. Diagnosis. The shape of the abdomen, the epigynum having a large bordered depression (Fig. 246) and the palpus (Fig. 248) distinguish this species from 7’. sermaculatum. Record. Oaxaca: Oaxaca (B. Malkin). Panama: El Volean (A. M. Chickering, W. J. Gertsch). LEVI: ACHAEARANEA, THERIDION AND SPHYROTINUS 121 THERIDION PALLISTERORUM, new species Figures 242-244 Type. Male type from Contreras, Distrito Federal, 8-9000 ft., May 30, 1946 (C. J. and D. L. Pallister), in the American Mu- seum of Natural History. Description. Carapace yellow with a narrow black median line and black border. Legs yellow, femora brown to black, other segments with some black spots underneath. Abdomen white, sprinkled with black on dorsum, a white spot on the widest point on each side; venter with a median black spot. Anterior median eyes almost their diameter apart, one-third from laterals. Pos- terior median eyes almost their diameter apart, slightly more than a diameter from laterals. Total length of male 2.4 mm. Carapace, 1.1 mm. long, 1.1 mm. wide. First femur, 2.9 mm.; patella and tibia, 2.7 mm.; metatarsus, 1.8 mm.; tarsus, 0.7 mm. Second patella and tibia, 2.2 mm.; third, 1.0 mm.; fourth, 1.3 mm. A female, which was slightly larger than the male, was in too poor condition for measurements. Diagnosis. Only details of the palpus (Fig. 244) separate this species from 7. dotanum. It is possible that this belongs to a northern form of the latter species. Record. Distrito Federal: Santa Rosa, 2700 m., July 13, 1946, 2 (H. Wagner). THERIDION MINUTISSIMUM Keyserling Figure 249 Theridion minutissimum Keyserling, 1884, Die Spinnen Amerikas, vol. 2, no. 1, p. 34, fig. 17, 6, 2, (46 Lectotype here designated from Montana di Nancho, Peru, in the Polish Academy of Sciences, Warsaw.) Description. Carapace, sternum orange. Most leg segments orange-white; first and second tibiae dark gray except for ends. All metatarsi dark gray. Abdomen black except for a broad horseshoe-shaped white band which encircles sides and anterior of dorsum, a white spot above spinnerets and four small white spots in center of dorsum. Anterior median eyes slightly larger than others, three-quarters diameter apart, almost touching lat- erals. Posterior eves one-third diameter apart. Total length of 22, BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY male 1.2 mm. Carapace 0.71 mm. long, 0.52 mm. wide. First femur, 0.91 mm.; patella and tibia, 0.96 mm.; metatarsus, 0.60 mm.; tarsus, 0.28 mm. Second patella and tibia, 0.74 mm.; third, 0.45 mm.; fourth, 0.57 mm. The type of this species was exam- ined. Record. Panama Canal Zone: Experimental Gardens, Aug. 12-19, 1954 (A. M. Chickering). THERIDION BARROANUM, new species Figures 234-235 Type. Female type from Barro Colorado Island, June 1950 (A. M. Chickering), in the Museum of Comparative Zoology. Description. Carapace, sternum, legs, abdomen vellow-white. Abdomen wider than lone covered by white pigment on sides of dorsum and sides, but not in center of dorsum. Anterior median eyes one diameter apart, one-fourth from laterals. Posterior median eyes their radius apart, their diameter from laterals. Total length, 1.9 mm. Carapace 0.89 mm. long, 0.82 mm. wide. Kirst femur, 1.10 mm.; patella and tibia, 1.10 mm.; metatarsus, 0.78 mm.; tarsus, 0.45 mm. Second patella and tibia, 0.98 mm. ; third, 0.70 mm.; fourth, 0.85 mm. Diagnosis. This species can be distinguished from other spe- cies, with an abdomen wider than long, by the position of the connecting ducts (Figs. 234, 235). THERIDION CHIRIQUI, new species Figures 268-271 Type. Male type from El] Voleén, Panama, Feb. 26, 1936 (W. J. Gertsech). in the American Museum of Natural History. Description. Yellow-white except for white dorsum of abdo- men which is wider than long. Venter of abdomen anterior to pedicel dusky. In male. white pigment band across abdomen (Fig. 269). Anterior median eyes their diameter apart, less than one-fourth from laterals. Posterior median eyes two-thirds diameter apart, less than one-fourth in female, one radius in male from laterals. Total length of female 1.3 mm. Carapace, 0.65 mm. long, 0.55 mm. wide. First femur, 1.10 mm.; patella and tibia, 1.04 mm.; metatarsus, 0.80 mm.; tarsus, 0.39 mm. LEVI: ACHAEARANEA, THERIDION AND SPHYROTINUS 123 Second patella and tibia, 0.78 mm.; third, 0.52 mm.; fourth, 0.79 mm. Total length of male 1.2 mm. Carapace, 0.61 mm. long, 0.55 mm. wide. First femur, 1.10 mm.; patella and tibia, 1.24 mm.; metatarsus, 0.83 mm.; tarsus, 0.42 mm. Second patella and tibia, 0.78 mm.; third, 0.50 mm.; fourth, 0.72 mm. Diagnosis. Only the structure of the genitalia (Figs. 268, 271) distinguishes this species from 7. nudum and others whose abdo- men is wider than long. The posterior median eyes of the female almost touch the laterals. Record. Panama: El Volean, 2, 6 paratypes (W. J. Gertsch ). THERIDION PANAMENSE, new species Figures 263-267 Type. Male type from El Volean, Aug. 9-14, 1950 (A. M. Chickering), in the Museum of Comparative Zoology. Description. Carapace dark dusky yellow with a dark border. Sternum dark dusky. Legs vellow with irregular black patches and stripes. Dorsum of abdomen with a median dorsal white stripe, on sides black: sides of abdomen whitish, venter as illustrated (Fig. 267). Anterior median eyes one diameter apart, one-fourth diameter from laterals. Posterior eyes one diameter apart. Anterior median eyes sometimes slightly larger than others. Abdomen usually wider than lone in female, not in male. Total length of females 1.6-2.5 mm. Female allotype measured total length 1.8 mm. Carapace 0.81 mm. long, 0.78 mm. wide. First femur, 1.36 mm.; patella and tibia, 1.51 mm.; metatarsus, 1.14 mm.; tarsus, 0.52 mm. Second patella and tibia, 1.05 mm.; third, 0.64 mm.; fourth, 0.98 mm. Total length of males 1.6-1.8 mm. Male type measured total length 1.6 mm. Carapace 0.88 mm. long, 0.78 mm. wide. First femur, 1.58 mm. ; patella and tibia, 1.75 mm.; metatarsus, 1.48 mm.; tarsus, 0.52 mm. Second patella and tibia, 1.25 mm.; third, 0.71 mm.; fourth, 0.95 mm. Diagnosis. The dark coloration, wider abdomen and epigynum with an anterior lip (Fig. 264), and the palpus, of the male (Figs. 265, 266) distinguish this from Theridion atropunctatum. Records. Panama: Boquete (many specimens); El Valle; 2 allotype, ¢ paratypes (all A. M. Chickering) ; El Volean (A. M. Chickering; W. J. Gertsch). 124 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY THERIDION ARTUM, new species Figures 313-314 Type. Female type from Forest Preserve, Panama Canal Zone, Jan. 1958 (A. M. Chickering), in the Museum of Com- parative Zoology. Description. Carapace dusky black, lighter in center, reddish around eyes. Sternum black. Legs yellowish with black rings on distal ends of femora, tibiae, metatarsi and irregular black spots on venter of segments. Abdomen with a median dorsal gray band, bordered by white, a white area anterior and posterior of the band. Venter black with tongues of black extending to the sides and a pair up the posterior. A pair of white spots between genital groove and spinnerets. Anterior median eyes shghtly larger than others, their diameter apart and almost touching laterals. Posterior eyes their diameter apart. Total length 2.2 mm. Carapace 0.98 mm. long, 0.85 mm. wide. First femur, 1.92 mm.; patella and tibia, 1.96 mm.; metatarsus, 1.50 mm.; tarsus, 0.63 mm. Second patella and tibia, 1.22 mm.; third, 0.73 mm.; fourth, 1.23 mm. Diagnosis. The epigynum (Fig. 314) which is a ight area in a heavily pigmented region separates this species from Theridion akron. The openings are difficult to discern. THERIDION AKRON, new species Figures 250-252 Type. Male type from Summit, Panama Canal Zone, Aug. 1950 (A. M. Chickering), in the Museum of Comparative Zool- ogy. Description. Carapace dusky yellow, sternum dusky. Legs yellowish; ends of segments dusky, with dusky patches on ven- ter. Dorsum of abdomen with a median black band on spotted white background; continuation of band white in color anterior and posterior; sides of anterior and posterior portion of dorsum darker. Venter black with a pair of white spots. Anterior me- dian eyes slightly larger than others, one diameter apart, almost touching laterals. Posterior median eyes their radius apart, three-quarters their diameter from laterals. Total length of fe- male 2.0 mm. Carapace 0.78 mm. long, 0.66 mm. wide. First femur, 1.50 mm.; patella and tibia, 1.60 mm.; metatarsus, 1.17 LEVI: ACHAEARANEA, THERIDION AND SPHYROTINUS 125 mim. ; tarsus, 0.49 mm. Second patella and tibia, 0.95 mm. ; third, 0.92 mm. Total length of male 1.7 mm. Carapace 0.84 mm. long, 0.71 mm. wide. First femur, 1.67 mm.; patella and tibia, 1.68 mm.; metatarsus, 1.32 mm.; tarsus, 0.53 mm. Second patella and tibia, 1.10 mm.; third, 0.66 mm.; fourth, 0.99 mm. Diagnosis. The palpus (Fig. 252) resembles that of Dipoena or Achaearanca; it has, however, a radix hidden behind the con- duetor, and the duct does not traverse the median apophysis as it does in Dipoena. The epigynal depression (Fig. 251) may have been covered with material in the specimens examined. The eenitalia differentiate this species from T. panamense. Records. Canal Zone: Barro Colorado Island, 29 (A. M. Chickering). THERIDION RESUM, new species Figures 253-254 Type. Kemale holotype from Forest Reserve, Panama Canal Zone, Aug. 10-12, 1989 (A. M. Chickering), in the Museum of Comparative Zoology. Description. Carapace, sternum, legs dusky yellow. Legs with faint indications of rings or spots on venter. Dorsum of abdo- men spotted white with a darker median band; venter black with a pair of white spots and a white spot anterior to pedicel, a white line above spinnerets which continues into the dark dorsal band. Anterior median eyes slightly larger than others, their diameter apart, almost touching laterals. Posterior eyes less than a di- ameter apart. Abdomen as wide as long. Total length of female 2.0 mm. Carapace 0.78 mm. long, 0.73 mm. wide. First femur, 1.42 mm.; patella and tibia, 1.45 mm.; metatarsus, 1.08 mm.; tarsus, 0.52 mm. Second patella and tibia, 0.98 mm. ; third, 0.62 mm.; fourth, 0.99 mm. Diagnosis. The epigynum which has a median depression with a posterior sclerotized rim (Fig. 254) distinguishes this species from 7. panamense. The latter species has one ventral white spot; 7. resum has two. THERIDION AKME, new species Figures 210-211 Type. Female type from Summit Park, Panama Canal Zone, Dee. 1957 (A. M. Chickering), in the Museum of Comparative Zoology. 126 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY Description. Carapace, sternum yellowish. Legs golden yel- low. Abdomen yellow-white. Anterior median eyes slightly larger than others, two-thirds their diameter apart, one-fourth their diameter from laterals. Posterior median eyes less than their diameter apart, two-thirds their diameter from laterals. Legs short. Abdomen widest slightly behind middle. Total leneth 2.7 mm. Carapace 0.98 mm. long, 0.94 mm. wide. First femur, 1.43 mm.; patella and tibia, 1.30 mm.; metatarsus, 1.00 mm.; tarsus, 0.55 mm. Second patella and tibia, 0.98 mm.; third, 0.75 mm.; fourth, 1.14 mm. Diagnosis. The epigyvnum (Fig. 211) and internal genitalia distinguish this species. THE THERIDION MOCTEZUMA GROUP THERIDION RESERVUM, hew species Figures 318-321 Type. Male type from Forest Reserve, Canal Zone, July 4-6, 1939 (A. M. Chickering), in the Museum of Comparative Zool- ogy. Description. Carapace, legs yellowish. First and second fe- mora of female with two black lines on venter, and black rings around tibia-metatarsal joint. Abdomen yellow-white with two dorsal white lines and series of black spots (Fig. 320). Anterior median eyes one diameter apart, one-fourth from laterals. Pos- terior medians separated by two-thirds diameter in female, by one diameter in male. Total length of female 1.9 mm. Carapace 0.68 mm. long, 0.65 mm. wide. First femur, 1.03 mm.; patella and tibia, 0.96 mm.; metatarsus, 0.80 mm.; tarsus, 0.36 mm. Second patella and tibia, 0.78 mm.; third, 0.52 mm.; fourth, 0.80 mm. Total length of male, 1.7 mm. Carapace 0.78 mm. long, 0.62 mm. wide. First femur, 1.42 mm.; patella and tibia, 1.55 mm.; metatarsus, 1.36 mm.; tarsus, 0.45 mm. Second patella and tibia, 1.00 mm.; third, 0.62 mm.; fourth, 0.95 mm. Diagnosis. The striking coloration of this species as well as details of the genitalia (Figs. 319, 321) distinguish it from other species of Theridion. Record. Canal Zone: Barro Colorado Isl. (several records). Forest Reserve, 2, ¢ paratypes; Madden Dam (all A. M. Chick- ering). LEVI: ACHAEARANEA, THERIDION AND SPHYROTINUS 127 THERIDION LATHROPI, new species Figures 272-276 Type. Male type from Lathrop Trail, Barro Colorado Island, Canal Zone, July 6, 1954 (A. M. Chickering), in the Museum of Comparative Zoology. Description. Carapace, legs dusky yellow. Sternum dusky. Abdomen black with a median longitudinal white stripe on dor sum which is widest in middle, narrower at ends; white patches on sides and a white spot behind genital groove on venter. Anterior median eyes three-quarters diameter apart, one-quarter from laterals. Posterior median eyes one-third diameter apart, three-quarters from laterals. Total length of females 1.8-2.2 mm. A female measured total length 1.8 mm. Carapace 0.83 mm. long, 0.75 mm. wide. First femur, 1.06 mm. ; patella and tibia, 1.12 mm.; metatarsus, 0.78 mm.; tarsus, 0.41 mm. Second patella and tibia, 0.81 mm.; third 0.61 mm.; fourth, 0.91 mm. Total length of males 1.6-1.8 mm. A male measured total length 1.6 mm. Carapace 0.82 mm. long, 0.73 mm. wide. First femur, 1.17 mm.; patella and tibia, 1.35 mm.; metatarsus, 1.04 mm.; tarsus, 0.45 mm. Second patella and tibia, 1.00 mm. ; third, 0.65 mm.; fourth, 0.92 mm. Diagnosis. The palpus (Figs. 275-276) and the epigynun which has a projecting flat hook (Fig. 274) distinguish this species. Records. Panama: Porto Bello (A. M. Chickering). Canal Zone: Barro Colorado Isl. (common) ; Forest Reserve; Ft. Davis (A. M. Chickering). THERIDION PETRUM, new species Figures 280-284 Type. Male type from Barro Colorado Island, Canal Zone, Panama, Aug. 14, 1954 (A. M. Chickering), in the Museum of Comparative Zoology. Description. Carapace, sternum, legs dusky yellow. Abdomen black with white lines on dorsum (Fig. 283) and a white spot on venter between spinnerets and genital groove. Diameter of an- terior median eyes one and one-half that of other eyes. Anterior median eyes one-half their diameter apart, one-third from lat- 128 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY erals. Posterior medians two-thirds their diameter apart, one- half from laterals. Total length of female 1.2 mm. Carapace 0.62 mm. long, 0.55 mm. wide. First femur, 0.65 mm.; patella and tibia, 0.65 mm.; metatarsus, 0.42 mm.; tarsus, 0.31 mm. Second patella and tibia, 0.53 mm.; third, 0.43 mm.; fourth, 0.55 mm. Total length of male 1.8 mm. Carapace 0.62 mm. long; 0.53 mm. wide. First femur, 0.68 mm.; patella and tibia, 0.73 mm.; metatarsus, 0.50 mm.; tarsus, 0.34 mm. Second patella and tibia, 0.57 mm.; third, 0.44 mm.; fourth, 0.60 mm. Diagnosis. The palpus (Fig. 284) differentiates the male from T. marvum. The epigynum has a knob (Figs. 281, 282) re- sembling that of 7. neomexicanum; however, coloration and leg leneth differentiate the two species. Records. Canal Zone: Barro Colorado Isl., 6, 2 paratypes (sev. collections). Lesser Antilles: Trinidad: Piareo (A. M. Nadler). THERIDION MARVUM, new species Figure 326 Type. Male type from Barro Colorado Island, Panama Canal Zone, July 30, 31, 1936 (A. M. Chickering), in the Museum of Comparative Zoology. Description. Carapace, sternum, legs dusky yellow. Abdomen gray with white lines on dorsum similar to 7. petrum (Fig. 283) ; venter with a white spot anterior to spinnerets. Anterior median eyes two-thirds their diameter apart, one-third from laterals. Posterior medians two-thirds their diameter apart, one-half their diameter from laterals. Total length of male 1.3 mm. Carapace 0.71 mm. long, 0.55 mm. wide. First femur, 0.91 mm.; patella and tibia, 0.99 mm.; metatarsus, 0.67 mm.; tarsus, 0.39 mm. Second patella and tibia, 0.78 mm.; third, 0.53 mm.; fourth, 0.67 mm. Diagnosis. Lighter coloration, slightly larger size, smaller an- terior median eyes and the structure of the palpus (Fig. 326) separate this species from 7. petrwm. Records. Canal Zone: Barro Colorado Island, 4 paratypes (A. M. Chickering) ; Boquete (A. M. Chickering). LEVI: ACHAEARANEA, THERIDION AND SPHYROTINUS 129 THERIDION MOCTEZUMA, new species Figures 285-288 Type. Male type from La Planta de Moctezuma, near Fortin, Veracruz, July 7, 1947 (C. and M. Goodnight), in the American Museum of Natural History. Description. Carapace, sternum, legs, dusky yellow-brown. Abdomen black with a white median dorsal band (Fig. 287), several lateral spots and a white spot on venter behind genital groove. Anterior median eyes one diameter apart, one-third from laterals. Posterior median eyes their radius apart, two- thirds from laterals. Abdomen of male with posterior muscle impressions sclerotized. Total length of female 2.5 mm. Cara- pace 1.04 mm. long, 0.93 mm. wide. First femur, 1.43 mm. ; patella and tibia, 1.60 mm.; metatarsus, 1.20 mm.; tarsus, 0.58 mm. Second patella and tibia, 1.15 mm.; third, 0.91 mm.; fourth, 1.30 mm. Total length of male 2.3 mm. Carapace 1.04 mm. long, 0.88 mm. wide. First femur, 1.50 mm.; patella and tibia, 1.80 mm.; metatarsus, 1.40 mm.; tarsus, 0.63 mm. Second patella and tibia, 1.30 mm.; third, 0.98 mm.; fourth, 1.30 mm. Diagnosis. The epigynum which has a translucent projecting knob (Fig. 286) and the palpus can be used to separate this species from 7’. lathropi. Records. Veracruz: La Buena Ventura, 1909 (A. Petrunke- viteh). THERIDION COBANUM, new species Figures 291-292 Type. Female holotype from Coban, Guatemala, July 1947 (C. and P. Vaurie), in the American Museum of Natural His- tory. Description. Carapace yellow-brown with a median dusky line. Sternum, legs yellow-brown; legs with faint indications of darker rings. Dorsum of abdomen with two rows of large black spots, their diameter apart; almost no white pigment present; venter dusky. Anterior median eyes their diameter apart, one- third their diameter from laterals. Posterior eyes two-thirds diameter apart. Total length of female, 2.5 mm. Carapace, 1.30 mm. lone, 1.04 mm. wide. First femur, 1.80 mm.; patella and 130 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY tibia, 1.93 mm.; metatarsus, 1.46 mm.; tarsus, 0.69 mm. Second patella and tibia, 1.48 mm.; third, 1.06 mm.; fourth, 1.61 mm. Diagnosis, The epigynum (Fig. 292) distinguishes this species from 7. moctezwma. Record. Tabasco: Teapa (C. and M. Goodnight). THERIDION ASPERSUM (I. P. Cambridge) Figures 311-312 Steatoda aspersa F. P. Cambridge, 1902, Biologia Centrali Americana, Araneidea, vol. 2, p. 384, pl. 36, fig. 183, 2 (92 type from Guatemala, in the British Museum, Natural History). Theridion aspersum, Roewer, 1942, Katalog der Araneae, vol. 1, p. 490. THERIDION QUANTUM, new species Figures 293-294, 317 Type. Male type from Summit, Canal Zone, July 21-29, 1950 (A. M. Chickering), in the Museum of Comparative Zoology. Description. Carapace vellowish with a median dusky line and dusky sides. Sternum dusky. Lees yellowish with indistinct dusky rings. Abdomen black with a white dorsal stripe, which is narrower at both ends; some white spots on sides; venter black. Anterior median eyes one diameter apart, a radius from laterals. Posterior median eyes of female less than one diameter apart, two-thirds from laterals. Anterior median eyes of males slightly smaller than others. Posterior medians of male two-thirds diame- ter apart, one from laterals. Total length of female 2.2 mm. Carapace 0.91 mm. long, 0.78 mm. wide. First femur, 1.15 mm.; patella and tibia, 1.15 mm.; metatarsus, 0.91 mm.; tarsus, 0.49 mm. Second patella and tibia, 0.96 mm.; third, 0.71 mm.; fourth, 1.04 mm. Total leneth of male 1.6 mm. Carapace 0.78 mm. long, 0.68 mm. wide. First femur, 1.15 mm.; patella and tibia, 1.25 mm.; metatarsus, 0.98 mm.; tarsus, 0.50 mm. Second patella and tibia, 0.92 mm.; third, 0.62 mm.; fourth, 0.87 mm. Diagnosis. The epigynum has a pair of light areas divided by a small lobe (Fig. 294) ; the palpus has a conductor of distinctive shape (Fig. 317). Records. Costa Rica: Orosi (Piecado) ; San José (KE. Schmidt). Canal Zone: Summit, 2° paratypes; Forest Reserve; Experi- mental Gardens (A. M. Chickering). LEVI: ACHAEARANEA, THERIDION AND SPHYROTINUS 131 THERIDION MALKINI, new species Figures 295-296 Type. Female type from Tamazunchale, San Luis Potosi, June 26, 1947 (B. Malkin), in the American Museum of Natural History. Description. Carapace dusky yellow, duskier on sides. Ster- num dusky yellow. Legs yellow with faint dusky rings and dusky femora. Abdomen with a median dorsal white band whose edges are scalloped and which narrows towards the spinnerets ; sides with some white spots. Anterior median eyes two-thirds diameter apart, one-quarter from laterals. Posterior median eyes their radius apart, two-thirds diameter from laterals. Total length of female 2.0 mm. Carapace 0.97 mm. long, 0.91 mm. wide. First femur, 1.24 mm.; patella and tibia, 1.40 mm.; metatarsus, 1.01 mm.; tarsus, 0.52 mm. Second patella and tibia, 1.04 mm.; third, 0.78 mm.; fourth, 1.17 mm. Diagnosis. The epigynum whose openings are anterior (Fig. 296) distinguishes this species from 7. quantum. THERIDION RUINUM, hew species Figure 325 Type. Male type from Las Ruinas de Palenque, Chiapas, July 14, 1949 (C. J. Goodnight), in the American Museum of Natural History. Description. Carapace yellow with a median dusky stripe and dusky sides. Sides of sternum dusky. Legs dusky yellow. Abdo- men black with a median dorsal white stripe having scalloped edges; venter without much pigment. Eyes large. Anterior medians three-quarters diameter apart, almost touching laterals. Posterior medians one-third diameter apart, one-half from lat- erals. Total length of male, 1.5 mm. Carapace 0.67 mm. long, 0.62 mm. wide. First femur, 1.00 mm.; patella and tibia, 1.04 mm.; metatarsus, 0.87 mm.; tarsus, 0.42 mm. Second patella and tibia, 0.85 mm.; third 0.62 mm.; fourth, 0.85 mm. Diagnosis. Details of the palpus (Fig. 325) separate this spe- cies from T. quantum. 132 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY THERIDION SIGNUM, new species Figures 297-298, 332 Type. Male type from Boquete, Panama, July 10-25, 1939 (A. M. Chickering), in the Museum of Comparative Zoology. Description. Carapace dusky yellow-brown. Sternum dusky vellow. Legs yellow-brown with indistinct dusky bands on middle and ends of segments. Abdomen mottled white on black with a median dorsal white band and a series of five white spots on sides; venter with an indistinct spot behind epigastric furrow. Anterior median eyes one diameter apart less than one-fourth from laterals. Posterior medians their radius apart, less than a diameter from laterals. A female measured total length 2.2 mm. Carapace 1.0 mm. long, 0.89 mm. wide. First femur 1.33 mm. ; patella and tibia, 1.44 mm.; metatarsus, 1.05 mm.; tarsus, 0.55 mm. Second patella and tibia, 1.10 mm.; third, 0.78 mm.; fourth, 1.18 mm. A male measured total length, 1.9 mm. Carapace 0.88 mm. long, 0.78 mm. wide. First femur, 1.28 mm.; patella and tibia, 1.48 mm.; metatarsus, 1.01 mm.; tarsus, 0.51 mm. Second patella and tibia, 1.10 mm.; third, 0.75 mm.; > fourth, 1.01 mm. This species is distinguished by the following characters : Diagnosis. The epigynum has a small hookshaped projection (Fig. 298); the palpus has the radix enlarged, covering the conductor on the ventral side by a transparent lobe, wrapped all around the back of the bulb and terminating in a large flat pro- jection. Embolus is small and hidden by conductor and radix (Fig. 332). Records. Panama: Boquete, 2 allotype July 10-25, 1939; 9 paratype Aug. 4-11, 1954 (A. M. Chickering). THERIDION CHILAPA, hew species Figures 309-310 Type. Female type from Chilapa, Guerrero, Mexico, Oct. 29, 1934 (L. Schultze), in the American Museum of Natural History. Description. Carapace yellow-brown. Legs yellow-brown with some dusky rines on middle and ends of distal segments. Abdo- men black with a median dorsal black band bordered by a wavy white line. Some light patches on sides. Eyes subequal in size. Anterior median eyes one diameter apart, two-thirds diameter LEVI: ACHAEARANEA, THERIDION AND SPHYROTINUS 133 from laterals. Posterior median eyes two-thirds diameter apart, a little more than one from laterals. Total length 2.5 mm. Cara- pace, 1.1 mm. long, 1.1 mm. wide. Second patella and tibia, 1.1 mm.; third, 0.9 mm.; fourth, 1.3 mm. Diagnosis. The epigynum having openings in a pair of nipples facing each other and a median depression (Fig. 310) separate this species from Theridion paidiscum. THERIDION SCIIMIDTI, New species Figure 331 Type. Male type from San José, Costa Rica (E. Schmidt), 1 the American Museum of Natural History. Description. Carapace, sternum, legs yellow. Abdomen gray with a median dorsal white band whose borders are irregular. Anterior median eyes one diameter apart, less than one-fourth from laterals. Posterior median eyes their radius apart, almost their diameter from laterals. Total length of male 1.9 mm. Cara- pace 0.80 mm. long, 0.63 mm. wide. First femur, 1.22 mm.; patella and tibia, 1.35 mm.; metatarsus, 1.08 mm.; tarsus, 0.53 mm. Second patella and tibia, 1.03 mm.; third, 0.74 mm.; fourth, 0.93 mm. Diagnosis. Details of the palpus (Fig. 331) separate this spe- cies from T. quantum. THERIDION PATIDISCUM, new species Figures 299-300, 329-330 Type. Male type from Boquete, Panama, Aug. 4-11, 1954 (A. M. Chickering), in the Museum of Comparative Zoology. Description. Carapace, sternum, dark dusky yellow. Legs yellow with dark dusky rings on middle and ends of segments. Abdomen mottled black with a median dorsal, white stripe, whose borders are irregular; several white spots on sides; venter without marks. Anterior median eyes one diameter apart, al- most touching laterals. Posterior eyes three-quarters diameter apart. Measurement of female total length, 2.3 mm. Carapace, 1.04 mm. long, 1.00 mm. wide. First femur, 1.32 mm.; patella and tibia, 1.46 mm.; metatarsus, 1.03 mm.; tarsus, 0.57 mm. Second patella and tibia, 1.12 mm.; third, 0.85 mm.; fourth, 1.18 134 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY mm. Male total length 1.7 mm. Carapace 0.91 mm. long, 0.78 mm. wide. First femur, 1.10 mm.; patella and tibia, 1.30 mm.; metatarsus, 0.78 mm.; tarsus, 0.52 mm. Second patella and tibia, 1.00 mm.; third, 0.73 mm.; fourth, 0.98 mm. Diagnosis. The epigynum (Fig. 300) has a pair of small selero- tized protuberances along the margin and a median small pro- tuberance which distinguishes it from 7. signaculum. The palpus (Mig. 330) resembles that of Sphyrotinus species; the various sclerites have shifted to the back of the bulb and are reduced in size. Record. Panama: Boquete, 2° paratype (A. M. Chickering). THERIDION SIGNACULUM, new species Figures 301-302, 333 Type. Male type from Forest Reserve, Canal Zone, July 4-6, 1939 (A. M. Chickering), in the Museum of Comparative Zool- ooy. Description. Carapace dusky white to dusky brown. Sternum dusky. Legs white with faint indications of rings or dusky patches on venter. Abdomen dark with a median dorsal white stripe, which is narrow on ends and has wavy sides; venter black with a white spot posterior to genital groove. Anterior median eyes less than one diameter apart, one-fourth a diameter from laterals. Posterior eyes separated by less than one diameter. The anterior median eyes of the male are larger than the others and less than their radius apart. Total length of females 1.6-2.0 mm. A female paratype measured total length 1.8 mm. Carapace 0.75 mm. long, 0.68 mm. wide. First femur, 0.91 mm. ; patella and tibia, 0.95 mm.; metatarsus, 0.65 mm.; tarsus, 0.39 mm. Second patella and tibia, 0.75 mm.; third, 0.56 mm.; fourth, 0.82 mm. Total leneth of male 1.6 mm. Carapace 0.71 mm. long, 0.62 mm. wide. First femur, 1.00 mm.; patella and tibia, 1.05 mm.; metatarsus, 0.75 mm.; tarsus, 0.42 mm. Second patella and tibia, 0.79 mm.; third, 0.60 mm.; fourth, 0.75 mm. Diagnosis. The female has an epigynum similar to 7’. paidis- cum, but it lacks the median protuberance (Fig. 302) ; the palpus of the male resembles 7. signum,; the radix, however, is smaller (Fig. 333). LEVI: ACHAEARANEA, THERIDION AND SPHYROTINUS 135 Records. Panama: Porto Bello (A. M. Chickering). Canal Zone: Madden Dam; Barro Colorado Isl. (several collections) ; Forest Reserve, 2, ¢ paratypes (all A. M. Chickering). THERIDION ATLIXCO, new species Figures 303-304 Type. Female type from Atlixeo, Puebla, June 26, 1947 (L. I. Davis), in the American Museum of Natural History. Description. Carapace dark gray, head region brown, reddish around eyes. Sternum black. Legs white with black spots or irregular narrow black rings. Abdomen with a median white band most distinct above spinnerets; and a white cross band on anterior half of dorsum. Venter with a black spot on each side of pedicel; a black ring around spinnerets; a wide black band from spinnerets to epigynum bearing a large white spot in cen- ter. Anterior median eyes one and one-quarter diameters apart, less than one-quarter from laterals. Posterior eyes two-thirds their diameter apart. Abdomen slightly wider than long, an- terior end widest, subtriangular. Total length, 1.6 mm. Carapace 0.70 mm. long, 0.69 mm. wide. First femur, 0.91 mm.; patella and tibia, 0.99 mm.; metatarsus, 0.66 mm.; tarsus, 0.32 mm. Second patella and tibia, 0.68 mm.; third, 0.50 mm.; fourth, 0.75 mm. Diagnosis. The indistinct epigynum (Fig. 304), covered by black pigment, differentiates this species from other Theridions. THERIDION PROGUM, new species Figures 305-306 Type. Female type from El Valle, Panama, July 1936 (A. M. Chickering), in the Museum of Comparative Zoology. Description. Carapace, sternum, legs dusky yellow-brown. Legs with indistinct darker rings. Abdomen black with a white stripe above spinnerets and several spots on sides. Anterior median eyes two-thirds diameter apart, almost touching laterals. Posterior median eyes one-third diameter apart, a radius from laterals. Total length 2.4 mm. Carapace 1.00 mm. long, 0.88 mm. wide. First femur, 1.32 mm.; patella and tibia, 1.40 mm.; metatarsus, 1.04 mm.; tarsus, 0.55 mm. Second patella and tibia, 1.04 mm.; third, 0.80 mm.; fourth, 1.15 mm. 136 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY Diagnosis. The openings of the ducts are on the ends of a swollen area on the epigynum (Fig. 306). The swollen area separates this species from 7. signaculum. THERIDION COLIMA, hew species Figures 277-279 Type. Female type from Miramar, Manzanillo, Colima, Jan. 15, 1948 (F. Bonet), in the American Museum of Natural His- tory. Description. Carapace, sternum, legs yellow. Legs dusky at joint of femur and patella. Abdomen whitish with a small dusky spot on dorsum and a dusky ring around spinnerets. Anterior median eyes one diameter apart, one-third of a diameter from laterals. Posterior median eyes one diameter apart, a radius from laterals. Total length of female 1.8 mm. Carapace 0.78 mm. long, 0.70 mm. wide. First femur, 1.14 mm.; patella and tibia, 1.12 mm.; metatarsus, 0.92 mm.; tarsus, 0.53 mm. Second patella and tibia, 0.78 mm.; third, 0.52 mm.; fourth, 0.88 mm. The generic placement of this species is not certain. It is pos- sible that it belongs to the genus Achaearanea. Diagnosis. The epigynum has a slight median elevation on whose sides are openings (Figs. 278, 279). This distinguishes the species from other theridiids known. THERIDION CLEMENS, new species Figures 322-324 Type. Male type from Blue Mountains, 5-7000 ft.; Jamaica, sritish West Indies, Aug. 17-19, 1934 (P. J. Darlington), in the Museum of Comparative Zoology. Description. Carapace, sternum, legs yellow. Abdomen with two dorsal longitudinal black bands, white in between and on sides ; sometimes with series of black spots. Diameter of anterior median eyes half that of others. Anterior median eyes one and one-third diameters apart, two-thirds diameters from laterals. Posterior median eyes one diameter apart, one and one-quarter diameters from laterals. Chelicerae of female enlarged. Measure- ment of female 2.4 mm. total length. Carapace 0.83 mm. long, 0.78 mm. wide. First femur, 1.56 mm.; patella and tibia, 1.69 LEVI: ACHAEARANEA, THERIDION AND SPHYROTINUS 17) mm.; metatarsus, 1.30 mm.; tarsus, 0.57 mm. Second patella and tibia, 1.19 mm.; third, 0.78 mm.; fourth, 1.24 mm. Male 1.9 mm. total length. Carapace 0.85 mm. long, 0.60 mm. wide. First femur, 1.69 mm.; patella and tibia, 1.91 mm.; metatarsus, 1.56 mm.; tarsus, 0.60 mm. Second patella and tibia, 1.26 mm.; third, 0.78 mm.; fourth, 1.20 mm. Diagnosis. The small anterior median eyes, the projecting median apophysis of the palpus (Fig. 324), and the structure of the epigynum (Fig. 323) distinguish this species from others. Records. Jamaca: Blue Mts., 5-7000 ft., 2 allotype, 9°, 4 paratypes (P. J. Darlington) ; Hardwar Gap, Blue Mts. (many collections) ; N. Moreces Gap, St. Andrew Par. (R. P. Bengry). THERIDION TEMPUM, new species Figures 315-316 Type. Female from the Forest Preserve, Panama Canal Zone, Dee. 24, 1957 (A. M. Chickering), in the Museum of Compara- tive Zoology. Description. Carapace dusky gray, darkest in eye region. Sternum dusky, lighter yellow in the middle. Endites and la- bium yellowish. Legs dusky gray. Abdomen yellow-gray with some white pigment spots on each side of dorsum, one black spot on each side of spinnerets. Eyes subequal in size. Anterior median eyes their diameter apart, almost touching laterals. Ab- domen globular, as igh as long. Posterior median eyes one diameter apart, two-thirds from laterals. Total length 2.3 mm. Carapace, 0.84 mm. long, 0.68 mm. wide. First femur, 1.04 mm.; patella and tibia, 1.04 mm.; metatarsus, 0.78 mm.; tarsus, 0.45 mm. Second patella and tibia, 9.74 mm.; third, 0.53 mm. ; fourth, 0.79 mm. Diagnosis. The epigynum having a sheht central depression and the long connecting ducts (Fig. 315) separate this species from Theridion plantatum. THERIDION PLANTATUM, new species Figures 255-256 Type. Kemale holotype from Plantation, Barro Colorado Island, Panama Canal Zone, July 19, 1954 (A. M. Chickering), in the Museum of Comparative Zoology. 138 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY Description. Carapace yellowish orange. Sternum orange. Legs yellow brown. Abdomen orange-white; posterior portion and area around spinnerets dusky; venter dusky. Anterior me- dian eyes shghtly larger than others, their diameter apart, one- quarter from laterals. Posterior eyes two-thirds diameter apart. Legs very strong. Total length of female 2.4 mm. Carapace 1.04 mm. long; 0.97 mm. wide. First femur, 1.56 mm.; patella and tibia, 1.85 mm.; metatarsus, 1.17 mm.; tarsus, 0.52 mm. Second patella and tibia, 1.04 mm.; third, 0.78 mm.; fourth, 1.24 mm. Diagnosis. The unusual coloration, the epigynum and internal evenltaha (Fig. 256) distinguish this species from others. THERIDION ARMOURI, new species Figures 259-260 Type. Kemale holotype from Armour Trail, Barro Colorado Island, Panama Canal Zone, July 17, 1954 (A. M. Chickering), in the Museum of Comparative Zoology. Description. Carapace orange-brown. Sternum orange. Legs dusky gray, distal segments darker. Abdomen dark gray above, reddish below. Anterior median eyes one diameter apart, almost touching laterals. Posterior eyes their diameter apart. Total length of female 2.0 mm. Carapace 0.71 mm. long. First femur, 0.88 mm.; patella and tibia, 0.98 mm.; metatarsus, 0.70 mm.; tarsus, 0.41 mm. Second patella and tibia, 0.71 mm.; third, 0.59 mm.; fourth, 0.85 mm. The relatively high abdomen may indicate that this species belongs to Achaearanea. Diagnosis. The epigvnum (Fie. 260) has a depression. The internal genitalia (Fig. 259) distinguish this species from others with a similar epigynum. THERIDION BRIDGESI, New species Figures 307-308, 328 Type. Male holotype from Pujal, San Luis Potosi, outside cave, March 1940 (W. Bridges), in the American Museum of Natural History. Description. Carapace yellowish with a median dusky band, sides dusky. Sternum dusky. Legs with dusky spots on ends LEVI: ACHAEARANEA, THERIDION AND SPHYROTINUS 139 of segments. Abdomen dusky black with a median dorsal white stripe whose edges are irregular. Sides with some white spots; venter with a white spot. Anterior median eyes two-thirds diameter apart, almost touching laterals; posterior eyes their radius apart. Total length of female 2.0 mm. Carapace 0.89 mm. long; 0.79 mm. wide. First femur, 1.30 mm.; patella and tibia, 1.387 mm.; metatarsus, 1.04 mm.; tarsus, 0.52 mm. Second patella and tibia, 1.12 mm.; third, 0.85 mm.; fourth, 1.17 mm. Total length of male 1.6 mm. Carapace 0.78 mm. long, 0.69 mm. wide. Diagnosis. The epigynum which is heavily pigmented and has two dark spots on the anterior end of a lightly sclerotized plate (Fig. 308) and the palpus which bears a thorn (conduc- tor) on the tegulum (Fig. 328) differentiates this species from others. A radix and median apophysis is present. Records. San Luis Potosi: Pujal, ¢@, ¢ paratypes (W. Bridges) ; Tamazunchale (W. J. Gertsch). THERIDION MUSAWAS, new species Figure 327 Type. Male type from Musawas, Rio Waspuc, Nicaragua, Oct. 10-31, 1955 (B. Malkin), in the American Museum of Natural History. Description. Carapace dusky yellow, darker in middle and near margins. Sternum, legs dusky yellow. Abdomen dark eray to black with indications of a median, dorsal white band. Anterior median eyes three-quarters diameter apart, less than one-quarter from laterals. Posterior median eyes one-third diameter apart, two-thirds from laterals. Posterior muscle scars on abdomen sclerotized. Total length of male 1.6 mm. Carapace, 0.80 mm. long, 0.58 mm. wide. First femur, 1.05 mm.; patella and tibia, 1.17 mm.; metatarsus, 0.78 mm.; tarsus, 0.36 mm. Second patella and tibia, 0.88 mm.; third, 0.65 mm.; fourth, 0.88 mm. Diagnosis. Details of the palpus (Fig. 327) separate this spe- cies from T. bridgest. THERIDION RINCONENSE, lew species Figures 261-262 Type. Female type from Rincon, Chiapas, April 6, 1953 (1. T. Davis), in the American Museum of Natural History. 140 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY Description. Carapace yellow, slightly dusky in center. Ster- num, legs yellow. Legs with indications of bands on venter. Abdomen spotted black, with a white line above spinnerets; venter yellowish. Anterior median eyes their diameter apart, one-fourth from laterals. Posterior median eyes their radius apart, two-thirds from laterals. Total length of female 2.5 mm. Carapace 1.04 mm. long, 0.91 mm. wide. First femur, 1.42 mm.; patella and tibia, 1.56 mm.; metatarsus, 1.05 mm.; tarsus, 0.68 mm. Second patella and tibia, 1.17 mm., third, 0.89 mm.; fourth, 1.20 mm. Diagnosis. The epigynum (Fig. 262) differentiates this spe- cies from other Theridion. THERIDION PANUM, new species Figures 289-290 Type. Female type from El Volcan. Chiriqui, Panama, March 7, 1986 (W. J. Gertsch), in the American Museum of Natural History. Description. Carapace yellow with dusky head* and a black line around margin. Legs yellow with indications of rings. Abdomen black with a median dorsal white band having seal- loped edges and narrowing at the posterior ; a white band around anterior edge; some white pigment on each side of pedicel on venter. Anterior median eyes slightly larger than others, their diameter apart, one-fourth their diameter from laterals. Pos- terior eyes one diameter apart. Total length of female 2.6 mm. Carapace 1.04 mm. long, 0.78 mm. wide. First femur, 1.87 mm. ; patella and tibia, 2.05 mm.; metatarsus, 1.60 mm.; tarsus, 0.73 mm. Second patella and tibia, 1.42 mm.; third, 0.91 mm.; fourth, 1.46 mm. Diagnosis. The structure of the very small epigynum (Fie. 290) separates this species from 7. moctezwma. THERIDION QUEMADUM, new species Figures 257-258 Type. Kemale type from Cruz Quemada, 2000 m., nr. Amate- nango, Chiapas, July 24, 1950 (C. and M. Goodnight), in the American Museum of Naturai History. LEVI: ACHAEARANEA, THERIDION AND SPHYROTINUS 141 Description. Carapaee yellow, head dusky, border with a black line. Sternum dusky yellow. Legs yellow with ends of segments dusky. Abdomen black with lighter patches on sides of dorsum and a light stripe above spinnerets. Anterior median eyes slightly smaller than others, their diameter apart, one- fourth from laterals. Posterior eyes two-thirds diameter apart. Total leneth 2.9 mm. Carapace 1.15 mm. long, 1.04 mm. wide. First femur, 1.71 mm.; patella and tibia, 1.89 mm.; metatarsus, 1.41 mm.; tarsus, 0.65 mm. Second patella and tibia, 1.28 mm. ; third, 0.94 mm.; fourth, 1.49 mm. Diagnosis. The epigynum (Fig. 258) distinguishes this species from 7. morulum. SPHYROTINUS Simon Sphyrotinus Simon, 1894, Proc. Zool. Soc. London, p. 524. Type species by monotypy: S. luculentus Simon. Hypobares Simon, 1894, op. cit., p. 552. Type species H. unisignatus Simon by original designation and monotypy. Philto Simon, 1894, op. cit., p. 552. Type species P. subtilis Simon by origi- nal designation and monotypy. Hubba O. P. Cambridge, 1897, Biologia Centrali Americana, Araneidea, vol. 1, p. 231. Type species by monotypy H. insignis O. P. Cambridge. Thonastica Simon, 1909, Bull. Biol. France et Belgique, vol. 42, p. 92. Type species 7. praemollis Simon by original designation and monotypy. Garricola Chamberlin, 1916, Bull. Mus. Comp. Zool., vol. 60, p. 233. Type species by original designation and monotypy G. sanctus Chamberlin. Paidisca Bishop and Crosby, 1926, Jour. Elisha Mitchell Sci. Soe., vol. 41, p. 178. Type species by original designation and monotypy Histagonia marai Crosby. Spelobion Chamberlin and Ivie, 1938, Publ. Carnegie Inst. Washington, no. 491, p. 138. Type species by original designation and monotypy S. spukilum Chamberlin and Ivie (= Sphyrotinus luculentus Simon). Thymoella Bryant, 1938, Bull. Mus. Comp. Zool., vol. 100, p. 377. Type species by original designation 7. banksi Bryant. Brontosauriella Bristowe, 1938, Ann. Mag. Nat. Hist., ser. 11, vol. 2, p. 72. Type species B. melloleitaoni Bristowe by monotypy. The members of this genus appear similar. The cephalothorax with appendages are brown and the abdomen, which may have dusky marks, is whitish. The legs are shorter than those of Theridion. The carapace may be modified in males like that of erigonids, sometimes the first legs are enlarged. In the northern 142 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY part of the region the species are often heavily sclerotized. Figs. 1-3. Achaearanea turquino, new species. 1. Female genitalia, dor- sal view. 2-3. Epigynum. 2. Ventral view. 3. Lateral view. Figs. 4-8. A. hermosillo, new species, 4. Female genitalia, dorsal view. 5. Epigynum) 6-7. Left palpus. 6. Ventral view. 7. Ectal view. 8. Female. Figs. 9-12. A. apex, new species. 9. Female genitalia, dorsal view. 10. Epigynum, 11. Memale abdomen, lateral view. 12. Palpus. Figs. 13-15. 4. anna, new species. 13. Palpus. 14. Female genitalia, dor- sal view. 15. Epigynum. Fig. 16. A. florens (O. P. Cambridge), palpus. Fig. 17. A. florendida, new species, palpus. Figs. 18-19. Achaearanea florens (O. P. Cambridge). 18. Female geni- talia, dorsal view. 19. Epigynum. Figs. 20-21. A. florendida, new species. 20. Female genitalia, dorsal view. 21. Epigynum. Figs. 22-23. A. serax, new species. 22. Female genitalia, dorsal view. 23. Epigynum. Figs. 24-25. A. schraderorum, new species. 24. Female genitalia, dorsal view. 25. Epigynum. Fig. 26. A. schneirlai, new species, left palpus. £ > ’ Figs. 27-28. A. manzanillo, new species. 27. Female genitalia, dorsal view. , ) 28. Epigynum. Figs. 29-31. 4. trinidensis, new species. 29. Palpus. 30. Female genitalia, dorsal view. 31. Epigynum. Figs. 32-34. A. zonensis, new species. 32. Female genitalia, dorsal view. 33. Epigynum. 34. Palpus. Figs. 35-38. Achaearanea hirta (Taczanowski). 35. Female genitalia, dor- sal view. 36. Epigynum. 37. Left palpus. 38. Female. Figs. 39-41. 4A. taeniata (Keyserling). 39. Female genitalia, dorsal view. 40. Epigynum. 41. Palpus. Figs. 42-44. 4. pura (O. P. Cambridge). 42. Female genitalia, dorsal view. 43. Epigynum. 44. Palpus. Figs. 45-50. A. maricaoensis (Bryant). 45. Female. 46. Epigynum, cleared. 47. Female genitalia, dorsal view. 48. Epigynum. 49-50. Palpus. 49. (Panama). 50. (Puerto Rico). Fig. 51. A. machaera, new species, palpus. Fig. 52. Achaearanea terex, new species, left palpus. I , Figs. 53-56. A. mesax, new species. 53. Female. 54. Female genitalia, dorsal view. 55. Epigynum. 56. Palpus. Figs. 57-60. A. picadoi (Banks). 57, 59. Female genitalia, dorsal view. 58, 60. Epigynum. 57-58. (Panama). 59-60. (Costa Rica). Figs. 61-63. A. oblivia (O. P. Cambridge). 61. Female genitalia, dorsal view. 62-63. Epigynum, 62. Ventral view. 63. Posterior view. Figs. 64-66. A. rostrata (O. P. Cambridge). 64. Female genitalia, dorsal view. 65. Epigynum. 66, Lateral view. Figs. 67-69. A. nayaritensis, new species. 67. Female genitalia, dorsal view. 68-69. Epigynum. 68. Ventral view. 69. Lateral view. Figs. 70-71. ( > 234 236 eS ~~ Bae 250-252. Figs. sal view. 251. Epigynum. 252. Left palpus. Oe Figs. 253-254. 7. 254. Epigynum. Figs. 255-256. VT. plantatum, new species. view. 256. Epigynum., Figs. 257-258. 7. quemadum, new species. view. 258. Epigynum. Figs. 259-260. view. 260. Epigynum. T. armouri, new species. Figs. 261-262. view. 262. Epigynum. T. rinconense, new species. Figs. 263-267. 7. pananense, new species. resum, new species. 253. Theridion akron, new species. 250, Female genitalia, dor- Female genitalia, dorsal view. 255. Female genitalia, dorsal 257. Female genitalia, dorsal 259. Female genitalia, dorsal 261. Female genitalia, dorsal 263. Female genitalia, dorsal view. 264. Epigynum. 265-266. Palpus (same locality). 267. Female abdo- men, ventral view. Figs. 268-271. 7. chiriqui, new species. 268. Palpus. 269. Male abdomen, dorsal view. 270. Female genitalia, dorsal view. 271. Epigynum. Figs. 272-276. Theridion lathropi, new species. 272. Female genitalia, dorsal view. 273. Epigynum, lateral view. 274. Epigynum. 275-276, Left aed palpus. 275. Mesal view, expanded. 276. Ventral view. Figs. 277-279. T. colima, new species. 277. Female genitalia, dorsal view. 278. Epigynum, lateral view. 279. Epigynum. Figs. 280-284. 7. petrum, new species. 280. Female genitalia, dorsal view. 281. Epigynum, lateral view. 282. Epigynum. 283. Female. 284. Palpus. Figs. 285-288. 7. moctezuma, new species. 285. Female genitalia, dorsal view. 286. Epigynum. 287. Female abdomen, dorsal view. 288. Palpus. Figs. 289-290. T. panum, new species. 289. Female genitalia, dorsal view. 290. Epigynum. Abbreviations: C, conductor; EL, embolus; M, median apophysis; &, radix, Figs. 291-292. Theridion cobanum, new species. dorsal view. 292. Epigynum. Figs. 293-294. T. quantum, new species. view. 294. Epigynum. Figs. 295-296. 7. mallini, new species. view. 296. Epigynum. Figs. 297-298. T. signwm, new species. view. 298. Epigynum, Figs. 299-300. 7. paidiscum, new species. view. 300. Epigynum. Figs. 301-302. 7. signaculum, new species. view. 302. Epigynum. Figs. 303-304. 7. atlixco, new species. view. 304. Epigynum., Figs. 305-306. 7. progum, new species. view. 306. Epigynun. Figs. 307-308. T. bridgesi, new species. view. 308, Epigynum. 293. 295 297 299. 301. Female 303% 305. DOM. Female Female Female Female Female Female Female genitalia, genitalia, genitalia, genitalia, genitalia, genitaha, genitalia, genitalia, 291. Female genitalia, dorsal dorsal dorsal dorsal dorsal dorsal dorsal dorsal OO 00" aoe See ; Figs. 309-310. Theridion chilapa, new species. 309. Female genitalia, dorsal view. 310. Epigynum. Figs. 311-312. 7. aspersum (F. P. Cambridge). 311. Female genitalia, dorsal view. 312. Epigynum. Figs. 313-314. 7. artum, new species. 313. Female genitalia, dorsal view. 314. Epigynum. Figs. 315-316. 7. tempum, new species. 315, Female genitalia, dorsal view. 316. Epigynum. Fig. 317. Theridion quantum, new species, left palpus. Figs. 318-321. 7. reservum, new species. 318. Female genitalia, dorsal view. 319. Epigynum. 320. Female abdomen, dorsal view. 321. Palpus. Figs. 322-324. T. clemens, new species. 322. Female genitalia, dorsal view. 323. Epigynum. 324. Palpus. Fig. 325, 7. rwinwm, new species, palpus. Fig. 326. T. marvum, new species, palpus. Fig. 327. 7. musawas, new species, palpus. Fig, 328. 7. bridgesi, new species, palpus. Figs. 329-330. T. paidiscum, new species, palpus. 329. Expanded, dorsa! view, cymbium removed. 330. Ventral view. Fig. 331. 7. schmidti, new species, palpus. Fig. 332. T. signum, new species, palpus. Fig. 333. 7. signaculum, new species, palpus. = Abbreviations: C, conductor; #, embolus; W, median apophysis. Figs. 334-338. Sphyrotinus luculentus, Simon. 334. Female genitalia, dor sal view. 335. Epigynum (St. Vincent Island). 336-338. Left palpus. 336. (Syntype from St. Vincent Island). 337 .(Panama). 338. (Chiapas). Figs. 339-341. S. chickeringi, new species. 339. Female genitalia, dorsal view. 340. Epigynum. 3841. Palpus. Figs. 342-344. S. illudens (Gertsch and Mulaik), (Panama). 342. Female genitalia, dorsal view. 3843. Epigynum. 344. Palpus. Figs. 345-349. S. guanicae (Petrunkevitch), (Jamaica). 345, 347. } Epigynum. 346. Female genitalia, dorsal view. 348-349. Palpus. 335 340 Figs. 350-356. Sphyrotinus maderae (Gertsch and Archer), left palpus. 350. (Nuevo Leén). 351. (Veracruz). 352-353. (Costa Rica). 354. (Pan- ama). 355. Female genitalia, dorsal view (Panama). 356. Epigynum (Pan- ama). 7 Figs. 357-359. S. verus, new species. 357-358. Carapace of male. 359. Palpus. Figs. 360-362. 8S. delicatulus, new species. 360. Palpus. 361. Female genitalia, dorsal view. 3862. Epigynum. Figs. 363-364. S. marai (Crosby), (Chiapas). 363. Female genitalia, dorsal view. 364. Epigynum. Figs. 365-366. S. expulsus (Gertsch and Mulaik), (Cuba). 3865. Female genitalia, dorsal view. 366. Epigynum. Figs. 367-368. Sphyrotinus chiapensis, new species. 367. Female geni- talia, dorsal view. 368. Epigynum, Figs. 369-370. S. boneti, new species. 369. Female genitalia, dorsal view. 370. Epigynum, Figs. 371-372. S. bradti, new species. 371. Female genitalia, dorsal view. 372. Epigynum. Figs. 373-374. S. corus, new species. 373. Female genitalia, dorsal view. 374. Epigynum. Figs. 375-376. S. reservatus, new species. 375. Female genitalia, dorsal > = ise (a>) 4 ww =~] ron ies) pigynum, Figs. 377-381. S. indicatus (Banks). 377. Female genitalia, dorsal view. 378. Epigynum. 379-380. Male carapace. 381. Left palpus. Fig 382. S. confraternus (Banks), palpus. Figs. 383-384. S. orilla, new species. 383. Palpus. 384. Male abdomen, lateral view. < 367 ™ A 369 | 37 Figs. 385-389. Sphyrotinus stylifrons (Simon). 385-386. Male carapace. 387. Female genitalia, dorsal view. 388. Epigynum. 389. Left palpus. Figs, 390-394. 391. Epigynum. Figs. 395-397. Palpus. Figs. 398-400. Figs. 401-403. S. piarco, new species. 390. Female genitalia, dorsal view. 392-393. Male carapace. 394. Palpus. S. matachic, new species. 395-396. Male carapace. 397. S. banksi (Bryant). 398-399. Male carapace. 400. Palpus. S. simla, new species. 401-402. Male carapace. 403. Palpus. Ss a / eS 385 yO —~ esi 390 | ay ) 395 ING fs \ mu ys / \ « fyi SS —~D \ ea ey wie — CES Figs. 404-410. Sphyrotinus insignis (O. P. Cambridge). 404. Female geni- talia, dorsal view. 405. Epigynum. 406-407. Male carapace. 408-410. Left palpus. 408. Mesal view, expanded. 409. Mesal view. 410. Ventral view. Figs. 411-416. S. notabilis, new species. 411. Female genitalia, dorsal view. 412. Epigynum. 413-414. Male carapace. 415-416. Palpus. 415. Mesal view. 416. Ventral view. Figs. 417-418. S. bogus, new species. 417. Female genitalia, dorsal view. 418. Epigyniun. Abbreviations: C, conductor: FE, embolus; WM, median apophysis: FR, radix. Figs. 419-422. Sphyrotinus boquete, new species. 419. Left palpus. 420. Epigynum, lateral view. 421. Female genitalia, dorsal view. 422. Epigynum. Figs. 423-426. S. prolatus, new species. 423. Male cephalothorax and palpus. 424. Palpus. 425. Female genitalia, dorsal view. 426. Epigynum. Figs. 427-428. S. deprus, new species. 427. Female genitalia, dorsal view. 428. Epigynum. Figs. 429-430. S. vivus (O. P. Cambridge). 429. Epigynum cleared. 430. Epigynum. pace oot) : Vira sor ech tet : Ee Ses! F A ar, ae _ Bulletin of the Museum of Comparative Zoology ATH ARVIAR DC Olin GE VoL. 121, No. 4 THREE NEW GENERA AND ONE NEW SPECIES OF THE FAMILY GONOSTOMATIDAE By MARION GREY Chicago Natural History Museum CAMBRIDGE, MASS., U.S.A. Pee DT ED) ae OF i TEE) DP MisUsS Br UCM JuLy, 1959 PUBLICATIONS ISSUED BY OR IN CONNECTION WITH THE MUSEUM OF COMPARATIVE ZOOLOGY AT HARVARD COLLEGE BULLETIN (octavo) 1863 — The current volume is Vol. 121. Breviora (octavo) 1952 — No. 111 is current. Memoirs (quarto) 1864-1938 — Publication was terminated with Vol.55. JOHNSONIA (quarto) 1941 — A publication of the Department of Mollusks. Vol. 3, no. 38 is current. OCCASIONAL PAPERS OF THE DEPARTMENT OF MOLLUSKS (octavo) 1945 — Vol. 2, no. 22 is current. PROCEEDINGS OF THE NEW ENGLAND ZoouoaicaL CuuB (octavo) 1899-1948 — Published in connection with the Museum. Publication terminated with Vol. 24. The continuing publications are issued at irregular intervals in num- bers which may be purchased separately. Prices and lsts may be obtained on application to the Director of the Museum of Comparative Zoology, Cambridge 38, Massachusetts. Of the Peters ‘‘Cheek List of Birds of the World,’’ volumes 1-3 are out of print; volumes 4 and 6 may be obtained from the Harvard Uni- versity Press; volumes 5 and 7 are sold by the Museum, and future volumes will be published under Museum auspices. Bulletin of the Museum of Comparative Zoology PX De ANG ARS WEIN 1 DY (OPO) ADIL, DOCEP 1S. Vote iZin Now THREE NEW GENERA AND ONE NEW SPECIES OF THE FAMILY GONOSTOMATIDAE By MARION GREY Chicago Natural History Museum CAMBRIDGE, MASS., U.S.A. Pan Lee LOR. SEES Mw SUM JuLY, 1959 No. 4 — Three New Genera and One New Species of the Family Gonostomatidae By Marion GREY Chieago Natural History Museum In the course of revising the family Gonostomatidae for a forthcoming volume on the Fishes of the Western North Atlan- tic, it was found necessary to create several new genera. And since it has been decided that descriptions of new genera and species should not be included in that monograph, we hasten to give such original descriptions in short papers which will ante- date the larger work. Two of the genera described below are based on previously known species. The third genus, hitherto unknown, is described from specimens trawled in 1957 aboard the U.S. Fish and Wild- life Service research vessel Oregon. Figure 1 has been prepared by Mr. John Pfiffner, Staff Artist, Chicago Natural History Museum, and Figures 2 and 3 by Mrs. Myvanwy M. Dick, Museum of Comparative Zoology. Terminology. The following symbols are used to represent the photophores: ORB, those situated close to the eye; OP, opereu- lar photophores; SO, a pair often found near the symphysis of the lower jaw; BR, organs on the branchiostegal membranes ; IV, pre-ventral photophores of the ventral series; VA, those of the ventral series found between the ventral bases and the anal origin; AC, photophores of the ventral series posterior to the anal origin; IC, total number in the ventral series, from tip of isthmus to base of caudal; OA, photophores of the lateral series. Photo- phore counts in parentheses indicate that these organs are grouped in a common gland. POLLICHTHYS, new genus Yarrella Jespersen and Taning, 1919, Vidensk. Medd. Dansk Naturh. Foren., 70:223; Norman, 1930, Discovery Rep., 2:288 (part); Fowler, 1936, Bull. Amer. Mus. Nat. Hist., 70:1202 (part); Lozano Rey, 1947, Mem. R. Acad. Cien. Madrid, Ser. Cien. Nat., 11:171. Type species: Yarrella mauli Poll. Generic characters. Eye normal, moderate. Snout slightly longer than orbit. Interorbital width at center of eye about 168 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY equal to or a little less than diameter of orbit. Mouth large, oblique; edge of premaxillary straight ; toothed edge of maxillary curving downward from juncture with premaxillary, becoming almost straight posteriorly, nearly reaching preopercle. Pre- maxillary about half as long as toothed portion of maxillary. Angele of preoperecle acute. Upper jaw with a single row of teeth, consisting of rather evenly spaced longer teeth and one or two smaller ones in each interspace, some of the interspace teeth curving inward. Teeth of lower jaw similar but smaller, and, anteriorly only, an outer row of very small teeth. Vomer toothless or with one or two small teeth on each side. Palatines each with a short row of small teeth. Pterygoids with or without a patch of very minute teeth on each side. Tongue usually with an irregular row of minute teeth centrally, not reaching tip. Gill rakers 11-12 + 4-5 on first arch. Spines on inner edge of first gill areh very short, a cluster of minute prickles below each one. No pseudobranchiae. Scales present but very decidu- ous. Anus closer to anal fin than to ventral bases, beneath fifth to seventh VA photophore. Head and trunk longer than tail. Dorsal origin about in middle of body length or slightly posterior to it. Anal origin beneath front or middle of dorsal fin. Ventral bases well ahead of dorsal origin. Adipose fin rather long-based, situated above a vertical from shghtly behind middle of anal fin. ORB 2, one close to and below front margin of eye, the other beneath posterior margin of eye, the anterior one larger and apparently double, the posterior one similar in form to body photophores. OP 3, about equal in size, upper one level with upper edge of pupil, lower ones level with end of maxillary. SO present. BR 8. Body with two rows of photophores; photo- phores present on isthmus. IV 21-23, eighth and/or ninth slightly displaced. VA 7-9. AC 18-21, straight, four or five of them behind anal fin. IC 47-50. OA 19-21, reaching to a vertical from last one or two VA or first AC photophore. No other photo- phores, and no luminous tissue, on head or body as far as known. Fin rays: dorsal 10-12, anal 22-30, pectoral 8, ventral 6-7. Branchiostegal rays 11-12, no spines at bases. Vertebrae perhaps around 40 (count made on a faint X-ray photograph of one specimen ). GREY: NEW GENERA OF GONOSTOMATIDS 169 Remarks. Pollichthys is probably most closely related to Vin- ciguerria Jordan and Evermann, from which it differs principally in having a longer and more forwardly placed anal fin. It is also similar to Polymetme MeCulloch in some respects, and its dentition is rather like that of Gonostoma Rafinesque. It is not closely related to Yarrella Goode and Bean, with which it has been confused for many years. Yarrella blackfordi Goode and Bean has been found to possess serial photophores on the body above the IC and OA, arranged like those of Lychnopoles argen- teolus Garman. And the genus Yarrella, of which Lychnopoles Garman is a synonym, contains only two species, blackfordi and argenteola. The new genus has been named for Dr. Max Poll of the Musée Royale du Congo Belge, Tervuren, Belgium. POLLICHTHYS MAULI (Poll) Yarrella mauli Poll, 19538, Rés. Sci. Exp. Océanogr. Belge (1948-1949), vol. 4, fase. 2, pt. III, p. 59, fig. 24, (5°15’ S., 11°29’ E., 225-240 meters). Yarrella blackfordi Jespersen and Taning, 1919, Vidensk. Medd. Dansk Naturh. Foren, 70:223, pl. 17, fig. 13; Breder, 1927, Bull. Bingham Oceanogr. Coll., 1, (1), p. 18 (ve-identified as P. mauli by Morrow, 1958, in litt.); Norman, 1930, Discovery Rep., 2:288, fig. 7; Jespersen, 1933, Faune Ichth. Atl. Nord, 14, fig.; Parr, 1934, Bull. Bingham Oceanogr. Coll., 4 (6), p. 5 (ve-identified as P. mauli by Morrow, 1958, in litt.); Fowler, 1936, Bull. Amer. Mus. Nat. Hist., 70:1203 (part), fig. 518; Beebe, 1937, Zoologica, 22:201; Parr, 1937, Bull. Bingham Oceanogr. Coll., 3 (7), p. 45 (re-identified as P. mauli by Morrow, 1958, in litt.); Lozano Rey, 1947, Mem. R. Acad. Cien. Madrid, Ser. Cien. Nat., 11:172, fig. 42; Grey, 1955, Fieldiana, Zool., 37:272. Specimens reported as Yarrella blackfordi by Marshall (1951, p. 3, and 1954, p. 204, fig. VIII, 10) probably belong to this species also. The generic diagnosis has been based both on published de- scriptions and the following specimens, some of them hitherto unrecorded. Bermuda, Caryn: Thirteen specimens, all somewhat shrunken since earlier measurements were made (Grey, 1955, p. 272). Haul 20, 400-450 meters, one, standard length 32 mm. Haul 22, 730-820 meters, three, 45-48.5 mm. Haul 23, 400-450 meters, two, 29.5-ea. 31.5 mm. Haul 25, 400-450 meters, one, 39 mm. Haul 170 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 26, 590-660 meters, two, 38 and ca. 45 mm. Haul 27, 1280-1370 meters, one, 34.5 mm. Haul 44, 400-450 meters, one, 38 mm. Haul 56, 260-275 meters, one, ca. 38 mm. Haul 62, 1000-1100 meters, one, 39 mm. Bermuda: Five specimens, Stanford University No. 46475, recorded as Yarrella blackfordi by Beebe (1937, p. 201), Ber- muda Oceanographic Expeditions, 1929-1930, of the New York Zoological Society. Net 861, 1280 meters, one, 16 mm. Net 867, 1463 meters, one, 20.5 mm. Net 888, 732 meters, one, 33.5 mm. Net 897, 1463 meters, one, ca. 12.5 mm. (but shrunken). Net 954, 1829 meters, one, ca. 13.5 mm. (but shrunken). 3ermuda: One specimen, U.S.N.M. No. 100537, 36 mm., Grampus (Bache) Station 10176, 32° 30’ N., 65° 48’ W., 5 Feb- ruary 1914, 750 meters. Atlantie off Florida: One specimen, U.S.N.M. No. 100627, 39.5 mm., Grampus (Bache) Station 10202, 25° 34’ N., 79° 24’ W., 19 March 1914, 75 meters. Atlantie off Florida: Fifteen specimens, ca. 16-37.5 mm., Miami University Marine Laboratory Collection, taken in plank- ton hauls made in ca. 25° N., 79° W., under the auspices of the National Geographie Society. Depth of capture 503 or 686 meters, one, front part of body only; 457 meters, one, standard leneth 25 or 26 mm.; 439-541 meters, one, 22 mm.; 212-366 me- ters, one, 24.5 mm.; 366 meters, one, metamorphosing, 17 mm.; 210 meters, one, 23.5 mm.; 155 meters, one, metamorphosing, 16-16.5 mm.; 137 meters, one, 26.5 mm.; 119 meters, one, 27.5 mm.; 110 meters, one, 17 mm.; 55 meters, one, 24 mm.; 29-66 meters, one, 22.5 mm.; 15 meters, one, 37.5 mm.; depth un- known, two, 17.5 and 36 mm. Caribbean Sea: One specimen, U.S.N.M. No. 108281, ea. 37 mm., Caroline, off the Virgin Islands, 18° 35’ 80” N., 65° 23’ 54” W., 26 February 1933, 300 fathoms (549 meters). Caribbean Sea off Nicaragua, Oregon: One specimen, ea. 44.5 mm., Station 1905, 12° 22’ N., 82°. 27’ W., 11 September 1957, 275 fathoms (503 meters). One specimen, 45 mm., Station 1907, 12° 25’ N., 82° 23’ W., 11 September 1957, 400-425 fathoms (732- 777 meters). Western Pacific off eastern Negros, Philippines: Six specei- mens, 16.5-40 mm., Albatross Station 5190, 10° 8’ 15” N., 123° 16’ 45” B., 295 fathoms (540 meters). GREY: NEW GENERA OF GONOSTOMATIDS sya! P. mauli has not been found, so far, in the Gulf of Mexico, but is known from a large area iu the North Atlantic west of 35° N. and south of 40° N., as well as off the African coast between 5° and 11° 8. Although it has not previously been reported outside the Atlantic, the Chicago Natural History Museum collection contains six specimens from the Philippines, received in ex- change from the United States National Museum. The following counts have been made on Atlantic specimens : dorsal rays 10-12, anal rays 25-26, pectoral rays 8, ventral rays 6-7, branchiostegal rays 11-12, gill rakers on first arch 11-12 + 5. The photophores are small but conspicuous: BR 8; IV 22, eighth and/or ninth slightly elevated; VA 7-9, scarcely separable from AC; AC 18-20, four or five of them behind anal fin; IC 49-50; OA 19-21, reaching a vertical from the next-to-last VA, the last VA, or the first AC photophore. Individual variation is rather sheht and it is interesting that specimens from the Pacific differ very little from Atlantic speci- mens. The largest of these, 40 mm. in standard length, has a slightly greater depth and more posterior insertion of the dorsal fin, lacks the vomerine and pterygoid teeth, and has only 47 photophores in the ventral row (IC), but is otherwise indis- tinguishable from Atlantic specimens. Of the smaller examples, 16.5-26 mm. in standard length, four also have an IC count of 47, and in the fifth it is 48. Counts of the largest Philippine specimen are as follows: dorsal rays ca. 10, anal rays ca. 24, branchiostegal rays 11 or 12, gill rakers on first arch 11 -+ 4. Photophores: BR 8; IV 21, the eighth and tenth slightly below, and the ninth slightly above, the level of the first seven; VA 7; AC 19, five of them behind the anal fin; IC 47; OA 21, reaching a vertical from the first AC photophore. The following measurements are expressed in per cent of the standard leneth, 40 mm., and are followed in parentheses by similar measurements of nine western Atlantic specimens, whose standard lengths are 37.5-48.5 mm. Depth 15.0 (11.9-14.5) ; head 23.8 (20.6-ca. 24.4) ; snout ca. 5.0 (5.1-5.55) ; orbit ca. 3.75 (2.56-4.45) ; interorbital width at center of eye ca. 2.5 (2.22-3.33) ; upper jaw 16.2 (15.4-17.9) ; premaxillary 5.0 (5.12-6.4) ; toothed edge of maxillary 11.2 (10.0-11.5) ; tip of snout to dorsal origin 52.5 (49.3-51.5), to anal origin 55.0 (53.3-56.6), to ventral bases 172 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 41.5 (ca. 39.0-43.3) ; distance between anal origin and base of middle caudal rays 42.5 (41.0-44.7), last anal ray and base of middle caudal rays 11.2 (10.2-13.1), last dorsal ray and base of middle caudal rays 33.8 (ca. 33.3-37.1), last dorsal ray and adipose fin 13.7 (12.8-15.8), ventral bases and anal origin ¢a. 12.5 (18.5-15.4) ; least depth of caudal peduncle 3.75 (3.1-4.45) ; dorsal base ca. 8.75-10.0 (10.2-12.4); anal base ea. 31.2 (ea. 28.9-ca. 31.6). Metamorphosing specimens. Postlarval stages of P. mauli have not yet been described, but among the young fishes in the collee- tion of the Miami University Marine Laboratory are two meta- morphosing specimens. The younger of these is about 17 mm. in standard length. It is somewhat more slender than the older fish, which is 16-16.5 mm. in standard length; and it has the anal insertion farther back on the body, beneath the end of the dorsal fin. The posterior position of the anal fin might have prevented positive identification if the older specimen had not been at hand for comparison. The two little fishes obviously belong to the same species and it can only be assumed that during meta- morphosis the body shrinks in length between the ventral bases and the anal origin, resulting in a more anterior insertion of the anal fin. This shrinkage undoubtedly occurs just in front of the anal fin, which, in the younger postlarva, is separated by a short gap from the last VA photophore. In appearance, the two postlarvae from Florida are similar to metamorphosis stages of Vinciguerria. They differ most notice- ably in the complete absence of pigment at the end of the caudal pedunele, and in having a longer tail and longer anal fin. The development of Pollichthys probably more or less parallels that of Vinciguerria in 1) having the photophores mostly formed, but unpigmented, prior to metamorphosis; 2) having most of the photophores pigmented simultaneously; 3) a rather sudden transition from the postlarval stage to the metamorphosis stage ; and 4) the changes in body proportions proceeding rapidly toward the adult form. The smallest metamorphosed specimen known was 16 mm. in standard leneth (Jespersen and Taning, 1919, p. 223). It was not described. The smallest adolescents seen from the western Atlantic (off Florida) are 17 and 17.5 mm. in standard leneth and differ from adults only in having a GREY: NEW GENERA OF GONOSTOMATIDS hie more slender body, less dorsal pigment, and only 14-15 OA photophores. Some of the more posterior OA and middle AC photophores are not fully developed on these little fishes, and even in specimens as large as 26-27 mm. the most posterior OA organs are still not fully developed. The SO must appear quite early in this species as it is present in all adolescent specimens seen, including four from the Philip- pines. As in Vineiguerria it is the OA photophores that are last to develop, the full complement not being acquired until a standard length of 23-24 mm. has been reached. The number of OA on small Atlantic specimens is 14-15 at a length of 17 and 17.5 mm., 17-18 at a length of 22-23.5 mm. In the small Philip- pine specimens counts are 12 at 16.5 mm., 14 at 18 mm., 13 and 16 at 20 mm. Description of younger metamorphosing specimen, standard leneth ca. 17 mm.: Eye oval shaped. Dorsal rays 10 or 11. Anal rays uncertain, at least 22 or 23, origin of anal beneath last dorsal rays. Pectorals small, pedunculate. Ventrals very small, membranous. Adipose fin long-based, well developed. Ventral finfold present on posterior half of distance between ventrals and anal. End of eut shehtly protruding just in front of anal fin. Photophores on head: ORB 2, situated as in adult ; lower posterior OP; BR 8; 1V, 6 on isthmus, 13 from just in front of pectoral base to ventral base, the two groups separated by a short space in which one unpigmented organ is barely discern- ible. VA 9, the last two small, ending a short distance in front of anal origin. AC small and faintly pigmented above anal fin, 9 on left side, 11 on right side, the last two scarcely pigmented ; and 2 larger ones on caudal peduncle. OA, 3 on left side, 1 on right side, faintly pigmented. Head and body entirely without pigment excepting the brown photophores; end of caudal ped- uncle transparent. Description of older metamorphosing specimen, standard leneth ca. 16-16.5 mm.: Eye round. Dorsal rays 10. Anal rays uncertain, more than 20, origin of anal fin beneath fifth dorsal ray. Pectorals short, not pedunculate. Ventrals short, with rays developed. Adipose fin long-based, well developed. End of gut shghtly protruding, anus below space between seventh and eighth VA photophore. Photophores on head: ORB 2; OP 3, the lower posterior the largest, the lower anterior and upper 174 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY ones small and lighter in color; BR 8, IV 22, the ninth shghtly elevated. VA 8, not separated from anal origin or from AC. AC 18, the fifteenth poorly developed, the two or three pre- ceding it very small, and the last three well developed. OA 5. Head without pigment except on photophores (brown). Body mostly colorless but with a streak of black internal abdominal pigment and very faint indications of the internal spots along base of anal fin. Woobs!a, new genus Type species. Photichthys nonsuchae Beebe, 1932. Generic characters. Kye normal, large. Snout shorter than orbit. Interorbital width at center of eye less than diameter of orbit. Mouth large, oblique ; edges of premaxillary and maxillary straight ; maxillary reaching well past eye but not quite to edge of preopercle. Premaxillary more than half as long as toothed portion of maxillary. Angle of preopercle slightly acute. Teeth of upper jaw uniserial, larger on premaxillary than on maxil- lary, a few minute teeth between larger ones. Lower jaw with a series of widely spaced teeth, a few small interspace teeth and, anteriorly only, an outer row of a few widely spaced smaller teeth. Vomer toothless or with two rudimentary teeth. Palatines each with a long row of curved teeth decreasing in size pos- teriorly. No teeth on pterygoids or tongue. Gill rakers 2-4 at angle, minutely denticulate, otherwise represented only by short, smooth spines. Spines on inner edge of first gill arch short. No pseudobranchiae. Anus close to anal fin. Head and trunk more than twice as long as tail. Origin of dorsal fin behind middle of body length. Origin of anal fin well behind end of dorsal fin. Ventral bases slightly in front of dorsal origin. Adipose fin well developed, above middle of anal fin. ORB 2, one close to front margin of eye, the other close to its posterior margin, the anterior one a little larger. OP 3, upper one slightly below a level from top of eye; lower anterior one about level with end of maxillary and smaller than either the upper one or the lower posterior one; the latter placed a little higher than the anterior lower organ. SO present, large. BR 14. No additional photo- phores on head. Body with two rows of photophores; photo- phores present on isthmus. IV 8 + 3 + 14 = 25, the first eight GREY: NEW GENERA OF GONOSTOMATIDS 175 straight, the next three smaller and in an ascending cross line, the ninth being slightly below the eighth; and the last fourteen straight and on a level with the ninth (thus slightly lower than the first eight). VA 11-12. AC 12, straight, five or six of them behind anal fin. IC 48-49. OA 29-31, extending to, or nearly to end of anal fin. No additional photophores and no luminous tissue on body as far as known. Fin rays: dorsal 12, anal 14, pectoral 9-10, ventral 7-8. Branchiostegal rays 17, no spines at bases. Remarks. Woodsia is closely related to Photichthys Hutton and, more remotely, to Vineiguerria Jordan and Evermann and Ichthyococcus Bonaparte. The structure of the photophores is similar in all of these genera and they also share such characters as two suborbital photophores, similar positions of the dorsal and anal fins, and a short tail. Photichthys and Woodsia are alike in their dentition, in having a longer premaxillary than other related genera, and a larger number of branchiostegal rays and BR photophores. The new genus differs from all gonostoma- tids in having the gill rakers reduced in number, and from Photichthys it differs further in the shorter anal fin, the deeper body form, and lower photophore counts. The new genus has been named for Mr. Loren P. Woods, Curator of Fishes, Chicago Natural History Museum. WoopstA NONSUCHAE (Beebe) Figure 1. Photichthys nonsuchae Beebe, 1932, Zoologica, 13:61, fig. 11; 1937, op. Cut:; 22:201. Measurements and other information about the holotype of this species, U.S.N.M. No. 170938, have been obtained through the kindness of Dr. Giles W. Mead; and to Dr. Alfred W. Ebeling and Dr. Carl L. Hubbs I owe the opportunity of studying a specimen from the Gulf of Panama, Scripps Institution of Oceanography No. S1O-55-244, standard length 76.5 mm. It was taken in 730-0 fathoms (1335-0 meters), 14 November 1955, 4° 04’ N., 78° 43’ W. The following description is based on the Pacific specimen. Dorsal rays 12, anal rays 14, pectoral rays 10, ventral rays 8, branchiostegal rays 17. MUSEUM OF COMPARATIVE ZOOLOGY BULLETIN 176 VIUBUBG JO Jimny “uw Gg, YYSuoT,T prvpueys FHRZ-CG-OLG “ON sydeasourdg Jo wornqysuy Sddig “(eqoogq) apyonsuou pispoo 4, “TT *BIgL MaN4diad GREY: NEW GENERA OF GONOSTOMATIDS 17 a Measurements in millimeters: depth 15.5; head ca. 23; snout ea. 5.5; orbit ca. 7.5; interorbital width at center of eye ea. 5; upper jaw 16; premaxillary 7.5; toothed portion of maxillary 8.5; lower jaw 17; tip of snout to dorsal origin 43.5, to anal origin ca. 56, to ventral base 39.5; distance between anal origin and base of middle caudal rays 20.5, last anal ray and base of middle caudal rays 11, last dorsal ray and base of middle caudal rays 26.5, last dorsal ray and adipose fin 11, ventral base and anal origin 17; least depth of caudal peduncle 7; dorsal base 8; anal base 10.5; peetorals and ventrals broken. Seales and most of skin lost. Bones of head fragile, jaws and snout somewhat damaged. Gill rakers on first arch developed only at angle, two on left side, three on right side; otherwise represented by 3-4 very short, smooth spines on upper limb and 11 clusters of two or three little spines on lower limb; total count of gill rakers and rudiments 13 + 4 on left side, 13 + 3 on right side. Premaxillary with a single row of six slender, widely spaced teeth, curving slightly inward, the first and last of these smaller ; sometimes a few minute teeth in interspaces. Maxillary with a single row of about ten smaller teeth, and a few minute teeth in each interspace. Lower jaw with a series of seven to nine widely spaced slender teeth, most of them as long as the longest pre- maxillary teeth; a few sparse, smaller interspace teeth; and anteriorly only, an outer row of four widely spaced teeth, smaller than those of inner row. Vomer toothless. Each palatine with a row of about eleven widely spaced, curved teeth, the posterior ones minute. Pterygoids and tongue without teeth. Photophores conspicuous, many of them **double,’’ the upper black-sheathed portion enlarged and, in some cases (especially on the head) apparently with an exposed luminous area; this double aspeet particularly noticeable in the ORB, SO, BR, OP, AC, OA and the pre-pectoral I1V. BR 14. IV 8+3-+ 14 = 20. VA 11. AC 12, five of them behind anal fin, the last three smaller and slightly separated from the rest. OA 29, the last four quite small, ending above the twelfth anal ray. Myomere count probably about 45. A series of pores along lateral line area anteriorly; and on sides from behind head to anal origin smaller pores, which follow 178 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY the lines of the septa between the muscle bands, both above and below the lateral line. Other characters as in generic diagnosis. Color probably brown or black, judging from remains of skin. Linings of mouth and gill covers dusky. Adipose fin blackish. Remarks. The newly discovered Pacific specimen differs re- markably little from the holotype, which was caught off Bermuda at a depth of 1097 meters. Dr. Giles W. Mead has examined the latter, and percentages based on his measurements are shown in the table. The most important correction of the original deserip- tion is the length of the premaxillary, which, in the figure of the type, was shown to be less than half as long as the toothed portion of the maxillary. Actually it extends to a vertical from the anterior edge of the pupil and bears eight longer teeth (on each side). Dr. Mead has also confirmed the presence of the SO and upper OP photophores. As shown in the table, the Pacific specimen differs in counts from the holotype only in having one more ventral ray, one more pectoral ray, one less VA and two less OA photophores. The two specimens are almost identical proportionately with one notable exception, 1.e., the lengths of trunk and tail. The Atlantic fish has a longer trunk and shorter tail, the insertion of the anal fin being farther behind the end of the dorsal fin than in the Pacific specimen, and the distance between the ventral bases and the anal origin is also a little longer. If future material should prove this difference to be invariable, the Pacific form would probably require a new spe- cifie name, but in view of the fact that the two specimens are otherwise so similar, they are for the present considered to belong to a single species, nonsuchae. Table. Counts and measurements of Woodsia nonsuchae taken from the literature where indicated. Beebe, 1932, p. 61 U.S.N.M. SIO-55-244 170938 (Mead, 1958, in litt.) Standard length 89 ca. 85.4 76.5 Dorsal rays 12 2 12 Anal rays 14 14 14 Pectoral rays 9 9 10 Ventral rays 7 7 8 GREY: NEW GENERA OF GONOSTOMATIDS Beebe, 1932, p. 61 U.S.N.M. 170938 (Mead, 1958, in Litt.) Branchiostegal rays — 16? Gill rakers on first arch 4 — Total number of gill 13-+5 = rakers and rudiments ORB 2 2 OP -— 3 SO = ot BR 14 — IV 11+14=24 — VA 12 — AC 12) — IC 49 -— OA 31 —- Long teeth in upper jaw 19 — Long teeth in lower jaw 10 -— Vomerine teeth 2 rudiments os Palatine teeth ial — per cent of standard length Depth 20.2 20.5 Head 29.8 30.4 Snout fis} 6.41 Orbit 9.0 10.5 Interorbital width at cen- 5.05 7.6 ter of eye Upper jaw — 21.0 Premaxillary — ca. 9.36 Toothed portion of max- — 12.3 illary Lower jaw 20.8 ca. 21.6 Tip of snout to: dorsal origin -—- ca. 57.4 anal origin — a ventral base — 51.5 Distance between: anal origin and caudal — 25e5 base end of anal and caudal — 13.45 base 1 Snout now in poor condition. 179 S1O-55-244 13-¢:3-4 14 813414=25 11 12 48 29 ca. 73. 180 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY Beebe, 1932, p. 61 U.S.N.M. SIO-55-244 170938 (Mead, 1958, in litt) per cent of standard length end of dorsal and — 33.4 34.6 caudal base end of dorsal and adi- — 13.45 14.4 pose ventral base and anal = 25.8 22.2 origin Least depth of caudal 8.43 8.9 9.15 peduncle Dorsal base — 9.37 10.45 Anal base aa. 14.0 LSAT Al + SONODA, new genus Type species. Sonoda megalophthalma, new species. Generic characters. Kye normal, very large. Snout shorter than orbit. Interorbital width at center of eye much less than diameter of orbit. Mouth moderate, almost vertical; edge of premaxillary slightly concave or nearly straight; toothed edge of maxillary convex, not quite reaching a vertical from posterior margin of pupil. Premaxillary shorter than toothed portion of maxillary but more than half its length. Angle of preopercle vertical. Teeth minute, uniserial in upper jaw, in two or more rows anteriorly in lower jaw. Vomer with three or four small teeth on each side. Palatines with a single row of minute teeth. Pterygoids and tongue probably toothless. Gill rakers 15-18 + 3 on first arch. No spines on inner edge of first gill arch except sometimes a single minute one on upper limb; second arch with one or two clusters of minute spines. Pseudobranchiae sometimes present, small. Seales present, deciduous. Head and trunk slightly shorter than tail. Anus close to ventral bases. Dorsal origin slightly nearer snout than base of caudal. Anal in two sections, connected by membrane, its origin beneath or slightly in advanee of first dorsal ray. Ventral bases well ahead of dorsal origin. No adipose fin. ORB 1, in front of lower portion of eye. OP 38, upper one close to eye and level with middle of eye; lower anterior one at end of maxillary; lower posterior one vertically GREY: NEW GENERA OF GONOSTOMATIDS 181 elongate, in a black sheath with luminous areas showing above and below (probably double). SO absent. BR usually (6), (7) on one side of one specimen. Body with two rows of serial photo- phores; photophores present on isthmus. 1V (6) + (10) ; the first six larger; the fourth, fifth and sixth curving upward toward pectoral base; the last ten close together and separated from those on the other side only by two black bands on belly. VA (7) or (8), joined above to a common gland. AC in two long eroups, (16-21) + (19-24) — 36-43, the organs small, sepa- rated below but joined above to narrow glands; the first group commencing over seventh or eighth anal ray, second group com- mencing above last anal ray. IC 59-67. OA (2) + 4-5 = 6-7, the first two joined above to a gland, others separate. No ad- ditional photophores or patches of luminous tissue on head or body as far as known. Fin rays: dorsal 8-9; anal 8-9 + 14-16 = 22-25, the first eight or nine rays thicker than the rest and separated from them by a short but definite space, although con- nected by a relatively thick membrane ; pectoral 13-15; ventral 6. Branchiostegal rays 8, no spines at bases. Vertebrae 40 including hypural, counted on an X-ray photograph of one specimen. Remarks. Sonoda is in some respects intermediate between Maurolicus Coeeo and Valenciennellus Jordan and Evermann. It is similar to the latter in the relative positions of the dorsal, anal and ventral fins, in having the trunk shorter than the tail, and in lacking the SO. In the arrangement of the AC photo- phores the new genus is closer to Maurolicus and there is at least a superficial similarity in the shape of the anal fin in these two genera also. However, in Maurolicus the anal fin does not seem to be actually in two separated sections as it is in Sonoda. The lower posterior OP, and some of the IC photophores, are similar in structure to those of Argyripnus Gilbert and Cramer. In having 7-8 VA photophores Sonoda differs from all other known maurolicid genera, in which these number only 4-6. The new genus is named for Miss Pearl Sonoda, Assistant in the Division of Fishes, Chicago Natural History Museum. SoNODA MEGALOPHTHALMA, new species Figures 2, 3 Holotype. Standard length 58 mm., Oregon Station 1918, 13° 25’ N., 82° 01’ W., 12 September 1957, 300 fathoms (548 meters). 182 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY Paratypes. Two specimens, standard lengths 54-55 mm., taken with the holotype. One specimen, 43.5 mm., Oregon Station 1924, 12° 50’ N., 82° 12’ W., 13 September 1957, 275 fathoms (503 meters). Four specimens, ¢a. 50-60 mm., Oregon Station 1928, 13° 14’ N., 82° 06’ W., 12 September 1957, 300 fathoms (548 meters). Five specimens, 44.5-52.5 mm., Oregon Station 1931, 13° 56’ N., 81° 50’ W., 14 September 1957, 275 fathoms (503 meters). Eleven speeimens, 48.5-58 mm., Oregon Station 1933, 14° 08’ N., 81° 49’ W., 14 September 1957, 300 fathoms (548 meters). Fig. 2. Sonoda megalophthalma n.sp. Holotype, standard length 58 mm., Oregon Station 1918, Caribbean Sea. Description. Dorsal rays 8-9 (holotype, 9); anal rays 8-9 + 14-16 = 22-25 (holotype 9+ 15 = 24); pectoral rays 18-15 (holotype 14) ; ventral rays 6; branchiostegal rays 8; gill rakers on first arch 15-18 + 3, counted on four specimens (holotype 16 + 3); pyloric caecae 6; vertebrae 40, including hypural, counted on holotype only. Measurements of the holotype in millimeters: standard length 58, greatest depth 12, length of head 14.5, snout 3, diameter of orbit 6.5, interorbital width at center of eye ca. 2, upper jaw ea. 9. Tip of snout to dorsal origin 26, to anal origin 25.5, to ventral bases ca. 20. Distance between anal origin and base of middle caudal rays 32.5, last anal ray and base of middle caudal rays 10, last dorsal ray and base of middle caudal rays 24. Least depth of caudal peduncle 5. Dorsal base 6. Anal base 23. Length of pectoral fin 11, of ventral fin ca. 5. GREY: NEW GENERA OF GONOSTOMATIDS 183 Measurements expressed in per cent of standard length, of nineteen specimens, standard length 43.5-60 mm.: depth 19.0- 21.8; head ca. 23.4-25.7 (and one, 27.3); snout 4.5-6.36; orbit 10.3-12.1; interorbital width at center of eye 2.19-3.54; upper jaw 15.0-16.4; tip of snout to dorsal origin 43.0-47.7, to anal origin 41.0-46.6, to ventral bases ca. 32.4-38.2; distance between first anal ray and base of middle caudal rays 54.0-57.6, last anal ray and base of middle caudal rays 15.2-19.1, last dorsal ray and base of middle caudal rays 37.1-45.4; least depth of caudal pedunele ca. 7.0-9.25; dorsal base 8.6-11.0; anal base ca. 36.4- 41.4; pectoral length 17.0-24.1; ventral length ca. 6.9-11.3. Se => ae, Ee Nic) © 0 soe OSes. Caso" G = aCIsGiorais ono a ad SS SCog06 ° Zo OSD SE OSS SoD 4 ZA (SSS OOO Ro (ere x®) Le Be = Fig. 3. Sonoda megalophthalma u.sp. Diagramimatie ventral view of an- . eS terior portion of body, showing position of light organs. Bones and skin of head very fragile, nostrils not apparent ex- cept on one specimen, in which they appear to be placed high on snout as in other gonostomatid fishes. Pseudobranchiae small, found on only a few specimens, perhaps lost in others because of their fragility. Skin of abdomen thin, often damaged. A large, thin, cycloid scale present beneath pectoral of one speci- men from St. 1933, no scale pockets visible, extent of scalation unknown. Fin rays fragile, mostly broken, probably normally all rather long, judging from remnants. Pectoral in some specimens reaching first or second anal ray. Ventrals, when complete, reaching anal origin. Photophores of BR, IV and OA series relatively large, the VA, and especially the AC, smaller. Total number of IV organs always 16, those of BR 6 except in one specimen, which has 6 on one side, 7 on the other. Number of VA and AC organs variable, the former 7 in twelve specimens, 8 in eight specimens and 7/8 in two specimens; those of AC often differing on the two sides of a single specimen. OA variable but usually 6. Fin rays also 184 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY varying in number: dorsal 9 in nineteen specimens, 8 in four specimens; first group of anal rays 9 in twenty specimens, 8 in two specimens; second group of anal rays 15 in ten specimens, 16 in seven specimens, 14 in one specimen; total number of anal rays 24 in ten specimens, 25 in six specimens, 22 in one speci- men; pectoral rays 14 in twelve specimens, 15 in three speci- mens, and 13 in two specimens; ventral rays always 6. Color in alcohol: body largely pale yellowish or whitish, sprinkled with black chromatophores ; abdomen black, sometimes with bluish or greenish iridescence anteriorly. Operecles and lower part of head black. Snout, lower jaw and top of head colorless and transparent. A yellowish body visible beneath top of head between and behind eyes. Fins colorless. Inside of mouth colorless. Linings of gill chamber and peritoneum black. At least five specimens are mature females. Characters otherwise as in generic diagnosis. LITERATURE CITED BrrBeE, W. 1932. Nineteen new species and four post-larval deep-sea fish. Zoologica, 13:47-107, figs. 8-31. 1937. Preliminary list of Bermuda deep-sea fish. Zoologica, 22:197- 208. Grey, M. 1955. Notes on a collection of Bermuda deep sea fishes. Fieldiana, Zool., 37:265-302, figs. 45-56. MaRSHALL, N. B. 1951. Bathypelagic fishes as sound scatterers in the ocean. Jour, Mar. Res., 10:1-17. 1954. Aspects of deep sea biology. London. 380 pp., 4pls., text figs. Bulletin of the Museum of Comparative Zoology A VAC VAL Di COW Le GE Vol. 121, No. 5 THE ANOLES OF THE EASTERN CARIBBEAN (SAURIA, IGUANIDAE) Parts IED By JULIAN S. KENNY Victor C. QUESNEL GARTH UNDERWOOD Ernest KE. WILLIAMS Wits ONE PLATE CAMBRIDGE, MASS., U.S.A. PRINTED FOR THE MUSEUM Juny, 1959 PUBLICATIONS ISSUED BY OR IN CONNECTION WITH THE MUSEUM OF COMPARATIVE ZOOLOGY AT HARVARD COLLEGE BULLETIN (octavo) 1863 — The current volume is Vol. 121. BREVIORA (octavo) 1952 — No. 111 is current. Memoirs (quarto) 1864-19388 — Publheation was terminated with Vol. 55. JOHNSONIA (quarto) 1941 —— A publication of the Department of Mollusks. Vol. 3, no. 38 is current. OccASIONAL PAPERS OF THE DEPARTMENT OF MOLLUSKS (octavo) 1945 — Vol. 2, no. 22 is current. PROCEEDINGS OF THE NEW ENGLAND ZooLogicaAL CuuB (octavo) 1899-1948 — Published in connection with the Museum. Publication terminated with Vol. 24. The continuing publications are issued at irregular intervals in num- bers which may be purchased separately. Prices and lists may be obtained on application to the Director of the Museum of Comparative Zoology, Cambridge 38, Massachusetts. Of the Peters ‘‘Cheek List of Birds of the World,’’ volumes 1-3 are out of print; volumes 4 and 6 may be obtained from the Harvard Uni- versity Press; volumes 5 and 7 are sold by the Museum, and future volumes will be published under Museum auspices. Bulletin of the Museum of Comparative Zoology AT HARVARD COLLEGE Volo 121. Nor 5 THE ANOLES OF THE EASTERN CARIBBEAN (SAURIA, IGUANIDAE) Parts [-IT] By JULIAN S. KENNY Victor C. QUESNEL GARTIL UNDERWOOD Ernest KE. WrinuiAMs Wir One Puate CAMBRIDGE, MASS., U.S.A. Pen DS HOU: TE Mi wiSaa UeM JuLY, 1959 No. 5 — The Anoles of the Eastern Caribbean (Sauria, Iguanidae). Parts I-I11 TABLE OF CONTENTS Page i. Pretace: “By ‘Hrnest He Wilhams ya. 2225. 2 187 Il. Two Sibling Species of Anoles in Trinidad. By Victor C.'Quesnel and Julian 8. Kenny .......... 189 III. Revisionary Notes. By Garth Underwood ........ IED Part I. Preface By Ernest E. WILLIAMS The papers that follow have resulted from an attempt, some years ago, to survey what was felt to be the simplest geographic segment of the genus Anolis. This study was intended as the first part of a projected general review of this very large and complex genus. The Lesser Antillean islands (defined as the islands from Anguilla to Grenada) had, so far as was definitely known, no more than two species of anoles on any one island, and a pre- ‘liminary survey of material in the Museum of Comparative Zoology seemed to indicate that no more than five full species were involved, all rather similar but still differing in clear and simple morphological details one from another. It seemed also from this survey, as carried out by myself (and also independ- ently for his own information by my co-worker Benjamin Shreve), that no major taxonomic problems existed and that the analysis at this level could be carried to completion simply and routinely. However, it had been my experience while studying the anoles of Jamaica, in collaboration with Garth Underwood, that colors in life enormously assisted in the recognition of forms that morphologically were often disconcertingly close. I therefore asked Garth Underwood, who was visiting the Lesser Antilles in 188 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 1956 for other purposes, to send me a few live anoles from each of the islands at which he stopped. This he faithfully did. The material that he collected very soon destroyed the pretty simplicity that had been fancied to exist; the problems that he presented us with are still only partially solved, although Garth Underwood himself has since studied the types of Lesser An- tillean Anolis in the Paris Museum, in the British Museum, and in the United States National Museum, and although he and I and Benjamin Shreve have restudied all the Museum of Com- parative Zoology specimens, increased as they have been by Underwood’s own collections. Prominent in the new emerging picture are certain new or rediscovered ‘‘sibling’’ forms. A supposedly unitary series of forms on the southern Lesser Antilles resolves itself on closer inspection into geographically replacing forms of at least two species somewhat erratically distributed over these islands. The species we here eall ‘‘sibling’’ can, it is discovered by close examination, be recognized and defined on conventional morphological characters. However, it is very probable that the rather subtle scale characters that distinguish them would have been dismissed — in the past have been dismissed — as ‘‘ varia- tion’’ had these species not first been recognized by color or behavior or both. It will be recalled that the segment of Anolis that is here re- ported on was believed to be, and may well be, the simplest areal segment of the genus in which more than one species occurs. (Only the portion of the southeastern United States in which A. carolinensis is the only member of the genus can be simpler.) Yet we have found the complications formidable and not amen- able to study on the basis of preserved specimens alone. It is even more evident in this genus than in others that we cannot be prisoners of our conventional museum techniques, that a very wide approach will barely encompass the things we need to know. Knowledge of the live animals is needed — of captives, if necessary — but preferably field knowledge. Anolis must be known intimately — anatomically, ecologically, ethologically — in all its relationships within and between species, if the many puzzles the genus poses are to be solved. How remote from this ideal our present knowledge is, anyone with even a little familiarity with the genus is aware. Many ANOLES OF THE EASTERN CARIBBEAN 189 forms are known from single specimens — rarely with any notes on color in life, much less on habits and habitat. Even the West Indian anoles, which often swarm wherever they are to be found, are extremely poorly documented ecologically and ethologically. It is thus no accident that the study of which we here publish a first portion has been a cooperative effort. It has required spe- cial opportunities, much travel, and a great deal of work to pro- duce what is transparently a bare beginning. One new form is described here by Underwood. Some evidence is already at hand that other new forms require to be described. Further parts of this study will deal with these new forms and with other data still accumulating. Part Il. Two Sibling Species of Anoles in Trinidad fo} By Victor C. QUESNEL AND JULIAN S. KENNY Two species of Anolis lizards have been recorded from Trini- dad by Parker, 1935: Anolis chrysolepis and Anolis aeneus. In 1955 it came to our notice that there was an apparently different, third, species which had not been recorded. A specimen was sent to the British Museum (Nat. Hist.) where it was identified as Anolis aeneus. This specimen, when alive, was predominantly green in color, whereas the lizard that we knew as A. aeneus is predominantly grey. A specimen of the grey form sent with the ereen one to the British Museum (Nat. Hist.) was also identified as A. aeneus. We believe that these two forms of A. aeneus may be, in fact, distinct species, and we give below our observations bearing on this point. The lizards are similar in squamation and limb measurements, and preserved specimens tend in time to assume the same color. Yet, in the field, the grey form appears the larger and more heavily-built and there is a suggestion of slight differences in the shape of the head. From the measurements we have so far been able to make, we would say that eggs and newly-hatched young are similar in size. Thus, seven eggs of the grey form ranged from 9.5 x 6.9 mm to 11.0 x 9.0 mm, and two eggs of the green form measured 10.7 x 5.8 mm and 10.3 x 8.0 mm. The eggs increase in size during incubation so that large numbers of eggs of known ages would be necessary for the detection of any differences there may be. 190 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY Similarly, four newly-emerged young of the grey form had the following measurements (snout to vent + tail): 20.5 + 36.5 mm; 21.8 + 40.0 mm; 20.7 + 37.3 mm; 20.5 + 35.5 mm; and one of the green form measured 19.5 + 33.0 mm. In spite of these general similarities we believe that the two forms are really distinct species. The differences in coloration and pattern of the living animals are striking. The grey form has a pattern of forward-pointing V-shaped cross bands, usually brown in color, and is frequently speckled, mottled or marbled. There is a great deal of variation between individuals and in the same individual at different times. The green form shows hardly any sign of V-shaped markings. The general body color is green- gold, rather paler in the mid-dorsal line than dorso-laterally, fading to a pure lemon yellow on the underparts. The head is brilliant green and the area surrounding the eye bright blue. This lizard can change its color to rich red-brown, and this is the color of the newly-hatched young. The dark coloration which the grey form sometimes assumes is also brown, but without the reddish tinge. There is a marked behavioral difference which we have con- firmed many times. The green form will frequently enter houses and make itself at home there; the grey form will not. In almost four years of living in a house whose garden is overrun with the erey Anolis, V. C. Quesnel has never yet found one in the house. On the other hand, we have had several reports of lizards enter- ing houses and on each occasion careful questioning and a check of the garden showed that the lizard concerned was the green Anolis. One informant related how the lizards lived in and around the furniture and took crumbs from the floor near the dining table. The specimen of the green form which was sent to the British Museum (Nat. Hist.) was caught inside a house. On one occasion some lizards of both forms escaped from a cage in the home of J. S. Kenny and the next day one or two of the green form were recaptured in the house; the grey ones had disappeared. So far, we have found the grey form in Port of Spain, Mayaro, and St. Augustine and the green form in Port of Spain, St. Augustine, San Fernando and Usine, Ste. Madeleine, but in districts where they both occur they are seldom found together. We have found only two mixed populations, and in these one ANOLES OF THE EASTERN CARIBBEAN 191 glance is usually enough to classify an individual as grey or green. Only one large male in one of these populations seemed to be intermediate in color. Their habitat preferences appear to be similar. In Port of Spain both abound in hedges and low shrubs. This is in contrast with Beebe’s observations of the grey form in British Guiana. (It is clear from his description that it is the grey form he observed.) Beebe (1944) says: ‘‘The Marbled Anolis is a tree-trunk lizard like Plica, rather than a frequenter of low, leafy shrubs.’’ Perhaps it is the near identity of ecological niche that accounts for the usual absence of mixture of populations of the two forms in Trinidad; or perhaps there are differences in preferred habitats that we have not detected. These observations show the necessity for a thorough study of the anatomy and behavior of the two forms in order to substan- tiate or reject our view that they are distinct species. REFERENCES BEEBE, W. 1944. Field Notes on the lizards of Kartabo, British Guiana and Caripito, Venezuela. Part 2. Iguanidae. Zoologica, 29: 195-216. PARKER, H. W. 1935. The lizards of Trinidad. Tropical Agriculture, 12: 65-70. Part III. Revisionary Notes By GartH UNDERWOOD Department of Zoology, University College of the West Indies I became involved in the problems of the anoles of the eastern Caribbean through a combination of circumstances. In the sum- mer of 1956 I visited Trinidad and Tobago travelling on a ship which called at St. Kitts, Antigua, Montserrat, Dominica, St. Lucia, Barbados, St. Vineent and Grenada. On each of these islands I collected anoles and sent living material to the Museum of Comparative Zoology to assist the studies of Dr. E. E. Wil- hams and Mr. B. Shreve. What was intended to be purely routine collecting revealed problems which were not apparent from the study of preserved material. The following summer I travelled on study leave and was able to visit the Museum d’Histoire Naturelle in Paris (M.H.N.P.), the British Museum 192 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY of Natural History (B.M.N.H.), the Museum of Comparative Zoology (M.C.Z.) and the United States National Museum (U.S.N.M.). I have thus seen living material from ten different islands, preserved material of all the forms and all of the type specimens, save three. For comparison with the types in Europe Dr. Williams sent me specimens from the French islands of Guadeloupe, Désirade, Marie Galante and Martinique. As I had made my collections, examined material and arrived at conclu- sions before having an opportunity to consult personally with the other two interested parties it seemed better that I should assume personal responsibility for my findings. Messrs. Williams and Shreve, however, very generously went further and invited me to review all the pertinent material in the M.C.Z. What began as an innocent offer of help has ended as an appropriation of their research project. I publish my findings in their present incomplete form because some corrections to Barbour’s arrange- ment are clearly necessary, and because it will certainly be some years before I or other interested parties will be able to ac- complish all the field work which is still required, and in order to point out the interest of the situation to other students. The taxonomic history of these anoles is of some interest. There appear to be 36 names of which 20 at present appear to be valid. Much of the older authors’ material is without data or, worse, with erroneous data. It is therefore evidence of Cope’s perspicacity that he correctly identified more of the earlier de- scribed species than any subsequent worker, and that his four new names still stand. Garman and Barbour worked with ma- terial of known provenance; however, they misidentified a num- ber of the older species. Consequently some of their names are synonyms of older names. From the observation that many of the islands have endemic forms they seem to have proceeded in the direction of the assumption that all the islands have endemic forms; some of their new species are in my opinion not signifi- cantly different from already named forms — a conclusion at which I may say that Williams and Shreve had already arrived. Of Garman’s twelve new names, at least seven appear to be good. Barbour proposed five new names, four of which appear to be synonyms; he misidentified four forms, and overlooked one un- named distinct form in the Museum of Comparative Zoology. ANOLES OF THE EASTERN CARIBBEAN 193 These lizards present the familiar nomenclatorial problem of island populations. Given a full knowledge of the relevant facts, which we certainly do not have, there remains a considerable element of personal taste in deciding how to represent them nomenclatorially. I therefore state my personal taste. If two separate island populations appear to have a common immediate origin and have closely similar habits then, although on details of coloration or squamation they may be 100 per cent distinct, I would regard them as subspecies. As I do not have the relevant information on the habits of many forms, the taxonomic rank accorded to them is to be regarded as provisional. In particular in Guadeloupe and its immediate neighbors there are a number of derivatives of the bimaculatus group. We do not know enough about their ranges and reproductive isolation to attempt a natural arrangement. I regard colors in life as of primary im- portance in the discrimination of Anolis species. Squamation differences I regard as only indirectly significant by-products of species differentiation, virtually impossible to assess without in- formation about the live animals, and in anoles often difficult to characterize in words. My account is largely confined to adult males since they are the most clearly differentiated. There appear to be two major groups of anoles in the eastern Caribbean: (1) the bimaculatus group, of Greater Antillean af- finities, extends through the Leeward group into Dominiea, and (2) the roquet group, of South American affinities, extends from the mainland through Trinidad, Tobago, Barbados and _ the Windward islands as far as Martinique. A third group is repre- sented by A. wattst in the northern Leeward Islands and St. Lucia. Happily even juvenile females are distinguishable on squamation characters. In both major groups the canthal ridge passes into a long supracilary seale; in the bimaculatus group it is followed by a series of differentiated small supraciliaries; in the roquet group there is no differentiated series of small supraciliaries. The caudal scales are arranged in whorls corresponding to the autotomy septa; in the bimaculatus group there are three or four dorsal crest scales in each tail whorl; in the roquet group there are usually five dorsal seales in each whorl; if there are less it is only as an individual variation and not as a population character. 194 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY The supraorbital scale-row continues forward of the orbits and meets the canthal ridge scale-row; in the bimaculatus group the forward continuation of the supraorbital row is well defined onto the antorbital area; in the roquet group it is ill-defined. There is a pronounced ridge on the supraorbital scale-row. In the bimaculatus group it continues onto the antorbital area at least halfway to the level of the nostrils, while in the roquet eroup it scarcely extends beyond the anterior edges of the orbits. Wedged between the canthal ridge and the supraocular secale-row Fig. 1. Sealation of the antorbital region. A, in the bimaculatus group, B, in the roquet group. are some scales decidedly larger than the granular supraoculars. In the bimaculatus group they are few in number and large; in the roquet group they are usually more numerous and smaller (see Fig. 1A and B). This difference is not always very sharp. The scales on the throat fan of the males are in well-defined rows; in the bimaculatus group the scales are somewhat rounded in outline and flat; in the roquet group they are in very well- defined rows of an elongated oval shape and swollen. In the males of the bimaculatus group there is a well-defined pair of enlarged post-anal seales; in the males of the roquet jad eo} Dt ANOLES OF THE EASTERN CARIBBEAN croup there is a pair of depressions in this position in which the scales may be somewhat differentiated but not enlarged (save in one specimen from Petit Martinique). Anolis wattsi is distinguishable from the bimaculatus group by its small size, narrower digital dilations, strongly keeled belly scales and by a double row of much enlarged vertebral scales which runs from the occiput to the mid-dorsal row of tail seales. The characters by which bimaculatus differs from roquet dis- tinguish this form as well from the roquet complex. Under each form I cite the name used by Barbour (1930a), who is the most recent previous reviser of these forms. Barbour’s later checklists alter the grouping of species and subspecies but not the allocation of names. I give for each form the snout-vent leneth of a large adult male. I add the range of the form and a list of MCZ specimens studied. For notes on the colors of the animals in life I have drawn upon the observations of Garman (1887) and Barbour (1930a), as well as on my own observations. I should add an apology to my colleagues for the incomplete- ness of my personal records for I did not regard myself as being involved in this study as a reviser until after returning from my travels. ANOLIS BIMACULATUS group Apart from the squamation characters which distinguish the bimaculatus group, a number of features of color pattern run through the group. The primary color appears to be a green which can turn to brown. Some stocks appear to be ‘‘fixed’’ in the brown condition (Saba, Anguilla bank group, Désirade). Conversely some forms are a somber green without a brown phase (Guadeloupe — alliaceus). There is a light flank stripe which ranges from conspicuous, as in the Antiguan form, to obsolete as in alliaceus from Guadeloupe. There are brown speckles which are generally small as in Antiguan animals but may be very large and dark as in those from Saba or altogether wanting as in Marie Galante specimens. On the ground color there may be light speckles, absent in the forms of northern Leeward Islands, just visible in Montserrat specimens, and forming a very conspicuous pattern as in marmoratus from Guadeloupe. It is probable that there are sympatric species and distinct subgroups within the group; study of such questions requires field work. 196 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY ANOLIS BIMACULATUS GINGIVINUS Cope Anolis gingivinus Cope, 1864, Proc. Acad. Nat. Sci. Philadelphia, 1864: 170. Type: B. M. N. H. 1946. 8, 29. 18-20. ‘‘ Anguilla rock, nr. Trinidad,’’ coll. W. J. Cooper. Anolis virgatus Garman, 1887, Bull. Essex Inst., 19:41. Type: M.C.Z. 6165, St. Barthélemy, coll. Lagois. Anolis gingivinus, Barbour. Not seen alive. From Garman’s deseription and the appearance of preserved specimens this is a grey lizard which does not have a green color phase; the markings are dark brown. A series from An- guilla has fine dark speckling which may extend from the back of the neck, down the trunk and onto the dorsal surface of the hind limbs or may be so reduced as to be almost lacking. There is subdued grey vermiculation on the anterior trunk. A light flank stripe starts over the shoulder and may extend past the elbow to reach knee level. A single specimen from St. Martin has transverse bands on the anterior trunk and tail and a dark stripe above the light flank stripe. A series from St. Barthélemy (10) shows better defined grey vermiculation, the flank stripe extending to the groin and, in two specimens, indications of transverse bands. These look like minor local population differences. Snout-vent. 57 mm. Range. Anguilla, St. Martin, St. Barthélemy. MCZ specimens examined. 6157 (Anguilla) ; 6165 (10) (St. Barthélemy ) ; 16546-595 (Anguilla) ; 20985 (St. Martin) ; 33372 (Anguilla). ANOLIS BIMACULATUS SABANUS Garman Anolis sabanus Garman, 1887, Bull. Essex Inst., 19:39. Type: M.C.Z. 6161, Saba, coll. Lagois. Anolis sabanus, Barbour. Not seen alive. This very distinct form resembles gingvinus in general color ; it has distinctive, large, very dark brown blotches; they show some tendency to arrangement in transverse bands. There is a vertebral stripe clear of markings. There is no flank stripe. The throat fan is ‘‘lemon yellow’’ (Barbour). Snout-vent. 65 mm. ANOLES OF THE EASTERN CARIBBEAN 197 Range. Saba. MCZ specimens examined. 6161 (5); 28501-26 + duplL.; 33376. ANOLIS BIMACULATUS BIMACULATUS (Sparrmann) Lacerta bimaculata Sparrmann, 1784, Nya Handl. Sven. Vet. Akad. Stock- holm, 5:169. Type: Mus. de Geer Roy. Stockholm, St. Eustatius, coll. Acrelius. Anolis edwardsi Merrem, 1820, Syst. Amph.: 45. No type, Nevis. Anolis reticulatus Gray, 1840, Ann. Mag. Nat. Hist., 5:114. Type: B.M.N.H. 1946.8.29.10, no locality, no collector. Anolis mayeri Fowler, 1918, Publ. Carnegie Inst. Washington, 252:8. Type: No. 3151 Princeton University. ‘‘ Virgin Islands,’’ coll. A. D. Brown. (Synonymy fide Williams and Shreve.) Anolis bimaculatus, Barbour. Seen alive. A series of six topotypes show a strong blue-green ground color. Creamy markings extend from the upper lip to the neigh- borhood of the earhole. The flank stripe, prominent on the anterior shoulder, may reach halfway to the groin. There may be some dark spots on the lips, sides of the head, flanks and the base of the tail but their density, form and distribution are very variable. There are vague indications of transverse mottling. A series from St. Kitts (9 males) is less blue-green and more chalky-green. This appearance is true of the living animals also. The creamy markings are more extensive on the head. The equally well marked flank stripe fades at elbow level. The dark spots may cover trunk, hind lmbs and tail or may be absent. These lizards can turn brown. ‘‘Throat, dewlap and venter lemon yellow’’ (Barbour). Nevis specimens are very similar and I agree that Anolis edwardst should be regarded as a synonym. The type of Anolis reticulatus is a female without data. The specimen is very dark without any color pattern. In squamation it shows good agreement with Antiguan specimens but some- what better agreement with St. Kitts specimens. I propose there- fore to place reticulatus as a synonym of bimaculatus. Snout-vent. 99 mm. Range. St. Eustatius, St. Kitts, and Nevis. MCZ specimens examined. 6156 (23), St. Kitts; 6184 (2), Nevis; 10379-402, St. Kitts; 16166, St. Kitts; 16169, Nevis; 198 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 16208, Nevis; 16278-95, St. Kitts; 16529-31, 16533-5, 16537, 16539, St. Eustatius; 28714-21, Nevis; 28722-4, Basseterre, St. Kitts ; 88277-8, 6 mi. E. of Charleston, Nevis; 54718-19, St. Eusta- tius. ANOLIS BIMACULATUS LEACHT Duméril and Bibron Anolis leachw Duméril and Bibron, 1837, Erp. Gen., 4:153. Type: M.H.N.P. Ig 34/1; ‘‘Antilles’’; pres. T. Bell. Anolis antiquae Barbour, 1915, Proe. Biol. Soe. Washington, 28:74. Type: M.C.Z. 10624; Antigua; coll. Wulsin and Noble. Anolis barbudensis Barbour, 1923, Oee. Pap. Mus. Zool. Univ. Mich., 132:4. Type: M.C.Z. 16167; Barbuda; coll. Forrest. Seen alive. The type specimen in Paris is an adult male with a well preserved color pattern. It bears a detailed resemblance to Antiguan specimens. There is speckling from the occiput to the loins, a flank stripe as far as the elbow and dark longitudinal speckling beneath the head. The specimen bears an old label Anolis bimaculatus. I suppose that it was initially identified as this species. This lizard is yellow-green in life with partly confluent brown speckles. There is more confluence of speckles on the anterior trunk, producing a coarsely vermiculate pattern, than on the posterior trunk which has independent speckles. The speckling may fade at the loins or may reach the tail. The flank stripe fades by elbow level. The lizard can turn brown. The throat fan is yellow. The types of Anolis barbudensis are two males. The larger has greatly reduced speckling. The smaller is typically Antiguan in pattern. I regard barbudensis as a synonym of leachi. In the London Zoo Mr. R. Lanworn kindly showed me a col- lection of anoles from Bermuda. Some were the Jamaican species Anolis g. grahami which I have myself encountered in Bermuda. The others were quite clearly the Antiguan species. They were correctly identified under the name Anolis leachi. Snout-vent. 89 mm. Range. Antigua, Barbuda. MCZ specimens examined. 10624-5, St. Johns, Antigua; 11981- 4, Antigua; 16167-8, Barbuda; 55455, Boggy Peak, Antigua ; 55456-7, Kensington House, St. John’s, Antigua; 55458-9, nr. Crab Hill Village, Antigua ; 55677-81, 55773, St. John’s Antigua. ANOLES OF THE EASTERN CARIBBEAN 199 ANOLIS BIMACULATUS LIVIDUS Garman Anolis lividus Garman, 1887, Bull. Essex Inst., 19:43. Type: M.C.Z. 6176; Montserrat; coll. Garman. Anolis lividus, Barbour. Seen alive. The predominant color is bright yellow-green grading to blue on the tail. There is russet around the eye and a variable exten- sion of russet onto the head and forequarters. Oblique rows of pale spots were sometimes present on the sides. The belly is yellow. The flank stripe is never strongly defined and is very variable in its extent. These lizards can turn a warm dark brown. The throat fan is light ochre. Snout-vent. 70 mm. Range. Montserrat. MCZ specimens examined. 6176 (35), 38379 (9), 55686-705, 559837, all from Plymouth, Montserrat. ANOLIS BIMACULATUS ?LIVIDUS Garman Anolis nubilus Garman, 1887, Bull. Essex Inst., 19:32. Type: M.C.Z. 6181; Redonda; coll. Richardson. Anolis nubilus, Barbour. Not seen alive. The larger male type of nubilus is larger than lividus. There is not the same clear indication of a green color phase. The light speckling is pronounced on the hindlimbs where the speckles are fairly large. There is almost no trace of the flank stripe. The distinctness of this population cannot be judged from three pre- served specimens. I propose that it be placed under lividus until a larger series can be compared directly with living material. Snout-vent. 80 mm. Range. Redonda. MCZ specimens examined. 6181 (3), Redonda. ANOLIS BIMACULATUS ALLIACEUS Cope Anolis alliaceus Cope, 1864, Proc. Acad. Nat. Sci. Philadelphia, 175. Type: B.M.N.H. 1946.8.28.96; no locality; purchased of M. Parduzaki. Anolis leachii, Barbour. Not seen alive. 200 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY The type is a fairly well preserved adult male which shows some pattern. It is quite clearly an example of the species com- mon in Guadeloupe. Above each shoulder is a light-edged hori- zontal black patch (divided on the left). Further forwards on the side of the neck are smaller similar marks; on the posterior head and anterior trunk are dark vermiculations. Most of the preserved specimens are very dark but it appears that they have a green color phase. Most have black light-edged blotches on the anterior trunk. In some specimens they may be partly joined to form transverse markings. There is commonly a well-defined horizontal black patch over the shoulder. There is no flank stripe. There may be indications of vermiculations as well as black markings on the anterior trunk. At the time of going to press I learned from Dr. Williams that he is satisfied that the series of specimens from St. Rose, Guade- loupe, which I have regarded as alliaceus, represent an unde- seribed species. One of the characters is uniform dark coloration without any black markings. Fortunately this character raises no question about the identity of alliaceus, under which name the St. Claude series will remain. It does, however, raise a question about the type of reticulatus. Snout-vent. 71 mm. Range. Guadeloupe, Basse-Terre only ? MCZ specimens eramined. 10314-3863, St. Claude, Guadeloupe ; 10403-425, St. Rose, Guadeloupe. ANOLIS MARMORATUS Duméril and Bibron Anolis marmoratus Duméril and Bibron, 1837, Erp. Gen., 4:139. Type: M.H.N.P. Ig. 43, paratype Ig 43/1; ‘‘Martinique’’; coll. Plée. Not seen alive. The two original specimens are adult males; the bellies are rather soft but the pattern is well preserved. As soon as I saw these specimens I was sure that I had not seen the species before and equally sure that I would recognize it when next I saw it. In particular, the color pattern does not resemble that of any of the specimens from Désirade in the M.C.Z. The species be- longs to the bimaculatus group and shows some resemblance in color pattern to the form in Dominica. [ was initially inclined to accept the Martinique locality as correct. As matters stand, only one representative of the roquet ANOLES OF THE BASTERN CARIBBEAN 201 group is known from Martinique; the occurrence there of a representative of the bimaculatus group, bearing some re- semblance to the Dominican form, seemed entirely plausible. I regarded Martinique as the meeting ground between Antillean and South American stocks. However, in the U.S.N.M., I found five specimens of marmoratus, all collected by L. Guesde, as fol- lows: 11174, ¢ ; 11180, ¢ — Guadeloupe; 11183, 2¢ ; 11184, 2 — Capesterre, Guadeloupe. A number of Plée localities are known to be erroneous. Is this Guesde locality more reliable? The addition af an exact locality (Capesterre is in S.E. Basse- Terre) to one of the collections lends authenticity. That Guesde collected in Guadeloupe and dependencies is clear from other material in the museum: There are specimens of ferreus from Marie Galante. Two males, USNM 11181, from ‘‘St. Barthélemy and St. Martin’’ are certainly not gingivinus and closely re- semble the Désirade form. This however still does not take us outside the political limits of Guadeloupe. (It is relevant that Louis Guesde was a resident of Guadeloupe well known locally for his interest in Amerindian remains.) Further the U.S.N.M. does not have anoles from Martinique collected by Guesde. I therefore assume that marmoratus represents a second Guade- loupe species. The forelimbs, hindimbs and posterior trunk are uniform and, as preserved, dark chocolate brown. On this dark ground color the head and dorsal neck have a very striking pattern of light speckles, partly confluent and more or less organized into stripes with irregular borders. A light stripe runs from the posterior part of the canthal ridge over the supraciliary seales, expands on the supraocular region to isolate two dark spots on the supra- ocular disc, passes across the temple and curves in towards the midline (save in 11174) behind the parietal region. Another light stripe follows each supraorbital scale row. There is a group of light speckles on the antorbital depression and a group of partially confluent speckles on each parietal area. A median light stripe runs along the nuchal crest (interrupted in smaller 11183). A light stripe along the upper lip, interrupted by dark speckles, passes into a row of light speckles running from the corner of the mouth across the earhole and onto the side of the neck where it becomes the middle member of three roughly or- ganised rows of light speckles. 202 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY The light speckles on the neck extend down the sides to about the shoulder level; they do not extend backwards as far as the shoulders. A light flank stripe extends back from over the shoulder and fades out before reaching elbow level (absent in 11174). There may be a few dark speckles on the lower lip and on the underside of the head. The single female is without evi- dent markings. In the U.S.N.M. register the Capesterre Guesde collection bears in the remarks column the note ‘‘ Anolis téte-rouge.’’ Guesde was presumably either reporting a local name or giving an indication of the appearance of this form. Snout-vent. 67 mm. Range. Guadeloupe, Basse-Terre only? No MCZ specimens examined. ANOLIS new species Anolis marmoratus, Barbour. A series of 35 specimens collected on Désirade by Richardson clearly represents an undescribed form. It appears to be brown with a greenish tinge. There are pale vermiculations on the trunk reminiscent of leachi. Brown speckles may be present on the trunk; when well defined they also extend onto the head. Solely on the basis of preserved material, without knowledge of the living animals, I do not feel that I can properly describe and name it. Snout-vent. 68 mm. Range. Désirade. MCZ specimens examined. 6166 (35), Désirade. ANOLIS FERREUS (Cope) Xiphosurus ferreus Cope, 1864, Proc. Acad. Nat. Sei. Philadelphia, 168. Type: B.M.N.H. 1946.8.5.59; ‘‘Guadeloupe’’; no collector. Anolis asper Garman, 1887, Proc. Essex Inst., 19:31. Type: M.C.Z. 6162; Marie Galante; coll. Richardson. Anolis asper, Barbour, Not seen alive. The type specimen, a large male, is quite unmistakably the larger Marie Galante species with crested tail. In the general color, as preserved, and in lack of pattern it further agrees with ANOLES OF THE EASTERN CARIBBEAN 203 this Marie Galante form. As Marie Galante les in the adminis- tration of Guadeloupe the erroneous locality ascription could easily occur. All the specimens which I have seen are uniformly light and patternless. According to Barbour the males are bright green or brown with a throat fan of grey with a wide lemon yellow margin. The proximal half of the tail of the male bears a very high crest supported by the neural spines. Largest male, snout-vent: 98 mm.; tail crest; 144% mm. high. Specimen with largest crest, snout-vent: 91 mm.; tail crest: 17 mm. high. Range. Marie Galante. MCZ specimens examined. 6162 (18), 28526-41 + dupl., Marie Galante. ANOLIS SPECIOSUS SPECIOSUS Garman Anolis speciosus Garman, 1887, Bull. Essex Inst., 19:42. Cotypes: M.C.Z. 6172; Marie Galante; coll. Richardson. Anolis speciosus, Barbour. Not seen alive. On the evidence of the five male cotypes, this form is distinct from alliaceus. In life they are evidently green lizards. The anterior head is a little darker than the rest of the body; they are otherwise quite uniform; there are no indications of spots, vermiculations or flank stripe. Snout-vent. 62 mm. Range. Marie Galante. MCZ specimens examined. 6172 (5), Marie Galante. ANOLIS SPECIOSUS TERRAEALTAR, Barbour Anolis terraealtae Barbour, 1915, Proe. Biol. Soe. Washington, 28:76. Co- types: M.C.Z. 10627; Terre d’en Haut, Iles des Saintes; coll. Noble. Not seen alive. On the basis of the type series terracaltae is obviously different from alliaceus; its distinction from speciosus is less striking however. The greenness is less marked, the anterior head is distinctly brown. U.S.N.M. 79002 (4) and 79003 (2?) from St. George (— Cabritt) appear to belong to this form and are plain brown. The four types (86, 12) bear curious irregular brown patches of various sizes on trunk, limbs and tail. They 204 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY look like artifacts but I am unable to visualize a preservation procedure which could produce such markings; perhaps the effect could be produced by splashing with formalin (which causes expansion of the melanophores). On this evidence they may be capable of a green and a brown color phase. The alloca- tion is provisional; both forms may well join the bimaculatus series. Snout-vent. 68 mm. Range. Iles des Saintes. MCZ specimens examined. 10627-30, Terre d’en Haut, Iles des Saintes. ANOLIS OCULATUS (Cope) fod NXiphosurus oculatus Cope, 1879, Proc. Amer. Phil. Soc., 18:274. Cotypes: U.S.N.M. 10139-48, -50, -51, -53 and 7426; Dominica; coll. Ober. Anolis alliaceus, Barbour. Seen alive. After two visits to the island and examination of more than sixty preserved specimens I remain uncertain about the situation in Dominica. The best way to explain my difficulties is to tell the story. On my first visit I collected a number of anoles in Roseau. I sent some alive to the M.C.Z. Williams reported that after making generous allowance for the effects of preservation he could not reconcile them with specimens taken by Barbour in the Botanical Gardens in 1928. I took brown lizards capable of only slight color change and quite incapable of a green phase. Barbour’s specimens were somber green lizards with black- edged white ocelli in oblique rows on the sides. I had apparently taken a new species in one hour ashore without moving outside the capital! On my return visit to Dominica I had therefore simply to make my way to the Botanical Gardens to collect a series of Barbour’s species and thus confirm this interpretation. After two hours of the most careful searching both in the Botanical Gardens and in the surrounding bush I took only two females of the species which I had already met. On my return to the ship a fellow passenger, Dr. W. G. Downs, Director of the Trinidad Regional Virus Laboratory, described to me a ‘‘beautiful Anolis’’ which he had seen by the waterfall. His description closely ANOLES OF THE EASTERN CARIBBEAN 205 matehed Willams’ account of Barbour’s material in the M.C.Z. There had evidently been some shift of population during the last 28 years, but Dr. Downs’ testimony settled the existence of two species. When I examined the large number of specimens in the M.C.Z. I confidently expected to be able to sort them into two species. I was therefore disconcerted to find every kind of an- nectant variant between specimens with black-edged white ocelli and plain brown specimens. The type series is, happily, rather uniform as compared with other series. Males 10139, -40, -42 to -45 bear black-edged white ocelli; only male 10147 is without black markings; the remainder are grown females, save a small juvenile no. 7426. A common pattern variant may be described as follows. The head is a plain brown, about four oblique rows of four white spots on each flank, the upper anterior spots edged with black, a few more white spots on the neck. Light speckles (rather than white) on the back of the neck and down the trunk (between the ocelli) onto the base of the tail. Vague lght mottling on the limbs. The amount of black around the white spots may be reduced to the point of absence. The number of white spots may be reduced. The hight speckling may extend all over the head or conversely may be inconspicuous, although still present. One specimen had small white spots (without black edges) all over the body ineluding the head, limbs and tail. Some specimens showed light speckling on the neck resembling that of marmoratus; oculatus may perhaps be placed as a subspecies of marmoratus. Apart from the difference between my collection and Bar- bour’s collection, I found other reason to suspect local population variation. A series in the U.S.N.M. from near Portsmouth (N. Dominica) had very conspicuous white spots extending onto the head, even onto the snout, showing a tendeney to form oblique rows on the flanks with very little if any black edging. Clearly, Dominica will require further careful examination. Snout-vent. 73 mm. Range. Dominiea. MCZ specimens examined. 6160 (45) Roseau, Dominica; 6159 (19) Portsmouth, Dominica; 28598, 55706-8, Roseau, Dominica. 206 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY ANOLIS ROQUET group Whether the name roquet should be appled to this group is not completely certain ; this is discussed under the species roquet. The group falls into three series of forms each distinguished by color characters. The roquet series consists of spotted lizards. The spots are much darker than the ground color, more or less black, varying in size from no more than one scale to mottles as large as the eye. The ground color is green which can turn dark brown or (in Trinidad) grey. There may be more or less well defined darker transverse bands on the ground color. The pineal window is surrounded by black pigment. This series is represented on the mainland and in Trinidad, Grenada, Grena- dines, St. Lucia, Martinique and Barbados. The trinitatis series consists of plain green lizards which can turn brown. The head is darker than the trunk; the pineal win- dow is surrounded by white pigment. This series is represented in Trinidad, St. Vincent, St. Lucia and perhaps in Martinique. The richardii series is comprised of large lizards, which as adult males show prominent bony ridges on the skull. The colors are rather dull, grey and olive, and there is not much power of color change. This series is represented in Tobago, Grenada and the Grenadines and St. Vincent. Only one form has hitherto been reported from St. Lucia, a member of the trinitatis series. In a few hours collecting in the botanical gardens in Castries I gathered sufficient material to be certain that there is a second species, a member of the roquet series. Last year Mr. G. R. Proctor making a visit of a few hours to St. Lucia went up into the rain forest. He brought back a single small Anolis which had blue eyes. I identified the speci- men as luciae and regarded the blue eyes as an aberration. On a second, extended visit Proctor reported the blue-eyed specimens from several localities and established what I believe to be the first reported case for the Lesser Antilles of two closely related forms with an allopatric distribution, this within an island which would go four times into the area of Rhode Island. I describe and name this form below. ANOLIS ROQUET ROQUET (Lacépéde) Lacerta roquet Lacépéde, 1788, Hist. Nat. Quad. Ovip. Serp., 1. No type; Martinique; no collector. ANOLES OF THE EASTERN CARIBBEAN 207 Anolis alligator Duméril and Bibron, 1837, Erp. Gen., 4:134. 3 cotypes: M.H.N.P. Ig55/1; 1 cotype Tg55/2; Martinique; coll. Droz and Plée. Anolis goudotii Duméril and Bibron, 1837, Erp. Gen., 4:108. Type: M.H.N.P. Ig 55; Martinique; coll. Goudot. Anolis roquet, Barbour. Not seen alive. By itself, Lacépéde’s description is not sufficient to make the species identifiable. On the testimony of Doflein (1908, p. 245), there are three species of Anolis in Martinique, a green, a brown, and a grey with spots. The museum material which I have examined does not appear to be referable to more than one species. In the hght of field experience I would suggest that Doflein’s evidence argues for the existence of at least two species. The green and the brown may well be color phases of the same species; I have met people in Jamaica who have thought the ereen and black phases of Anolis grahami to be different species. Williams suggests that this may be an undiscovered Martinican representative of the ftrinitatis series. The other form is de- seribed by Doflein as ‘‘hellgrau mit dunkleren Flecken mar- moriert.’’ This agrees with the form in museum collections, and Lacépede’s remark ‘‘couleur de feuille morte, tachetee de jaune et de noiratre’’ certainly agrees better with the form in collec- tions than with a plain green or plain brown lizard. It therefore seems reasonably sure that the name ‘‘roquet’’ is properly ap- plied. If we allow roquet as the correct name, then alligator is cer- tainly a synonym. The Droz cotypes are soft and faded but the color indicates clearly that they agree with Martinique speci- mens. The Plée cotype is shrivelled but the pattern shows clearly and unmistakably and there is no doubt that it is also the Mar- tinique form. The type of goudotu is a small, probably female juvenile which is virtually unidentifiable. A broad light stripe with dark edges, interrupted on the base of the tail, runs down the back. This pattern agrees quite well with some juvenile roquet. At present I see no reason to question the allocation of goudoti as a synonym of roquet. If, however, material of two or more spe- cies is collected on Martinique the type of goudotw will have to be re-examined. 208 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY It is evident from preserved material that roquet is a variable form. There appear to be three main elements in the various eolor patterns: white spots on the flanks which tend to be ar- ranged in oblique transverse rows; transverse bands, dark on a hehter ground, which include the rows of white spots when they are present; mottling of the ground color which may range from fine vermiculation to conspicuous marbling. The head is usually uniform and darker than the body but the mottling may extend onto the head. Conspicuous mottling, transverse banding and conspicuous white spots may be present simultaneously. Com- monly, the mottling is inconspicuous and the bands also incon- spicuous. Less commonly the white spots are reduced; in a few specimens they were absent. The general color of the lizard is evidently green in life. Barbour reports that the throat fan is a ‘‘rich orange yellow.’’ Snout-vent. 75 mm. Range. Martinique. MCZ specumens examined. 5757, 6008 (30), 6188 (24), 6189 (3), 20526. The other members of the roquet group are perhaps best un- ravelled by starting with a discussion of the forms on Trinidad. When I saw living examples for myself I was immediately con- vinced of the claim of Messrs. Quesnel and Kenny that ‘* Anolis aeneus’’ consists of two species. For the purposes of the follow- ing discussion | will refer to these two species as the ‘‘spotted”’ form and the ‘‘green’’ form. Two names are available: Anolis aeneus Gray 1840 and Anolis trinitatis Reimhardt and Liitken 1862. ANOLIS ROQUET AENEUS Gray Anolis aeneus Gray, 1840, Ann. Mag. Nat. Hist., 5:114. Type: B.M.N.H. 1946.8.28.7; no locality; pres. T. Bell. Anolis gentilis Garman, 1887, Bull. Essex Inst., 19:34. Type: M.C.Z. 6163, Petit Martinique; coil. Garman. Anolis aeneus (partim) Barbour. Trinidad specimens, but not those from the Grenadines, seen alive. The type is a half-grown male without locality. ‘* Tropical America’’ given by Barbour is evidently a later addition; in any case Barbour does not give the original literature citation. ANOLES OF THE EASTERN CARIBBEAN 209 There is no evidence of locality at all. Gray gave an account of material collected by MecLeay in Cuba. At the end, he de- scribed a few more odd specimens in the museum; the only information about this one is that it was presented by Thomas Bell. This proved to be the most difficult type specimen to allocate. It is quite well preserved but somewhat faded; as a half-grown male it would not show a distinctive color pattern in any ease. I am quite satisfied that it is not the green form; the pattern, as much as is visible, does not agree and, for what it is worth, the squamation also does not agree. It shows definite re- semblances to small specimens of the spotted Trinidadian form. I find it difficult to make a plausible allocation to territory. Relevant to this question is material from the Grenadines, Grenada, Barbados, Trinidad and British Guiana. I have seen only preserved material from British Guiana and the Grenadines ; of these latter I have seen series from Becquia (B.M.N.H.) in the northern part of the group, Cannouan (B.M.N.H. and M.C.Z.), near the center, and Carriacou (B.M.N.H. and M.C.Z.) and Petit Martinique (M.C.Z.) in the south. The Grenadian and Barbadian lizards are in life spotted on a green ground color. The Trinidadian lizards are different in that they are spotted on a grey ground color; they are incapable of a green phase. The preserved material from British Guiana agrees well with the Trinidadian and gives no hint of greenness. The pre- served specimens from the several Grenadines agree well with one another and, surprisingly, agree much better with Trini- dadian specimens than they do with Grenadian or Barbadian material. A few of the Grenadine specimens show a green tinge; none shows the clear indications of greenness shown by pre- served Grenadian specimens. The metallic tinge of the aeneus type can be better matched by small Trinidad and Grenadine specimens. In respect of color pattern and squamation it showed the best combined agreement with a specimen of similar size from Beequia; it agreed in respect of the transverse marks behind the ehin and the longitudinal marks on the throat and in the faint indication of pattern beside the neck and over the shoulder. I therefore consider aeneus to be the correct name for the grey Trinidadian species and gentilis provisionally to be a synonym 210 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY of aeneus. Mr. Shreve had come to the same rather surprising conclusion that the Grenadine specimens resemble the Trini- dadian, and the Grenadian resemble the Barbadian. The placing of gentilis as a synonym of aeneus is provisional pending a direct comparison of living Grenadine and Trinidadian material. If it transpires, as well it may, that there is a distinction between Grenadine and Trinidadian specimens, then the question of the provenance of aeneus will become acute and the type will have to be compared anew. Even then it may well be necessary to claim reviser’s privilege to assign a type locality to aeneus. I hope that Trinidad will be so designated, thus making the name gentilis available for the Grenadine form; this latter name is founded on a series of specimens of exact provenance and in- cluding adult males. Trinidadian aeneus is essentially a grey lizard; there may be a trace of copper green. There are fuzzy warm brown transverse bands, five on the trunk, extending onto the tail. The whole is liberally sprinkled with dark spots. The lizard is capable of little color change. The fan is a dirty grey with sometimes a faint wash of verdigris. The Grenadine specimens look as though they are sometimes a little greener; the majority resemble the Trinidad specimens closely. There were no obvious indications of differentiation within the Grenadine islands. Snout-vent. 75 mm. Range. Trinidad, the Grenadines and British Guiana. MCZ specimens examined. 6163, Petit Martinique, Grenadines ; 6187 (18), 6192 (8), Port of Spain, Trinidad; 20531-5, Carri- acou, Grenadines; 33387-92, Canouan Id., Grenadines; 55534, 55537, Trinidad: 55675, 55787-800, Port-of-Spain, Trinidad. ANOLIS ROQUET CINEREUS Garman Anolis roquet var. cinereus Garman, 1887, Bull. Essex Inst., 19:35. Type: MCZ 6182; St. George, Grenada; coll. Garman. Anolis roquet var. extremus Garman, 1887, Bull. Essex Inst., 19:35. Type: MCZ 6183 Bridgetown, Barbados; coll. Garman. Anolis roquet gentilis partim Barbour. Anolis aeneus partim Barbour, Seen alive. On the basis of preserved material both the Grenadian and Barbadian populations are very variable and overlap so broadly ANOLES OF THE EASTERN CARIBBEAN 211 that I cannot at present see any ground for making a nomen- clatural distinction between them. They both have a green ground color which can change to a warm dark brown. Some of the living specimens seen in St. George’s were rather yellowish-green, some in Bridgetown rather bluish-green. The skin around the eyes is dark; the rest of the body is sprinkled with dark spots. Indications of transverse bands which descend the flanks obliquely were very variable. In some specimens there is no indication of banding, while in others the ground color is a little darker in the form of bands; in still others there are pronounced dark bands. In these latter cases the bands may be associated with oblique rows of white spots. Throat fan medium yellow ochre. Snout-vent. 74 mm. Range. Grenada, Barbados. MCZ specimens examined. 2748 (11), Barbados; 6182 (19), St. George, Grenada; 6183 (17), Bridgetown, Barbados ; 8132-56, Sauteurs, Grenada; 55401-6, Jamestown, Barbados; 55536, Gre- nada; 55764, St. George, Grenada. ANOLIS ROQUET ?CINEREUS Garman I provisionally refer to cinereus the roquet representative from St. Lucia, here reported for the first time. The possibility that it is an introduction should be considered. Museum material of Grenadian and Barbadian cinereus shows considerable variation which may well be sufficient to include the St. Lucian cinereus. The St. Lucia specimens are known only from Castries. Fur- thermore, from personal recollection of the living animals and the appearance of the preserved specimens there is a special resemblance between Castries cinereus and Bridgetown cinereus. The description below of the colors in life is based on personal observations, Proctor’s field notes and three specimens received alive through the kindness of Mr. Gregory Williams. Ground eolor: head and back lavender grey, grades into yellow-green on the flanks, forelimbs, hindlimbs and tail; belly warm chrome- yellow; underside of head grey-blue. Throat fan bright orange- yellow. Body sprinkled with ragged black mottling of variable extent, heaviest on top of the head and dorsal neck, more sparse on back, extends only slightly if at all onto limbs and tail; in- 212 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY conspicuous but distinct lavender grey flank stripe from shoulder to groin. Snout-vent. 68 mm. Range. St. Lucia, Castries only? MCZ specimens examined. 56385-93, Castries, St. Lucia. ANOLIS TRINITATIS TRINITATIS Reinhardt and Liitken Anolis trinitatis Reinhardt and Liitken, 1863, Vid. Meddel. Kjobenk., 1862:269. Type: Universitetets Museum, Copenhagen; Trinidad; coll. Riise. Anolis aeneus (partim) Barbour. Seen alive. This is one of the species of which I have not seen the type specimen. As this specimen bears a precise locality, the only point at issue was whether it is referable to the spotty or the green species of Trinidad. I accordingly wrote to Dr. F. W. Braestrup and he very kindly checked the specimen and the original description. I am happy to say that his reply leaves no doubt but that Reimhardt and Littken described and named the green form. I say ‘‘happy’’ because these authors worked at a time when many workers were quite content to found new species on badly preserved immature specimens of unknown origin with- out a word to suggest that they realized that the animals had ever been alive. I quote from Dr. Braestrup’s letter: ‘‘Concerning Anolis trinitatis | am sorry to say that the type is very badly faded, but we have the statement by R. and L. towards the close of the Danish description (translated by me): ‘The color is bluish green on the body; more bluish violet, bronzy on the limbs. The head is darker and especially it seems to have a black area around the eyes; in some a light stripe along the upper lip under the eye is very distinct.’ (There is further a footnote to this stating ‘In the living animal these markings are perhaps not found at all.’) ... the specimen designated type is 70 mm. from snout to vent. R. and L. were, I believe, always very careful in their deserip- tions and I think that we may be sure that there were no trans- verse bars on the body. . . . The type designation is found in our old catalogues and it was no doubt made by the authors themselves even although they did not in print expressly desig- ANOLES OF THE EASTERN CARIBBEAN 213 nate it.’’ The type specimen is somewhat larger than the largest Trinidad specimen which I collected. In life trinttatis is a bright grass green, and can turn a dark brown. The skin around the eye is dark and on the head and anterior trunk is a little fine blue mottling. Snout-vent. 70 mm. Range. Trinidad. No MCZ specimens at the time of writing. ANOLIS TRINITATIS VINCENTIL Garman Anolis vincentii Garman, 1887, Bull. Essex Inst., 19:46. Types: MCZ 6178-9; St. Vincent; coll. Garman. Anolis vincentii Barbour. Seen alive. The St. Vineent anoles resemble triuitatis but the series of living specimens seen showed some small but clear differences. The ground eolor is the same but the hght blue mottles on the anterior trunk are more extensive in vincentu. The clearest dif- ference is that in vincentw the area around the eye is hight blue in the living animal. The throat fan is a similar orange-yellow. Snout-vent. 70 mm. Range. St. Vincent. MCZ specimens examined. 6178 (47), 6179 (22), 20542-8, 38185-9, 38382, 55532, 55993, Kingstown, St. Vincent. ANOLIS TRINITATIS LUCIAE Garman Anolis luciae Garman, 1887, Bull. Essex Inst., 19:44. Type: MCZ 6173; St. Lueia; coll. Garman. Anolis luciae, Barbour. Seen alive. I have placed this form in the trinitatis series at the sugges- tion of Dr. Williams; it is a suggestion with which I fully agree. This form reaches a greater length of body and is more slenderly built than the roquet representative in St. Lucia. The general color is brown to olive; the power of color change is auite limited ; the belly is white. The head is plain brown; the lips are paler, flecked with darker pigment. The pineal window is larger than in roquet and surrounded by white pigment. The iris is black ; 214 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY the fan has a pale yellow center. There is vermiculate speckling of brown on olive, especially on the anterior trunk; there is a light shoulder stripe which does not pass the elbow. Snout-vent. 81 mm. Range. St. Lucia, recorded from Castries (Barbour, Under- wood) and Vieux Fort (Proctor). MCZ specimens examined. 6173 (20), 6175 (1), Castries, St. Lueia; 20549, St. Lucia; 56394-411, Castries, St. Lucia. ANOLIS TRINITATIS PROCURATORIS subsp. nov. Type: M.C.Z. 57202, adult male, Savanne Edmund, St. Lucia, 13°47’N, 61°14’W, collected by G. R. Proctor, May 1958. @ At.procuratoris oO Atluciae 4 Ar.cinereus x A.wattsi Fig. 2. St. Lucia. Localities for the four anoles known from the island. Paratype: M.C.Z. 57203, adult female, same data as type. Color in life reported by G. R. Proctor. Diagnosis. An Anolis of the trinitatis series closely related to luciae. Differs from all the other members of the series in the blue iris and the greenness of the belly and the underside of the head; differs from luciae, additionally, in the power of color change from green to brown. ANOLES OF THE EASTERN CARIBBEAN PAILS) The general color is blue-green, the animal is capable of marked color change to an olive brown, the belly and the underside of the head are green. The head is brown; the area around the pineal window is a little more conspicuously white than in luciae. The iris is cobalt blue, the throat fan has a pale yellow center as in luciae. Indications of vermiculation are slight; the shoulder stripe is almost obsolete. No consistent differences of squamation were found. Apart from the type locality, this form is recorded from Mar- quis (Proctor, 1 specimen) and Bare de l’isle (Proctor, speci- men). I received three specimens from Soufriére collected by Mr. Gregory Williams. He reported that the eyes were blue in hfe; as preserved, they differ from the type in that they are yellow-green rather than blue-green. He also reports blue- eyed anoles from La Vigie about 114 miles northwest and La Clery about 1 mile north of Castries. The boundary zone evi- dently runs close to the capital. The records for this form suggest that the range of luciae may be discontinuous, interrupted by procuratoris; they further sug- gest that procuratoris may not be completely confined to forested country. We have a pair of forms, one showing marked color change and found (mostly) in forested country and one showing more limited power of color change and found in more open eountry. The St. Lucian pair procuratoris and luciae may thus be compared to the Jamaican pair Anolis lineatopus neckeri and All. lineatopus. Neckeri shows a considerable power of color change and is found in forested country; lineatopus shows a limited power of color change and is found in drier more open country. Caribbean reptiles bear many patronymics, some of which mean rather little. Mr. G. R. Proctor is a field botanist who has shown himself to be a discriminating collector of reptiles, on this and other occasions, and I am happy to make this recogni- tion of his contribution. The name Proctor is a syncopated form of procurator which is a classical Latin term. Snout-vent. 74 mm. Range. St. Lucia: Savanne Edmund, Marquis, Soufriére, Barre de 1’Isle, La Vigie, La Clery. 216 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY ANOLIS RICHARDIU RICHARDID Duméril and Bibron Anolis richardit Duméril and Bibron, 1837, Erp. Gen., 4:141. Type: M.H.N.P. Ig 53; ‘‘Tortola’’; coll. Richard. Anolis occipitalis Gray, 1840, Ann. Nat. Hist., 5:112. Types: B.M.N.H. 1946.8.12.59, 1946.8.29.11 ‘‘West Indies’’; coll. T. Bell. Anolis stenodactylus Gray, 1840, Ann. Nat. Hist., 5:114. Type: B.M.N.H. 1946.6.8.12.54 ‘‘Jamaica’’; coll. J. Winterbottom. Anolis trossulus Garman, 1887, Bull. Essex Inst., 19:38. Type: M.C.Z. 6181; Grenada; coll. Garman. Anolis richardii, Barbour. The type specimen is a half-grown male. No color pattern is visible. The specimen bears a detailed resemblance to Grenadian and Tobagonian material. The populations in the two islands so closely resemble one another that I could not guess from which island the specimen had more probably come. Adults are a dull olive green. There are cream markings on the lips which may extend onto the temporal region. There are variable indications of dark hour-glass markings across the back. The throat fan is a light ochre yellow and very large (but the basihyoid cartilage is not correspondingly long). The lizards ean turn a dull dark brown. The smaller individuals are a brighter green with a warm dark brown phase. Some have a light stripe on the shoulder which starts obliquely backwards and upwards to fade on the flank; a second parallel stripe starts below it on the flank and runs backwards and upwards to the eroin,. I did not see any juveniles in Grenada but the similarity of the adults is close. Snout-vent. 125 mm. Range. Tobago, Grenada (and the Grenadines fide Cochran 1934 and 1938). MCZ specimens examined. 6181 (12), 8108-31, St. George’s, Grenada; 10748, Tobago; 12016-26, Milford Bay, S.W. Tobago, 55676, Grenada. ANOLIS RICHARDII GRISEUS Garman Anolis griseus Garman, 1887, Bull. Essex Inst., 19:36. Type: M.C.Z. 6164; St. Vincent; coll. Garman, Anolis griseus, Barbour. ANOLES OF THE EASTERN CARIBBEAN PAT This form was seen only preserved. It appears to be somewhat lighter than richardu; the ground color has a greenish tinge. The body generally bears reticulate or vermiculate markings. There is a short light stripe over the shoulder only; a longer stripe starts below the axilla and runs backwards and upwards to the groin. There is no evidence of creamy markings on the lips. There may be dark brown spots on the anterior trunk which tend to form oblique rings. If the specimens examined represent a fair sample of the population, then these lizards do not grow quite as large as richard. Snout-vent. 117 mm. Range. St. Vincent. MCZ specimens examined. 6164 (5), St. Vincent; 38380-1, Fountain Estate, St. Vincent. ANOLIS WATTSI Boulenger Anolis wattsi Boulenger, 1894, Ann. Mag. Nat. Hist., (6)14:375. Type: B.M.N.H. 1946.8.29.12-13; Antigua; coll. Watts. Anolis forresti Barbour, 1923, Occ. Pap. Mus. Zool. Univ. Mich., 132: Type: MCZ 16170; Barbuda; coll. Forrest. Anolis wattsi, Barbour. Antigua and St. Kitts specimens seen alive. Antiguan specimens of this small lizard are attractively col- ored. Overall is medium brown. The head and especially the snout is bright russet. There is a luminous translucence about the colors. The temples are blue-grey. The throat fan is light orange. Only two specimens were seen in St. Kitts. They had only a little russet on the snout and lacked the blue-grey on the temples. Museum specimens indicate the occurrence of this lizard in Anguilla, St. Martin, St. Eustatius, St. Kitts, Nevis, Barbuda and Antigua; it presumably also occurs in St. Barthélemy. It has not been reported from Montserrat. In Proctor’s St. Lucia material Williams found specimens of Anolis wattsi collected in Castries. This startling discovery was confirmed by a collection of anoles from Castries which Mr. Gregory Williams shipped to me. Unfortunately, the three speci- mens of wattsi were dead and I can only report that the colors of the dried animals resemble Antiguan watts’. It may well turn out to be a distinct subspecies. 218 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY Snout-vent. 49 mm. Range. St. Martin, Anguilla, St. Kitts bank, Antigua bank, St. Lueia. MCZ specimens examined. 10134, St. Johns, Antigua; 12334- 6, St. Eustatius; 16170, 16201-5, Barbuda; 16213-15, Nevis; 16538-43, St. Eustatius; 16596-7, Anguilla; 20986, St. Martin; 21676-9, Antigua; 28563-68, St. John, Antigua; 28696-99, St. Kitts; 38375-6, 6 mi. E. of Charleston, Nevis; 54708, St. Eusta- tius ; 55460 nr. Crab Hill Village, Antigua; 55461 nr. All Saints, Antigua; 55462-4 nr. Old Road Town, Antigua; 55465, Boggy Peak, Antigua; 55535, 55682-5, St. John’s, Antigua; 57220-1, Castries, St. Lucia. F'TELD OBSERVATIONS Basseterre, St. Kitts, was the poorest collecting locality of my trip. In the higher class residential section of the town, to the northwest, there were numbers of bimaculatus. A search for cool and shady clumps of trees which might be expected to harbor wattsi was fruitless. The ‘‘Botanical Station’’ proved to be merely a rather bare nursery. Two watts: were eventually found in a churchyard. In Montserrat, Anolis lividus was abundant both in shady places in Plymouth and in open situations in the cemetery. No indications of the occurrence of a second species were seen. The botanical gardens in St. John’s, Antigua, proved to be a good collecting ground. Leachiw was the more obvious because it occurred in more open situations. In the gardens wattsi was common but appeared definitely to prefer shady situations; it is therefore probably more localized in its occurrence. All the anoles collected in Dominica were in open situations. Some were on walls and trees in Roseau, others were seen on trees outside the town. In woodland around the botanical gar- dens no anoles were seen. In St. Lucia there was good collecting in the botanical gar- dens; with the aid of small boys a good series was collected. My impression is that luciae prefers open situations and that the roquet form prefers shady places. This difference in habits did not appear to be sharply marked however. ANOLES OF THE EASTERN CARIBBEAN 219 There was no difficulty in obtaining a series in Bridgetown, Barbados, on trees, in open situations, in yards, and by the creek. I did not have time to get very far in Kingstown, St. Vincent; vincentii proved to be quite common on trees and fences in the town along the waterfront. I was told about a larger tree lizard, evidently griseus. It is known locally as ‘‘coco-bé.’’ Later I was told that a distinction is made between ‘‘coco-bé”’ and ‘‘coco- tom,’’? brown and green respectively. I am not clear whether these are color phases of the same species or griseus and vin- centi, respectively. In Grenada I made my way from St. George’s towards the botanical gardens but never arrived, for there was excellent col- lecting on roadside trees. Both cinereus and richardi were in evidence, and both appeared to be common. They were seen on the shaded side of trees out in the open. Richardi was not found on fence posts where cinereus was however seen. On the road up to the fort some cinereus were also seen on stone walls. In Trinidad, very fortunately, as it now turns out, trinitatis was quite common in the garden of the guest house in which we stayed near the southwest corner of Queen’s Park Savanna. Aeneus was not seen in this garden, which in the course of one month was the only locality in which I saw trinitatis. In gardens 100 metres away aeneus was found to the exclusion of trinitatis. Along a line of fence posts enclosing the dock area, in a very open and sunny situation, aeneus was fairly common. The one garden in which trinitatis was found was, on the other hand, cool and shady. The unrelated Trinidad species chrysolepis was not seen. I was informed that it occurs in the forest and is relatively rarely seen. In Tobago, richardii is fairly generally distributed. It was collected in and around Scarborough, at Store Bay and at Spey- side. It appeared to have a liking for coconut trees. When disturbed the anoles would run right up the coconut trees to the crown. This anole is known as ‘‘gumangala’’; this is the only Caribbean island in which I have met a genuine vernacular name for the common tree lizards which may be seen anywhere. I was puzzled by the apparent absence of the two Trinidad forms. | looked most carefully around clumps of trees and in old build- 220 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY ings in Searborough without discovering any suggestion of the presence of another species. DISCUSSION These anoles show well the dual influence of the Greater An- tilles and South America on the fauna of the Lesser Antilles. I regard the evidence as fairly good that the bimaculatus group, of Greater Antillean affinities, extends no farther south than Dominica. Complementary to this, we see the extension of the roquet group as far north as Martinique. There is every reason to believe that anoles disperse readily; I am therefore at a loss to explain why the two groups do not overlap. Further collecting may show that either Dominica or Martinique does in fact pos- sess two stocks of anoles. I find it difficult to believe that com- petition has excluded a second species from either of these two islands. On a single tree in Jamaica I have seen four species of anoles, and a total of six species in one group of trees. The distribution of Anolis wattsi through the Leeward Islands nicely illustrates the process of colonisation of a group of islands. We have the absence from the southeast and the shght measure of differentiation between the islands as evidence of relatively recent arrival from the west. Watts: should be sought on St. Barthélemy, a poorly known island. It is perhaps genuinely absent from the better known island of Saba. Saba also lacks the almost ubiquitous Ameiva; the lack of beaches may make random colonization of this island difficult. The differentiation of the bimaculatus stock within the Lee- ward Is. suggests that it is old established. West of Guade- loupe the differentiation runs along the sort of lines which might be expected. The stocks on the three islands of the An- euilla bank are closely alike; their separation is probably recent. It is a little surprising that the Saba stock should most closely resemble the Anguilla bank stock; it is however the nearest island to this bank. The St. Eustatius, St. Kitts and Nevis stocks re- semble one another, the closest resemblance being between the latter two. On the basis of scanty material the stocks on the two islands of the Antigua bank also resemble one another. The Leeward Islands consist of two chains: an outer chain of older islands running from the Anguilla bank to Marie Galante ANOLES OF THE EASTERN CARIBBEAN Di Anquilla gingivinus wallsi SL Marlin gingivinus walts7 St. Barthélemy gingivinus ? Antiqua leachii waltsi Barbuda keachii waltsi Saba g sabanus su St Eustatius & bimaculatus S wattsi St Kitts bimaculatus _ Wallsi Nevis = bimaculatus =~ wattsi Redonda S| nubilus = CSN sa fe Montserrat S| lividus yt ~Q Désivade Sp. NOV. Guadeloupe alliaceus — marmoxatus 2 ae Les Sainles lerraeallae ne = as Marie Galanle speciosus lerreus fas Dominica bs oculalus m7 fu Martinique roguet i: ost it St Lucia S AA, DD ca. = 14C. Nephropores unrecognizable and microscopic. First dorsal pore at 8/9 (2 specimens), ?9/10 (4 specimens), 9/10 (6 specimens). Clitellum, saddle-shaped, reaching ventrally to B, interseg- mental furrows obliterated, dorsal pores not occluded and pre- sumably functional, setae probably present but deeply retracted and exceedingly difficult to recognize, on (xil1?)Xiv-xix(xx? 1 specimen), (xiii?)xiv-xx (1 specimen), xiv-xix (2 specimens), Xiv-xx (8 specimens). 1 The discussion of classification is from a manuscript, written during tenure of a John Simon Guggenheim fellowship, 1952-1953, but with such minor changes as were required to bring it up to date, March 1959, 230 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY Spermathecal pores minute, superficial, at or slightly median to B, slightly in front of or on segmental equators, two pairs, in vill-ix. Female pores at or slightly lateral to A, about equidistant from 13/14 and eq/xiv or slightly nearer 13/14. Male pores minute (each on a very small tubercle at eq/xviii?). Prostatic pores minute, two pairs, in AB at eq/xvii and eq/xix. Seminal grooves nearly straight or slightly concave laterally, between eq/xvli and eq/xix, at or slightly median to B, deep and wide (i. e., not mere linear furrows), containing male pores and with prostatic pores at the ends. Each groove is within a protuberant, longitudinally elliptical area (of epidermal thickening?). A deep slit-like crease at mV crosses all of xvii-xix, reaching into xvi (2 specimens), through xiii (1 specimen). Genital markings unpaired, transversely placed, presetal, in BB, on xx and xxi (12 specimens). Additional markings of the same sort as follows: on xy-xvi (2 specimens), xvi (1 speci- men), Xv, xvi and xxii (1 specimen), xvi and xxii (3 specimens). A central portion of each marking is greyish and translucent. Internal anatomy. Septum 5/6 funnel-shaped, slightly strenethened and translucent, 6/7-10/11 thickly muscular. Lon- gitudinal muscle band at mD distinct from level of first dorsal pore or pore-Lke marking, unrecognizable anteriorly. Pigment unrecognizable in body wall except at mD, the muscle band, in one or more sc¢attered segments or through several consecutive metameres, often dark red. The subesophageal mesentery in X-xlli seems to be slightly strengthened. Gizzards well developed, lined with thick cuticle, two, in v-vi (12 specimens). Cuticular lining continued from gizzards through viii or ix (at least?). Esophagus widened and monili- form in xi-xiil where there are (internally) closely crowded rather lamelliform but low and vertieal ridges as well as a bifur- eated ventral typhlosole (12 specimens). Intestinal origin in region of 15/16. Intestinal typhlosole lacking (?). Dorsal blood vessel single, complete, bifureating under the brain, the branches passing ventrally and uniting over the sub- esophageal ganglion to become the ventral trunk. Latter also complete, with two pairs of branches between 4/5 and the an- terior bifurcation. Extra-esophageal trunks filled with blood anteriorly and traceable forward into i, posterior portions un- recognizable, median to hearts. Supra-esophageal trunk present GATES: CLASSIFICATION OF EARTHWORMS 24-3). | in 1x-xill. Subneural trunk lacking (?). Hearts of x-xill ap- parently latero-esophageal though blood is lacking in filamentous branches to the dorsal trunk. Last hearts in xiii (11 specimens). Hearts of ix-vi (all easily traced to ventral trunk), lateral. Hearts of v may pass to ventral vessel on one or both sides but usually are not traceable to that trunk. Brain in ii or in iii. Exeretory system meronephric. Astomate micronephridia cover body wall in (11?) iii-iv. At least from iv the number per segment decreases, to 40-50 in x, the small astomate tubules in the latter metamere and posteriorly in a row just in front of the septum and from region of A nearly to mD. Several nephri- dia are crowded together at the median end of a row but as far back as tubules are distinguishable all are astomate. Holandrie (12 specimens). Male funnels, and presumably also testes, free in x-xl. Male deferent ducts slender, becoming un- recognizable shortly behind funnel septa and after passing to parietes. Seminal vesicles medium-sized or smaller, finely aci- nous, low down in coelomic cavities, two pairs, in 1x and xu. Prostates medium-sized, flat discs, racemose,” two pairs, each usu- ally extending through three segments, usually three to six levels behind xvii or xix with their ducts passing anteriorly through septal perforations, occasionally in xvi-xvii and xvili-xx, once seemingly confined to xvii and xix but bulging septa far pos- teriorly. Duet slender but probably muscular, 4-6 mm. long. Spermathecae fairly large but not reaching dorsal parietes, always bound to anterior septum of their segment by fairly strong tissue from which the ectal half of the duct is free. Duet slender, probably muscular though sheen no longer recognizable, shghtly widened entally but an appearance of even greater widening is due to presence of one or two very short loops bound closely together. Seminal chamber ellipsoidal, sessile vertically on entalmost portion of duet into which it opens entally by a single aperture. GM glands represented by clusters of several very small spheroidal bodies, the clusters always covering the parietes over sites of genital markings except in two specimens with least obvious clitella and in which the little glands may not have 2 Without a central lumen. (Sections have been deposited in the Museum of Comparative Zoology.) Dae BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY reached definitive size. (These glands, in some of the specimens, are not easy to distinguish from the nephridia. ) Ventral follicles (a and b) of viii, ix, xvii and xix, completely buried within the parietes, contain setae that are much slenderer and shorter than those of neighboring segments. Shafts of these smaller setae are straight and taper ectally but not to a sharp point. Ornamentation on the penial setae, as those of xvii and xix presumably should be ealled in spite of their small size, is of several (four?) longitudinal rows of four or five excavations. Each pit is long, narrow, deepened entally where its floor is finely nodose. Excavations on the copulatory setae (a and b of vul-ix) are of the same sort but more numerous, in longitudinal rows of seven to nine. Eetalmost pits are closer to the tip than on the penial setae but ends of copulatory setae may have been subjected to more erosion. Ventral setae of xvili appear to be lacking as their follicles were not found. Reproduction. Tridescence on male funnels of each specimen (including the aberrant one) proves that sperm had been ma- tured. A similar iridescence in the spermathecal seminal cham- bers of each worm shows that copulation had taken place. Reproduction, accordingly, is assumed to be sexual and _ bi- parental. Ingesta. The gut of each worm is filled with a sort of humus in which plant parts or tissues are not distinguishable. Little or no sand and clay is present. These worms appear to be dis- eriminating feeders. Abnormality. Organs of right side belonging in vill-xix are (1 specimen) one segment anterior to their normal location; spermatheeal pores on vii-vili, female pore on xiii, prostatic pores on xvi and xviii, last heart in xii, testes in 1x-x, seminal vesicles in vili and xi, ete. The elitellum is on xili-xx on both sides. Both hearts of ix are lateral. Presumably a mesoblastic somite at the seventh level (or anteriorly?) was aborted early in development. Remarks. The worms may have been quickly killed and prop- erly preserved. Unfortunately, however, subsequent care had been lacking and all are macerated, especially from region of xxiil to hind end. The size of properly preserved, contracted specimens is estimated to be between 150-250 x 9-10 mm. Loea- tions of prostatic pores were confirmed by dissecting prostatic GATES: CLASSIFICATION OF EARTHWORMS 233 ducts out of the body wall. Location of male pores, inasmuch as vasa deferentia cannot be traced, requires confirmation. No evidence was found to indicate junction of those gonoducts with the prostatic ducts. Some of the seeming variation in antero- posterior extent of the clitellum may be due to incomplete de- velopment as the tumescence, in several of the specimens, is slight. The intestinal wall had been reduced to a transparent slime in which caeca and supra-intestinal glands might not have been recognizable. A typhlosole, if present, should have been distinguishable as a lamella of slime extending down into gut contents. When a typhlosole is absent supra-intestinal glands usually are lacking. Complete relaxation of the esophageal valve in each specimen, along with the maceration, obviated determi- nation of site of intestinal origin which is unlikely to be variable, except as a result of some aberration in embryonic development. Characterization of excretory organs behind xxx is impossible. If nephridia are exonephric throughout the body all may be small and astomate as in Octochaetoides, or the medianmost on each side of some posterior segments may be larger (as in various octochaetine genera) and provided with a preseptal funnel. Although less lkely perhaps, some or even all of the micro- nephridia, along a major posterior portion of the axis, may be enteronephrie and stomate. Presence in posterior segments of one to several pairs of large, stomate, enteronephric nephridia is not anticipated. Uniformity in number of prostates and in relation of their external apertures to the male pores among so many specimens, even including one that is aberrant, seems to warrant an as- sumption that the arrangement of the male terminalia is charac- teristic of the species to which these worms belong. An ‘‘acanthodriline’’ arrangement of racemose prostates has been unknown hitherto and requires, in the Oligochaeta where a single character rarely is diagnostic at any taxonomic level, erection of a new genus. Although diagnosis is assured, determination of relationships must await much more adequate description of digestive, vas- cular and excretory systems than is permitted by available material. Some generic characters cannot, of course, be recog- nized when only one species is known. Intestinal origin, for instance, is uniform in some genera, possibly even throughout one 234 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY family (Lumbricidae), but is subject to individual variation in the Moniligastridae. A typhlosole is lacking in some genera and even throughout one family (Moniligastridae), but is now known to be absent only in some of the species of several genera. Num- ber of gizzards is uniform in some genera and subfamilies, but in the Moniligastridae is subject to individual variation. Andry, in spite of the reliance placed on it in the classical system, does vary intragenerically, sometimes even intraspecifically. Hence, the definition below is tentative. In fact, any generic definition must be considered liable to revision, at least until all species have been satisfactorily described. A definition certainly does not become sacrosanct through publication in a thick mono- graph. Yet species, genera, even one family, have been erected because of unimportant difference from one or more characters as stated in definitions. Taxonomically important characters that are unknown or in need of confirmation, in accordance with previous practice, are indicated below at specific as well as generic level. Genus EHXxXUS gen. nov. Definition. Quadriprostatic, prostates racemose and of phere- tima sort but with ducts opening externally at equators of xvi and xix. Male pores (in seminal grooves that extend from eq/xvil to eq/xix) at eq/xvili (?). Setae, eight per segment and paired (throughout ?). Gizzards in v-vi. (Intestinal origin in xv?) Calciferous glands (typhlosole, intestinal caeca and supra-intestinal glands?) lack- ing. Vascular system with complete (single) dorsal trunk, with extra-esophageal trunks median to hearts, with a supra-esophageal trunk in ix-xiii but without a subneural(?), (lateroparietal trunks?), and with latero-esophageal hearts in x-xui. Exere- tory system meronephric, nephridia astomate and exonephric (throughout or enteronephrie in ii-iv?), massed on parietes in iii-iv, but posteriorly — on each side of each segment — in a transverse row, extending from A nearly to mD just in front of the septum. Type species. EHxxus wyensis sp. nov. Precis of E. wyensis. Quadrithecal, spermathecal pores mi- nute, superficial, two pairs, at or slightly median to B, on or GATES: CLASSIFICATION OF EARTHWORMS 235 slightly in front of segmental equators, in vill-ix. Female pores at or slightly lateral to A, about midway between 13/14 and eq/xiv. Male and prostatic pores minute and superficial. Genital markings unpaired, in BB, presetal, on xx and xxi, often with one or more similar markings in some of xv-xvi, xxii. Setae, rather closely paired, DD ca. = 144C (throughout?), ventral setae of xviii lacking at maturity (?), @ and b of xvii and xix (penial) as well as of viil-ix (copulatory) concealed in small follicles within parietes and ornamented ectally by several lon- gitudinal rows of narrow pits that are deeper entally and there with a nodose floor. Clitellum, saddle-shaped, on xiii, xiv-xix, xx. First dorsal pore at 8/9 or 9/10. Prostomium, epilobous(?). Pigmentation, red(?). (Segments?) Size, 150-250 by 9-10(?) mm. Septa 6/7-10/11 thickly muscular. Atyphlosolate(?). Intes- tinal origin in xv (?). MHolandric. Seminal vesicles, small, acinous, two pairs, in ix and xu. (Ovaries fan-shaped and with numerous ege strings?) Spermathecae medium-sized, each with a vertically ellipsoidal seminal chamber sessile on duct near ampulla and opening into duct entally. GM glands small (com- posite ?), numerous, nearly covering parietes over sites of genital markings. Reproduction. Presumably sexual and biparental. Distribution. Unknown. DISCUSSION One important part of the puzzle mentioned in the title is finding a place for the new genus in the ‘‘classical’’ system of the Oligochaeta. That classification, as presented in the tenth volume of Das Tierreich (Michaelsen, 1900), was regarded as ‘‘a triumph of arrangement which brought order into confusion and constituted a remarkable advance in our understanding of the group’’ (Stephenson, 1930, p. 716), more especially be- cause of the ‘‘clear and logical division into subfamilies, on a phylogenetic basis, of the large family Megascolecidae’’ (italies not in original). Few of the various modifications proposed during the last half century (cf. Table 2 and subjoined notes) have been universally acceptable. Accordingly, the classification in the latest monograph on the order, The Oligochaeta (Stephen- 236 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY son, 1930), is recognized, for greater convenience in the ensuing discussion, aS a currently terminal stage in development of the classical system. The new genus clearly belongs in the Megascolecidae, where the male genitalia appear, from the definitions, to be of con- siderable taxonomic importance. Arrangement of the terminalia in Hxrus is ‘‘purely acanthodriline,’” i. e., there are two pairs of prostates having apertures at equators of segments xvil and xix but with the male gonoducts opening typically to the exterior on xvii midway between the other pores. Acanthodrilin termi- nalia are allowable in the Acanthodrilinae, Ocnerodrilinae, Diplocardiinae and Octochaetinae, but the genus cannot go in either of the first three subfamilies where, by definition, prostates must be tubular. Inclusion in the Octochaetinae at first does seem permissible as prostates are not mentioned in the definition. The omission probably was unintentional as all genera do have tubular prostates and Michaelsen (1900, p. 319) did specify ‘*Prostaten schlauchformig.’’ Glands of the racemose kind found in LH. wyensis are permissible in the last remaining subfamily where the prostates are ‘‘tubular or racemose (pheretima pros- tates).’’ Those organs, however, by definition, are limited to one pair ‘‘with each prostatic duct uniting with the vas deferens of the same side and opening in common with it (except in Diplotrema)’’ on xviii. The exception provides no loophole as prostates in Diplotrema are tubular and their pores, though discrete, are on xviii, close to the male apertures. Erection of a new subfamily for a single species, in spite of all those difficulties, seems unwarranted, in which case modifica- tion of the classification becomes necessary. Any ex tempore changes, solely for accommodation of one troublesome form, might later prove to be as ill advised as some of those proposed during the last forty years. Accordingly, a review of the entire classical arrangement of the Megascolecidae seems advisable and this can begin appropriately with an examination of family and _ sub- family definitions. 3 Various patterns of morphological organization long have been characterized as lumbricine, microscolecine, etc., though none are diagnostic of the named sub- families and some are common to several families. The adjective designating pattern is hereinafter distinguished from that for a subfamily by omission of the final vowel. GATES: CLASSIFICATION OF EARTHWORMS Baye Megascolecidae (cf. Stephenson, 1930, p. 818). The definition comprises eight sentences. One and parts of two others are un- qualified. ‘‘Setae sigmoid, single pointed,’’ correct, but equally applicable to all earthworm families and some Miecrodrili. ‘‘One pair of male pores,’’ not universally true throughout the family, as two pairs of male pores are present in species of Hoplochae- tella, and, in several subfamilies, parthenogenetic strains have no male pores. ‘‘One pair of ovaries in xiii.,’’ true of most fami- lies of earthworms but here inadequate. Ovaries in one genus, possibly two, are always in xii and at least one species (of Diplo- cardia) has two pairs of ovaries in xli-xiu. Other parts of the definition are qualified, by ‘‘usually,’’ ‘‘rarely,’’ exceptions or alternatives. One such statement, ‘‘Two pairs of testes in seg- ments x and xi, or one pair only, in x or xi’’ is equally applicable to the Eudrilidae, Glossoscolecidae, Lumbricidae and the micro- drilin Haplotaxidae, but is actually inadequate for the Megascole- cinae which contains one genus, possibly two, with testes in ix and x. Each of the characters mentioned in the definition is found in various other families, hence none (at least as state) is diagnostic. Acanthodrilinae (idem, p. 820). Two statements are unexcep- tional: ‘‘Meganephridial. Prostates tubular.’’ Neither is diag- nostic, tubular prostates being common in each of the sister subfamilies as also are ‘‘meganephridia.’’ The latter term, in its literally descriptive meaning, has no taxonomic value at fam- ily or subfamily levels. Redefining the word is unnecessary, as an accurately descriptive term, holonephric (or holonephridial), is available. Other characters mentioned in the definition are cualified by ‘‘mostly,’’ ‘‘more or less,’’ or are in pairs of alterna- tives, sometimes even with qualifications. None of the characters, with or without qualification, is diagnostic, being equally ap- plicable to sister subfamilies, other familes and even some Microdrili. Megaseolecinae (1dem, p. 828). Two characters here also are stated without qualification: ‘‘ Male pores on xviii. Prostates one pair.’’ Neither is diagnostic, each being applicable at least to some portion of the other subfamilies. Although male pores may sometimes be on xviii in other families they are by no means universally so in the Megascolecinae. In one genus, perhaps two, 238 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY the pores are always on xvii, in another they may be on xvii, XVlli, xix or xx; in Plutellus they may be on xviii, xix or xXx. Number of prostates in some megascolecine species is subject to individual variation. In Pheretima posthuma as many as three pairs have been found. Other statements in the definition, even with all their qualifications are inadequate. ‘‘Usually one giz- zard in front of the testis segments, sometimes two or three, exceptionally none,’’ should read somewhat as follows: Gizzard lacking, single, in a single segment or in a space formed by abor- tion of one or more septa, or double or triple, in front of testis segments, or one in front of testis segments and several more behind. Similarly, ‘‘Spermathecal pores, if present, one to seven pairs, in front of testis segments’’ should be: Athecal or thecal and then spermathecae usually in front of testis segments, rarely in or even behind those segments, pores unpaired, paired, in pairs of pairs or in pairs of groups. Octochaetinae (idem, p. 841). No statements unqualified and no character diagnostic. The prolixity of one characterization, ‘‘Axeretory system of meganephridia along with micronephridia or micronephridia alone, the latter never having the form of sacs’’ ean be avoided, with considerable gain in taxonomic accu- racy, by use of three words: Excretory system meronephric. Diplocardiinae (idem, p. 849). Two unqualified statements. Both are applicable to sister subfamilies, one to other families. No character is diagnostic. Oenerodrilinae (idem, p. 852). Three unqualified statements, two being the same as in the definition of the Diplocardiinae ; the third is ‘‘Meganephridial.’’ Characterization of the cal- eiferous section of the gut should, strictly speaking, exclude an important section of the subfamily containing two of Stephen- son’s genera. This was, however, avoided by interpreting certain microscopic spaces in the esophageal wall as vestiges of paired extramural calciferous glands. These definitions, which contain no diagnostic characters, do not define but merely list some of the more obvious or better known structural diversities of a group, and now appear to have resulted from construction of the classification ‘‘on a phylo- genetic basis.’’ Morphological changes that mean anything from the evolutionary point of view, according to Stephenson, are GATES: CLASSIFICATION OF EARTHWORMS 239 few. We know, continues the argument, which characters are primitive and which are secondary. The principal pairs or eroups of primitive-secondary characters are: One gizzard — multiple gizzards. Lumbricin setae — perichaetin setae. Holo- nephrie — meronephriec. Tubular prostates — racemose pros- tates. Acanthodrilin genitalia — microscolecin, megascolecin, balantin genitalia. Characters of lesser importance are: Cal- ciferous glands absent — ecalciferous glands present. Holandry- proandry, metandry. Hologyny-progyny, metagyny. Those are just the characters by which megascolecid genera were defined. Genera so characterized could then be arranged serially in evo- lutionary lineages of a mother-daughter-granddaughter sort. The amazing result of the filiations was a phylogenetic tree with all of the stages in the evolution of the subfamilies still available for investigation, ‘‘a living paleontology — as if students of the Equidae had all the stages in the ancestry of the horse alive be- fore them today.’’ Correctness of the filiations was ‘‘proved”’ by forms that were found to be transitional between mother and daughter genera. Since anatomical changes that amount to anything from an evolutionary point of view are so very few, convergence is fre- quent in the Oligochaeta. Perichaetin setae, for instance, have appeared in the Acanthodrilinae, Octochaetinae, Megascolecinae and in the Glossoscolecidae, meronephry in various megascolecine lines of descent as well as in the Diploecardiinae and the Octo- chaetinae, calciferous glands in all megascolecid subfamilies (even including the Acanthodrilinae) and also in the Glossosco- lecidae and Lumbricidae. Necessarily, convergent genera re- sulted. The diplocardiine Monogaster of tropical Africa was distinguished from the octochaetine Octochaetus of Pacific areas only by its distribution as the ‘‘definitions of the two genera are the same.’’ Similarly, distribution distinguished the acantho- driline Udeina* of South Africa from the megascolecine Plutellus of Pacifie areas. Nor is the convergence always as simple taxo- nomically as in those two instances, for genera may be polyphy- «The terminalia were erroneously characterized and are not megascolecin as Pickford (1987) later discovered. The two species of Udeina may have arisen independently from different species groups of the acanthodriline Parachilota, according to Pickford (1937), by changes that now appear to be relatively un- important and of dubious value for generic distinction. 240 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY letic. Megascolex, the best example, evolved from Perionyz, Spenceriella and two different sections of Notoscolex. In that case aS well as in others, anatomy permitted recognition of diverse origins but how much polyphyly would remain morpho- logically or geographically undetectable was beyond estimation. Acquisition of further data occasionally might enable resolution of a fraction of the polyphyly, but usually Stephenson seems pes- simistic about taxonomic boundary lines. They are ‘‘bound to be merely arbitrary’’ (1923, p. 193), and ‘‘sinee all such lines are arbitrary interruptions in the record of a continuous proc- ess’’ (1930, p. 833) it does not matter very much where they are drawn. Convenience, presumably of the systematist, was the eriterion employed in various cases. No comment seems necessary as to sequence in some of the pairs or groups of primitive-secondary characters. The acantho- drilin male terminalia, however, may not always have been ancestral to the other kinds and, lke some of them, may have been derived from a more ancient (and possibly less uniform ?) arrangement. The prostate sequence certainly requires consid- eration. The pheretima kind lacks a central lumen throughout and the duct may even branch outside of the gland. Supposedly intermediate sorts of racemose prostates, having a central lumen into which more or less definite lateral canals open, really are tubular. The pheretima prostate develops ontogenetically (Ste- phenson and Ram, 1919) from peritoneal proliferation and acquires an external aperture by growth outward through the body wall. Tubular prostates, on the contrary, are epidermal in- vaginations (Pickford, 1937) — as can be seen in dissections of juveniles. Gradual evolution of an ectodermal ingrowth into a mesodermal outgrowth, especially in animals with a determinate embryology, is difficult to visualize and now appears improbable. Megascolecin terminalia, those in which male gonoducts unite with the ducts of a single pair of ectodermal or mesodermal pros- tates to open externally, through one pair of male pores on xviii, accordingly provide another but hitherto unrecognized example of convergence. The ‘‘arbitrary’’ selection of such terminaha as the distinguishing character of a subfamily produced an other- wise undefinable and markedly polyphyletic group. Delimitation of other megascolecid subfamilies, it may here be noted, was no more fortunate. Selection of a form with extra- GATES: CLASSIFICATION OF EARTHWORMS 241 mural and paired ecalciferous glands in ix and x as the initial stage in oenerodriline evolution necessitated deriving the un- paired condition in Curgiona and Gordiodrilus by fusion mid- ventrally of paired sacs, and interpreting microscopic spaces in the esophageal wall of Indian genera as rudiments of retracted extramural glands. Both derivations seem improbable. The Diplocardiinae, in which the initial evolutionary stage was dupli- eation of the gizzard, has gradually disappeared (cf. note 6, Table) into the two remaining subfamilies. The more recent, the Octochaetinae, with endemic genera in New Zealand, India, Africa and the Americas, now appears (Gates, 1958b) to be polyphyletic. The older Acanthodrilinae comprises the phylo- genetic ‘‘brushwood”’ that was left over from delimitation of sister subfamilies. The group does seem to have some little com- mon anatomy, but the wide discontinuities in its distribution (New Caledonia, Australia, New Zealand, the Americas, Africa, Madagascar, Cape Verde and Subantarctie Islands), especially if earthworm evolution is limited (Stephenson, 1930) to the Tertiary and Quaternary, suggests polyphyly. Delimitation of megascolecid genera in the classical manner has proved to be even more unsatisfactory. On various occasions, since 1900, Michaelsen himself shifted generic boundaries back and forth, or abolished them. As for the Acanthodrilinae, Ben- ham could not agree with Michaelsen, and after her study of the group Pickford differed from both of them. On megascolecine demareations, Stephenson disagreed with Michaelsen, and Gates eould follow neither. In the Ocnerodrilinae, Michaelsen’s treat- ment of boundaries that left Gordiodrilus another waste-basket of phylogenetic ‘‘brushwood’’ has been questioned (Gates, 1942). More recently (Gates, 1957b) validity of some hitherto undis- puted demareations was challenged. In the Octochaetinae, sev- eral lines (Gates, various publications) had to be changed, and now (Omodeo, 1958) two genera have been resurrected (as well as a subfamily) from synonymies, where they had been buried for 60 years. Many more changes can be expected and especially —if the past provides any basis for prophecy — whenever a group is studied by another person. Accordingly, further con- sideration of individual areas of controversy may well be left to the future. 242 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY Hitherto undisputed boundaries do, however, require some comment. Andry, for instance, is not mentioned in definitions of octochaetine as well as most megascolecine and diplocardiine genera. The three standard characters had been found to have taxonomic value only at species level. All acanthodriline and most ocnerodriline genera, on the contrary, are defined by andry which also has considerable importance in the basic phylogenetic esoteries. Even supra-generie groups, in the Acanthodrilinae, are defined by andry. Nematogenia, hitherto meroandric by definition, now includes (Gates, 1957b) a holandric species. Holandry and marked hyperandry each have been found in many individuals of a species of Pheretima, holandry and anan- dry in another species of that genus. Holandry, proandry, metandry, hyperandry, anandry, all have been found in worms with genital and somatic anatomy that require the lot to be in one and the same species. Individuals, if not also species, some- times are morphologically holandrie though functionally mero- andrie. Standard characters of the male terminalia, in six non- megascolecine genera, were allowed taxonomic value only at species level. Other genera in the Acanthodrilinae, Octochaeti- nae, Diplocardiinae and Ocnerodrilinae, on the contrary, were delimited from each other by their terminaha to which again considerable importance is attached in phylogenetic esoteries. Microscolex georgianus is supposed to have acanthodrilin termi- nala, but the posterior pair of prostates fails to develop in an occasional individual which is then microscolecin. Aprostatic individuals have been found in one species of Diplocardia. Varia- tion as to number and segmental location of prostates has been found in species of several ocnerodriline genera. More data of similar sorts and from various subfamilies can be expected when efforts are made to study individual variation on a significant seale. All of the male genitalia now appear to be liable to rapid or abrupt and perhaps macromutational modification. Accordingly, status of genera presently distinguished from each other by andry and/or characters of the terminalia only is dubious. Such genera will have to be united if good evidence to the contrary — preferably from somatic anatomy — is not forthcoming. GATES: CLASSIFICATION OF EARTHWORMS 243 The least departure from the very common and ancient lumbri- cin arrangement of setae constitutes (as Stephenson maintained ) a definite and natural line of demarcation. But, of what taxo- nomic value? The change has been made in one glossoscolecid subfamily and probably on more than one occasion in each of the megascolecid subfamilies except the Ocnerodrilinae. In every instance the extra setae presumably first appeared in individuals of one or more species of a good (7. e., natural or monophyletic) genus, aS now seems to have happened in the glossoscolecine Periscoler. Somatic anatomy, in that taxon, must have obviated a generic division according to setal numbers. A natural boun- dary in an evolutionary process can then be placed arbitrarily in a classification as would have been the case if Periscolex had been split in the same manner as some of the Megascolecinae. Such arbitrariness now seems to have been responsible for part of the much discussed megascolecine polyphyly. Investigation of somatic anatomy is expected to show that Megascolex, as well as several other megascolecine taxa, are quite unnatural congeries, resolvable into morphologically homogeneous and monophyletic genera. The digestive system provided two classical sets of standard characters. One of the organs, the gizzard, is not homologous throughout the Oligochaeta. Much of the argument over that structure early in this century was about taxonomic value of presence or absence of a single esophageal gizzard. Intermediate stages, characterized as weak, feeble, rudimentary and vestigial, had been found where they were not expected. Agreement that presence or absence is unreliable as a generic character was nearly reached, but Michaelsen’s opinions changed more than once. Segmental locations were not always mentioned in descriptions of species and those recorded occasionally have been thought subsequently to be wrong. Whether such errors have been re- sponsible for any of the supposed intrageneric variation in location remains to be discovered. Transfer of a single esopha- geal gizzard into the segment next behind has been believed to be gradual, but for such change there is very little good evi- dence. That usually mentioned is insertion of a thin septum at or near the middle of a gizzard. The funnel-shaped septum has been found (in most cases that have been checked) to be inserted 244 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY behind the gizzard, though adherent to a posterior portion of it in strongly contracted specimens. With increase in number, gizzards abruptly acquired taxo- nomic importance at generic and subfamily levels. The digastrin subfamily, however, was gradually disbanded, and Michaelsen seems to have concluded that a third gizzard was inadequate to distinguish Hutrigaster from Dichogaster. Eutrigaster was re- tained by Stephenson (1930), as was Perissogaster which, by definition, differs from Digaster only in possession of a third oizzard. Presence of a third pair of spermathecae, again by definition, is all that distinguishes Didymogaster from Digaster. That difference, per se, certainly does not warrant generic dis- tinction as the spermathecal battery seems to be subject to nearly as rapid evolutionary modification as the male genitalia. All three genera, restricted to a small section of Australia, are dis- tinguished from Notoscolex, according to their definitions, only by the multiple gizzards. An extra gizzard, by itself, scarcely seems to provide more justification for generic distinction than does the presence of an extra pair of setae. Considerable intra- specific variation in location and number of gizzards has been found in the moniligastrinae. Assumptions as to specific and generic uniformity of those characters in the Megascolecidae usually are unsupported by data obtained from more than a very few specimens. Pending acquisition of much more infor- mation, the taxonomic value of gizzard number and location (the only macroscopic characters the organ can supply) remains uncertain. Caleiferous glands were allotted a wide range of taxonomic values in the classical system. The organs, though sometimes present, are not even mentioned in definitions of Acanthodriline and Megascolecine genera. In other subfamilies the characters used in definitions are mainly segmental location and number. The latter often is erroneous. Dichogaster, by definition, has three pairs of calciferous glands. Yet, a group of common and widely peregrine species (Gates, 1942, 1958b) really has only one pair. Howascolex became even more of a congeries, after 1930, and by definition now has ‘‘Caleiferous glands either as mere swellings of the oesophagus in segm. 14, with or without a shght development of the same kind in the neighbouring segments ; or GATES: CLASSIFICATION OF EARTHWORMS 245 well developed calciferous glands in segm. 14 and 15, or in segms. 8-11.’’ Diplocardia, by definition, lacks calciferous glands, though a species was known to have one of a highly specialized sort. The North American genus provides (apparently within the limits of a monophyletic taxon) a beautiful series of evolu- tionary stages (Smith, 1924), beginning with absence of eal- ciferous lamellae and ending with an intramural gland very similar to that of the better known Lumbricidae. The standard characters of the remaining somatic sequence are meganephridial and micronephridial. Meganephridia may be holonephric, meronephric, exonephric, enteronephric, vesicu- late or avesiculate, with or without a bladder-lke caecum, of various sizes and occasionally smaller than some micronephridia, one to several pairs per segment or more numerous. Micro- nephridia may be stomate, astomate, exonephric, enteronephric with respect to pharynx or to intestine, vesiculate or avesiculate, several pairs per segment up to ‘‘forests’’ of hundreds. The standard characters obviously have little meaning in an im- portant sector of somatic structure. Holonephriec and mero- nephric have relegated the older terms to infrequent but more precise usage. They do characterize groups of genera or larger units more accurately than their predecessors, but even so they can have very little of the taxonomic value allowed the older pair in the classical system. The first appearance of meronephry undoubtedly constitutes a natural and definite line of division in organ evolution but initially can provide no more justification for generic separation than addition of a few setae. Increased knowledge of somatic anatomy can be expected to reveal, in good genera, earlier stages of meronephry than have been recognized hitherto. More ancient meronephric systems, as an interesting and im- portant series of contributions by Bahl (1919-1945) shows, are organized in a variety of dissimilar ways. A somewhat different sort of organization is less satisfactorily described (Gates, 1943) because of poor preservation. Existence of other kinds of meronephric systems is indicated by the literature. For some time it has been quite obvious that dissimilarities in structure, as well presumably as in embryological development and phylo- genetic evolution, are such as to require precise characterization at generic level. 246 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY Ever since Savigny (1826) amazed his colleagues by demon- strating the existence, in Paris, of many more than one species of earthworm, taxonomic descriptions have been mainly con- cerned with organs visible to the unaided eye or, in smaller forms, through a hand lens. A few organs, such as prostomium, setae, gizzard and segmentally paired holomeganephridia, doubt- less were the structures seen in the juveniles, that often must have been examined. With increased knowledge, inability to find ‘‘segmental organs’’ became acceptable proof for existence of a micronephridial excretory system. Mature worms, however, provided the taxonomist a clitellum of diverse lengths, cireum- ferential extent and location, an array of other epidermal modi- fications collectively designated as genital markings, genital pores in various locations, a battery of spermathecae, another of seminal vesicles permitting deduction of andry when male gonads were unrecognizable. Mature exotic material that in- creasingly became available, provided prostates of various sorts, as well as a bewildering variety in other genitalia. On the con- trary, guts, blood vessels and nervous system may well have seemed to be tediously uniform. The diversity of genital struc- ture was still far from complete elucidation during the period when Michaelsen (1900) was completing his masterpiece. The conclusion that ‘‘The sexual organs are the most important of all for systematic purposes’’ (Stephenson, 1923, p. 7) may have seemed unavoidable. Peremptory denial of evolutionary value (apparently equated in a phylogenetic classification with taxonomic value) to so much of the somatic anatomy, is unlikely to have stimulated investigation of it. Interesting characteristics of the vascular system in little known species were observed by Benham but being of ‘‘no taxonomic importance’’ they were not recorded. Inclusion in a taxonomic contribution of information about blood vessels and hearts in several genera of a little known family, even in 1930 seemed so unusual that it was mentioned in Stephenson’s monograph. Study of the octochaetine Eutyphoeus has enabled redefinition of the genus as given in the left column of Table 1. On the right, for comparison, is the classical definition (Stephenson, 1930). Descriptive characterizations, often in the past or still included in generic definitions, are listed after the distribution. GATES: CLASSIFICATION OF EARTHWORMS 247 TABLE 1 EUTYPHOEUS As now defined As previously defined Biprostatic, pores in region of Sexual apparatus purely micro- AB, near eq/xvii. Male pores minute, near but behind prostatice pores. Bithecal, pores superficial, never minute, at 7/8. Setae paired, arrangement lumbri- ein, Clitellum annular, on xXiv-xvi, in- tersegmental furrows obliterated, dorsal pores oceluded, setae retained. Septa 4/5-5/6 with muscular thick- ening, 6/7-7/8 aborted, 8/9-10/11 thickened, crowded together behind their normal locations, 11/12 ap- proximated to 10/11. An esophageal gizzard belonging to vi in space between 5/6 and 8/9. Caleiferous glands intramural, longi- tudinally hemi-ellipsoidal with flat faces mesially, numerous transverse vertical partitions and interlamellar spaces directly communicating dor- sally with the esophageal lumen here T-shaped in cross-section, 1 pair in xu. Intestinal origin in xv. Typhlo- sole terminating with a series of doubly-paired supra - intestinal glands. Unpaired, anteriorly di- rected, small, midventral caeca one each in a number of consecutive seg- ments in front of supra-intestinal glands. Dorsal blood single. No subneural. Lateroparietal trunks from posterior end of body pass to hind ends of caleiferous glands. Extra-esophageal trunks, median to hearts, pass to front of calciferous glands. Hearts four pairs, in x-xiii, last three pairs latero-esophageal. vessel scolecine (conjoined male and pro- static pores on xvii). Spermathecal pores, one pair, at 7/8. Setal arrangement lumbricin. An enlarged esophageal gizzard in a space formed by fusion of several segments. A pair of calciferous glands em- bedded in the esophageal wall in xii. 248 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY As now defined Exeretory system meronephriec, all nephridia small, numerous astomate nephridia of iii pharyngonephric and apparently in a_ circumferential parietal band but attached to a tightly-zig-zagged cord, remainder of system exonephrie and comprising astomate biramous nephridia which are numerous in next few segments but behind elitellum are in longi- tudinal ranks, the medianmost nephridium of each side behind supra-intestinal glands somewhat enlarged, saccular and with presep- tal funnel. Terminal portion of male gonoduct modified to function as a_bulbus ejaculatorius. Spermathecal diverticula open into ental end of short duct. Metagynous. Distribution: Burma, from Tenas- serim division and western margin of Shan plateau into the Gangetic Plain and through the Himalayas to beyond Nepal. Segments more than 150. Unpigmented or pigmented and then with dorsum brown, oceasion- ally green but with no corresponding pigment recognizable. Ventral setae of xvii lacking or penial. Male and prostatic pores as well as apertures of penisetal follicles in two slight fissures, each with a super- ficial porophore or more or _ less deeply invaginate, vestibula paired or unpaired and median, sometimes with protrusible penes. Lateral intestinal caeca lacking, rudimentary and/or sporadic, or one pair. As previously defined Purely micronephridial. India, Plain. especially the Gangetic GATES: CLASSIFICATION OF EARTHWORMS 249 As now defined As previously defined Holandrie or meroandric. Seminal Holandrie or metandrie. vesicles in ix and/or xii. Coelomie cavity of xi gradually reduced to an annular, then a U-shaped and even- tually a subesophageal testis sac. Two female pores, or right oviduct functionless or atrophied. The revised definition contains no alternatives and no excep- tions. Absence of variation with regard to the characters men- tioned, in normal specimens, was determined (Gates, in press) from external examination and dissection of hundreds of speci- mens. The reproductive system, not excluding the female gonoducts, has undergone considerable modification during in- trageneric evolution. The genitalia, from an evolutionary point of view, are not conservative. Much somatic anatomy, on the contrary, has remained uniform during a period in which the genus was spreading through rain-forests and semi-deserts, from tropical lowlands to Himalayan heights. Eudichogaster, the parent genus of Lutyphocus in the classical system, when redefined (Gates, 1939, 195?) with reference to more of somatic anatomy, cannot be ancestral. Phylogenetic filiation, as in the ease of Bahlia which is more closely related to Eutyphoeus, is possible only through common descent from some form that is no longer extant (Gates, In press). Scolioscolides, at present known only from a single species originally placed in the Megascolecinae, is (Gates, 1937) still more closely related to Eutyphoeus by all of its anatomy than is either of the other genera. The ‘‘living paleontology’’ of the Indian Octochaetinae has vanished. That portion of the gut in the nine to fifteen segments between gizzard and esophageal valve of Indian octochaetines has pro- vided a striking demonstration of the importance of previously neglected organization. Using only characters from such a short region, a key was constructed (Gates, 1958b) that identifies eenera (except Scolioscolides) and simultaneously indicates those likely to need drastic revision. The key is based on macro- scopic anatomy. Microscopie structure of the caleiferous glands undoubtedly will provide still other defining characters. In a 250 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY short terminal portion of holonephridia, Pickford (1937) found macroscopically recognizable characters that could be used in defining Acanthodriline genera. The classical Megascolides has marked discontinuities in its distribution: Peninsular India, the eastern Himalayas, Austra- ha, Tasmania, and Oregon-Washington in North America. M. bergtheili Michaelsen 1907, of the Himalayas, is so nearly identi- cal with the indubitably octochaetine and specialized Eutyphoeus that little more than the megascolecin terminalia is available to warrant even subgeneric separation. With that discovery (Gates, 1937), the single morphological distinction between the Octochaetinae and the Megascolecinae became taxonomically null and void. M. prashadi Stephenson 1920 and annandalei Stephenson 1921 are barely if at all distinguishable from the oetochaetine EHudichogaster barodensis Stephenson 1914 which had to be separated off from the rest of the genus (Gates, 1939a) beeause of its somatic anatomy. M. cochinensis Michaelsen 1910, M. duodecimalis and pilatus Stephenson 1915, M. chengannures Aiyer 1929, have been transferred (Gates, 1940) also because of somatic anatomy to the octochaetine Travoscolides. M. an- trophyes Stephenson 1924, known only from the holotype, un- like the other Indian species does seem to be megascolecine. Relationships, insofar as they could be determined in the circum- stances, are with a local group of genera. With those discoveries Megascolides disappeared from the Orient. The classical Woodwardiella of the Megascolecinae also has discontinuities in its distribution : India, Ceylon, Java, Australia. The Java record is due to transportation of a species from the west. Oriental species, because of somatic structure, had to be transferred to other genera including two, Nellogaster and Nelloscolex, that are not classical. Another genus, shared be- tween Australia and South India, that is terminal lke Wood- wardiclla in its own line of megascolecine evolution now (Gates, 1958b) has lost some of its morphological and phylogenetic heterogeneity in the same way. Megascolexr is penultimate in its line of evolution but has, like its ancestor Notoscolex, en- demie species only in South India-Ceylon and Australia, with or without New Zealand. A classical genus which is only one step removed from the parent of all octochaetines had endemic GATES: CLASSIFICATION OF EARTHWORMS 251 species only in New Zealand and peninsular India. To account for those distributions Michaelsen first postulated separate land bridges, from Australia and from New Zealand to India, and later invoked Wegenerian association of continental land masses. Independent origin of identical genera since the Cretaceous was Stephenson’s explanation. Octochaetus was subsequently split into two subgenera, that were later raised to generic status, but the neoclassical Octochaetoides probably still requires (Gates, 1958b) subdivision. Resolution of the notorious and somatically indicated polyphyly of Notoscolexr and Megascolex can be ex- pected with confidence. In the Megascolecidae few genera will be retained, just as in the classical system; one such may be Diplo- cardia; another —the largest of all earthworm genera — is Pheretima. Yet, even in Pheretima, as already suggested, all genital organs except ovaries, oviduets and cocoon-secreting elitellum can be eliminated within a species, while other strue- ture remains constant. The wide oceanic discontinuities in distribution, that are so common in the Megascolecidae, characterize several genera of other earthworm families. The Moniligastridae has no such dis- continuities, but genera as well as species are distinguished almost entirely by genital characters. Two genera of the little studied Glossoscolecidae provide some evidence (Gates, 1958a) of rapid evolutionary modification in genitalia. In the Lumbri- cidae, intermediate forms between most of the classical genera long have been known. A recent examination of a couple hun- dred specimens of an infrequently seen form (Gates, 1957a, p. 13) provided noteworthy instances of individual variation only with respect to those genital characters most used for defining and distinguishing species. In another lumbricid, all genital organs, except as in the above-mentioned pheretimas, have been eliminated, while other anatomical features remained constant. Most lumbricid genera appear (Gates, 1956a, p. 30) ‘‘to be only congeries of species associated because of relatively un- important or superficial convergences.’’ So little is known about so much of the taxonomically important structure in sO many species, that ex tempore transfers from one polyphyletic genus to another (Gates, 1956a, p. 26) seem inadvisable. The somatic anatomy of various unmentioned subfamilies and families, as Die, BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY well as of the Glossoscolecidae, certainly is much less known than that of the Lumbricidae. There is then little to indicate that other earthworms may have been classified, except perhaps by accident, more naturally than the megascolecids. The revision that is needed, through- out the megadrilous portion of the Oligochaeta, may have to be drastie. Since 1900, one family and two subfamilies (see explanatory notes 7-9, following Table 2), several genera and many species have been erected. Of the latter, the number in Pheretima and Dichogaster already had been doubled (to ca. 300 and 160) by 1930. Many species still are known only from descriptions of a very short series, or of a single type, that sometimes has been aberrant. Little information about reproduction (whether sexual or parthenogenetic), individual and geographic variation has been recorded. Somatic anatomy, rather generally, is too inadequately characterized to permit grouping species according to over-all similarity. Until that information is available polyphy- letic taxa cannot be made monophyletic with certainty by any reshufflings based on the literature. Earthworms have been systematically collected in few areas, even in those immediately surrounding or easily accessible from museum and university centers. Such material as still may be available’ in institutions usually is the casual spoil of other activities, almost always hastily preserved in the field, often more or less macerated, but if not, probably deteriorating slowly be- cause methods of retaining specimens in good condition are unknown. Even the best of field-preserved material cannot be expected to provide all of the information that is needed, espe- cially with regard to vascular and excretory systems. Decades must pass, so far as can be estimated from the rate of increase in knowledge during the last century, before data as to neglected portions of digestive and other somatic systems will be sufficient to permit grouping species and genera according to over-all similarity. The problem to which we now return is that of finding a place for a new genus of unknown affinities in an obviously obsolescent 5 A list of extant types could not be provided by Michaelsen’s own institution as late as 1958. GATES: CLASSIFICATION OF EARTHWORMS 253 system, at a time when relationships of earthworms cannot be determined from the literature or from extant collections. In such circumstances changes in the system clearly should be mini- mal to avoid needless extensions in the future of already ecompli- eated synonymies. Racemose prostates of the pheretima kind are present in genera that belong in a region extending through India, China, Malaysia and Australia, perhaps with some intrusion into New Zealand that is not due to human introductions. As wide oceanic intervals are lacking in that region it is possible that all of the forms under consideration have had a common origin. Accordingly, Exrus is assumed to be from the same Australasian region and to be- long in the Megascolecinae, which alone contains genera having truly racemose prostates without a central canal. The subfamily, as pointed out long ago, already was undefinable morphologically. It can be defined, by its prostates, if forms with tubular glands, regardless of presence or absence of lateral branches from the axial lumen, are excluded. Mesodermal origin of the prostate, in a group where diagnostic characters are few, appears to be an evolutionary innovation of sufficient importance to justify more than subfamily status. The Megascolecinae of Stephenson (1930), as now restricted, accordingly becomes a family in par- tial agreement with Michaelsen’s (1921, 1929) later proposals. Genera now excluded from the Megascolecidae are distributed among the other subfamilies, in accordance with precedents set when Stephenson, Michaelsen and Pickford disbanded the Trigastrinae and the Diplocardiinae. Thus, genera with holo- nephri¢ excretory systems throughout go into the Acanthodrili- nae, and meronephriec genera go into the Octochaetinae. Each of those two groups of genera seems to be entitled to the same rank as the Oenerodrilidae, which was separated off some years ago (Gates, 1939b). Such rearrangements, in the neoclassical manner, satisfy Stephenson’s eriterion of convenience. They permit reference to or discussion of groups, having a limited degree of common morphology, independently of the highly subjective phylogenetic esotery on which the classical system is really based. The synopsis below summarizes the proposed changes along with brief definitions and generic lists. 254 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY Ay. Prostates racemose, of pheretima type, without central canal and presumably of mesodermal origin. MEGASCOLECIDAE. Comprises the following genera, some of them perhaps in part only, Lampito, Pheretima, Perionyx, Plionogaster, Woodwardiella, Comarodrilus, Notoscolex, Megascolex, Digaster, Perissogaster, Didymogaster, Nel- logaster, Tonoscolex, Nelloscolex and Exxus. Ao. Prostates tubular, with central canal, of ectodermal origin. B,. Pre-intestinal region short with latero-esophageal hearts confined to x-xi and intestinal origin in or (usually) anterior to xiv. (Setal arrangement lumbricin. Calciferous glands or epithelial- lined diverticular spaces in thickened esophageal wall, in ix-x. Exeretory system holonephric.) OCNERODRILIDAE,. Genera as in Stephenson, 1930, except for Aphanascus which was united with Malabaria (Gates, 1942), Kerria and Curgia which are now known as Eukerria and Curgiona, and in addition Deccania. Bo. Pre-intestinal region longer, with intestinal origin in or behind xv and with hearts not confined to x-xi or their homoeotie equiva- lents. C,. Exeretory system holonephric. ACANTHODRILIDAE. Genera as in Stephenson, 19380, with addition of Hodrilus, Parachilota, Diplocardia, Zapotecia and from the megascole- cinae Diplotrema, Plutellus, Pontodrilus, Diporochaeta. Co. Excretory system meronephric. OCTOCHAETIDAE. Gen- era as in Stephenson, 1930, with addition from the Megascolecinae of Spenceriella, Megascolides, as well as the neoclassical Wegeneriona, Neogaster, also Scolioscolides, Lennogaster, Barogaster, Rillogaster, Priodochaeta, Prio- doscolex, Travoscolides, and Celeriella. The tubular prostates do seem to suggest a closer affinity to each other, than to the Megascolecidae, of the familes Oecnero- drilidae, Acanthodrilidae and Octochaetidae, which were all included in his Acanthodrilidae by Michaelsen (1921, 1929). Any attempt at formal taxonomic indication of the relationship, in the present circumstances, scarcely seems worth while. Inter- position of suborder and series between order and families, as in Michaelsen’s later schemes, still seems, in agreement with Stephenson (1930, p. 719), to be unwarranted by our present knowledge. Ken) 9! EARTHWORMS OF ATION Y ) CLASSIFIC oeprxeyo[de yy (¢) eeprproanypTy (F) SBprlayseVsT|Tuo jy aeprydours.iedg IVPT[LIPOLL) IVpIJOVIOLITIPY AV PLI4S¥5OULLO FL OB PT99[ODSOSSO[4) (1) eBploraquiy (€) 9Bpr[LIpo19ue9 IVPTJIVYVOJIO) BPI poyyUB Vy JVPLIB[OISBBITV oVPITII pny posodoig Uld.19 FJ oeprixeyo[dey eePIpromnqty (g) eepttzpouasusg IVUL[LIpOLIUIG, IBUTJOBYIOJIO dsVUl[LLpoyyueoy dVULDI[OISLBITNV (%) 9BplLsa[Oosve sayy pLOFyId oVprixeyo[de yy] sVPIploin LV sVUT[LIpoussuAg IVUTISVSI[IUO PL IVPTIYSVSIPLMOPY avurrydoursiedg OVUL[LI POTD AVULJOVIPPOLIIPL IVULIJSVSBOWLLO TT IVULII[OISOSSO[H aBpWo[0dsOsso[4 aVepHiiquny ICUI[LIpOLIIG, dVUTJOVYVOJIO (9) evurrpivooldiq IVUT[LIpoyUBDV ICVULDI[ODSBSII AVPLI[OISBSIWW SVPTIIPNOL OG6l uosuaydea4g ovprxeyoldepyy SVPIproini Ty dVUT[LIpoussuség OVULIISEST[IMO JL OBPIIISVST[IUO]\ ovurrydouvsaeds dVUT[LA POLL IVUTJOVYIOAOL] IVUTIISVSOULIO FT ABUTII[OISOSSOTL) avulotiqun'y] aVplolaquin'y] IVUT[LIpOLsudGQ IVULJILIPIOJIO dVULIPeBdO [At dVUI[LIpoyjuvoy IBUTII[ODSUHIT ovVUl[LIpagy IVPLOO[OSvHIIV EG6L wosueydeyzg *soqou L10}VULLAXd 03 IojJol (6)-(T) dB ptqdALOIIY SU PIPLoany Vi adVpT[LIpoussurcg SBPIIISESTIUO PY vUlpILIOOIY avprytydouvsiedg SVPT[LIpOtry) IV PTOVYIOIITY IVPLI4SVSOULIO FT aBPWa[OdSOsso[*) OVPLOlq way RULOLIG WT IBUT[LIPO1IUIO, aVUTJIVYIOJPIO, avulrpzed0[diq (Q) eVULI4SvVStLy, IBUT[LIPOYJUBIV aVpl[LIpoyyuBoy IBPTNI[OISLSI]Y oVplpLipagy VUTII[OISB.HIIY ([) BjJovIyooOsT[O-09N Lo6L WaspavyoT BJOBYPOSI[(Y) oY} JO SUOTJVOYISSL[D JULIA UL SolpIUIVZQNS PUB SOT[IMURF WIOMTILEG avprxeyoldeyy SEP LprOT OT hy, IVPLIYSV SIMO OVUr[IIpoOrry IVUTPIVIPOIONN IVULIYSVAOULLO FT IVULII[ONSOSSOTL) IBPLIILODSOSSO[L) OVPIOLIq uN] IVUL[LIpOLIWO, IVULJOVIPIOJIO ovUILpIvdO [AIC IVULIFSVSLL, dVUT[LIpOyJUBIY IVULIO[OISVBIIY ovUT[TIpuy IVPTII[ODSBH JW 0061 Uas [IBY TY 'g alavy, 256 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY EXPLANATORY NOTES (1) Suborders of the 1921 scheme, the Archioligochaeta and Neo-oligochaeta, were abandoned in 1929. The suborders then proposed are three, Oligochaeta plesiopora, prosopora and opis- thopora. (2) Pickford, 1937. (3) Gates, 1939, 1942. (4) Gates, 1945. In this article, an editor made the author say (p. 394) ‘‘Hearts of vi and vii connect the dorsal and ventral trunks to a longitudinal vessel that appears to be an extra- esophageal,’’ which is incorrect and unfortunately was repeated (Pickford, 1945) in a formal redefinition of the Syngenodrilidae. The ‘‘hearts,’’ so far as could be determined from the available material, connect the dorsal and ventral trunks only. Another editorial change was deletion of a statement to the effect that the ‘‘hearts’’ are median to the extra-esophageals. Those trunks, so far as is known and except in Syngenodrilus and the Monili- gastridae, are median to hearts and segmental loops. The pair of characters, lateral to or median to the hearts and segmental loops, eventually will prove to be of considerable taxonomic importance. (5) Pickford, 1945. Also cf. Gates, 1945. Recognition of a family, or even a subfamily, for Syngenodrilus alone, at present scarcely seems warranted from, quoting Stephenson, the point of view of convenience or by our knowledge of relationships. (6) The Trigastrinae of 1900 and 1921 disappeared when Stephenson transferred the Indian Hudichogaster (which had been split off from the Afro-American Dichogaster) to the Octo- chaetinae, Trigaster, Eutrigaster, Dichogaster and Monogaster to the Diplocardiinae. Michaelsen (1933) suggested transfer from the Diplocardiinae to the Octochaetinae of the meronephric Trigaster, Dichogaster (presumably including Eutrigaster) and Monogaster. Genera still left in the Diplocardiinae, the holoneph- rie Diplocardia and Zapotecia, were placed by Pickford (1937) in the Acanthodrilinae. (7) The Criodrilinae of 1900 became monogeneric by erection of a family for Sparganophilus, transfer to the Microchaetinae of the African Alma as well as the American Drilocrius (split off from Criodrilus). GATES: CLASSIFICATION OF EARTHWORMS 257 Pop (1949) included Criodrilus in the Lumbricidae without subfamily divisions. Omodeo (1956) exeluded Criodrilus and recognizes two subfamilies, Lumbricinae and Hiseninae. (8) The monospecific Hippoperidae was erected (Taylor, 1949) for a eudrilid supposedly distinguished from the rest of the family by presence of a second pair of male pores. (9) Megascolecid subfamilies are reduced to two in a publica- tion (Lee, 1959) received after the manuseript of this contribu- tion had been typed. Acanthodrilinae. ‘‘One pair of prostatic pores on xvi (rarely ) or XvVll or Xix, or two pairs on xvii and xix (rarely the two pairs may be further back) ; one pair of male pores, usually on xviii, sometimes on neighbouring segments, sometimes combined with a pair of prostatic pores (in which case, never on xvill) ; pros- tates with unbranched central canal.’’ Comprising Acanthodri- lus, Microscolex, Rhododrilus, Dinodriloides, Perieodrilus, Mao- ridrilus, Neodrilus, Plagiochaeta, Chilota, Yagansia, Udeina, Hodrilus, the diploeardiine Diplocardia, Zapotecia, Trigaster, EKutrigaster, Dichogaster, Monogaster, the octochaetine Howasco- lex, Octochaetus (ineluding Octochactoides), Dinodrilus, Hoplo- chaetina, Ramiella, Eudichogaster, EUTYPHOEUS, Hoplochae- tella, the neoclassical Leucodrilus, Decachaetus, Eudinodriloides, Sylvodrilus and Neochaeta, the Oenerodriline Maheina, Curgiona, Malabaria, Paulistus, Eukerria, Kerriona, Haplodrilus, Ocnero- drilus, Pygmasodrilus, Nematogenia, as well as WNelloscolex, Tonoscolex, Rillogaster and Lennogaster. Megascolecinae. ‘‘One pair of prostatic pores and one pair of male pores on xvili (Diplotrema only) or one pair of combined male and prostatic pores on xvill; prostates with unbranched or branched central canal.’’ Comprising Diplotrema, Plutellus, Pontodrilus, Woodwardiella, Comarodrilus, Megascolides, Spen- ceriella, Notoscolex, Megascolex, Pheretima, Plionogaster, Digas- ter, Perissogaster, Didymogaster, Diporochacta, Perionyx, the oenerodriline Quechua (should be Quechwona), as well as Baro- gaster, Priodochaeta, Priodoscolex, Travoscolides and SCOLIO- SCOLIDES. The oenerodriline Gordiodrilus and_ several neoclassical genera were not placed. These changes, like others in the neoclassical manner, are not based on any substantial increase in knowledge of somatic anat- 258 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY omy. Similarity, for each subfamily, is restricted to presence or absence of united male and prostatic pores in xviii only (megascolecin terminalia). Elucidation of the relationship between Hutyphoeus and Scolioscolides showed conclusively that the two subfamilies cannot be so distinguished. Further proof probably will be provided by the oenerodriline genus (Gordio- drilus) that could not be placed in either of the revised units. Branching of a central prostatic canal was recorded in some species of Diplocardia more than sixty years ago. A central canal is lacking in the prostates of Nelloscolex and Tonoscolex. Quechuona has a short pre-intestinal region (with hearts in x-xi only) such as is characteristic of nearly all oenerodriles. The megascolecin male terminalia probably are present in Gordiodri- lus, which clearly belongs in the same family with Quechuona and other oenerodriles. SUMMARY In the ‘‘classieal’’ system of the Oligochaeta, species are mu- sem taxa, 1. e., based on resemblance in a few macroscopically recognizable characters to a type specimen that sometimes was quite abnormal. Information as to individual and geographical variation usually is lacking. Genera, defined by a very few ‘‘key’’ characters and often obviously polyphyletic, are arranged in larger units according to a phylogenetic esotery. Definitions of subfamilies and families are statements of range of variation in certain characters most of which are used to define genera. The system is artificial and obsolescent. Revision of genera on a basis of over-all similarity of species 1s impossible because of absence in the literature of information about much somatic anatomy. Material needed for adequate characterization of the species (many without types) is unavailable in institutions and is unlikely to be secured for many years to come. To accommo- date Hxrrus wyensis, the Megascolecidae of the latest monograph on the Oligochaeta is redefined to include only genera having racemose prostates of the pheretima sort. Excluded species with holonephric and meronephric excretory systems respectively are transferred to the Acanthodrilidae and Octochaetidae. GATES: CLASSIFICATION OF EARTHWORMS 259 REFERENCES BauL, K. N. LOMDS 1922. 1924. 1926. 1941. 1942, 1945. 1946. GATES, G. 1937. 1938. On a new type of nephridia found in Indian earthworms of the genus Pheretima. Quart. Jour. Micros. Sei., 64: 67-119. On the development of the ‘‘enteronephric’’ type of nephridial system found in Indian earthworms of the genus Pheretima. Tbid., 66: 49-103. On the oceurrence of the ‘‘enteronephrie’’ type of nephridial system in earthworms of the genus Lampito. Ibid., 68: 67-99. The enteronephrie system in Woodwardia with remarks on the nephridia of Lampito dubius. Ibid., 70: 113-134. The enteronephric type of nephridial system in the genus Tonoscolex (Gates). Ibid., 82: 443-466. Studies on the structure, development and physiology of the nephridia of Oligochaeta. Part I. General introduction, and the nephridia of the subfamily Octochaetinae. Part II. Multiple funnels of the nephridia. Jbid., 83: 423-457. III. The branch- ing and division of nephridia and Eisen’s so-called ‘‘safety valves’? in Pontoscolex. IV. The enteronephrie system in Megascolex cochinensis, with remarks on vestigial nephridia. Ibid., 84: 1-34. Part V. The enteronephric system in Megascolex ceylonicus and M. sarasinorum, with remarks on the phagocytic organs in Megascolex templetonianus. Ibid., 85: 177-190. Part VIII. The enteronephriec type of nephridial system in earthworms belonging to three species of Megascolex Templeton and three species of Travoscolides Gates (Megascolides McCoy ). Ibid., 87: 45-60 E. Indian earthworms. II. Scolioscolides. Ree. Indian Mus., 39: 305-310. Indian earthworms. III. The genus Hutyphoeus. Ibid., 40: 39- 119. IV. The genus Lampito Kinberg. V. Nellogaster, gen. nov., with a note on Indian species of Woodwardiella, Ibid., 40: 403-429. 1939a. Indian earthworms. VI. Nelloscolex, gen. nov. Ibid., 41: 37-44. VII. Contribution to a revision of the genus Hudichogaster. Ibid., 41: 151-218. 1939b. Thai earthworms. Jour. Thailand Res. Soc. Nat. Hist. Suppl., 1940. 12: 65-114. Indian earthworms. VIII. Priodochaeta, gen. nov. IX. Prio- doscolex, gen. nov. X. Contribution to a revision of the Indian section of the genus Megascolides. XI. Travoscolides, gen. nov. Ree. Indian Mus., 42: 115-143. 260 1945. 1956a. 1956b. 1957a. 1957b. 1958a. 1958b. LEE, Ke. 1959. BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY Notes on various peregrine earthworms. Bull. Mus. Comp. Zool. Harvard College, 89: 63-144. On some American and Oriental earthworms. Ohio Jour. Sci., 43: 87-116. On the Oligochaete genus Syngenodrilus and its taxonomic rela- tionships. Jour. Washington Acad. Scei., 35: 393-396. Notes on American earthworms of the family Lumbricidae. ITI-IV. Bull. Mus. Comp. Zool. Harvard College, 115: 1-46. Reproductive organ polymorphism in earthworms of the oriental megascolecine genus Pheretima Kinberg 1867. Evolution, 10: 213-227. Contributions to a revision of the earthworm family Lumbrici- dae. I. Allolobophora limicola. Breviora, Mus. Comp. Zool., 81: 1-14. Contribution to a revision of the earthworm family Oecnerodrili- dae. The genus Nematogenia. Bull. Mus. Comp. Zool. Harvard College, 117: 427-445 On a hologynous species of the earthworm genus Diplocardia, with comments on oligochaete hologyny and ‘‘ consecutive hermaphroditism.’’ Amer. Mus. Nat. Hist. Novitates, no. 1886; 1-9. On Burmese earthworms of the megascolecid subfamily Octo- chaetinae. Ann. Mag. Nat. Hist., (13) 1: 609-624. On a species of the Indian earthworm genus Hudichogaster. (In Press. ) . The earthworms of Burma. VII. The genus Hutyphoeus, with notes on several Indian species. Rec. Indian Mus., 53: 93-222. (MS submitted in 1954. Page proof bearing publication date of 1955 corrected in 1956.) The earthworm fauna of New Zealand. New Zealand Dept. Sci. Indust. Res. Bull., 130: 1-486. MICHAELSEN, W. 1900. 1921. 1929. 1933. Oligochaeta. Das Tierreich, vol. 10, i-xxix, 1-575. Zur Stammesgeschichte und Systematik der Oligochaten, ins- besondere der Lumbriculiden. Arch. Naturgesch., (A) 86 (8): 130-141. Zur Stammesgeschichte der Oligochaten. Zeitschr. Wiss. Zool., 134: 693-716. Die Oligochitenfauna Surinames. Mit Hrorterung der ver- wandschaftlichen und geographischen Beziehungen der Octo- chiitinen. Tijdschr. Nederlandsch Dierk. Ver., (3) 3: 112-131. OmopEo, P. 1956. 1958. GATES: CLASSIFICATION OF EARTHWORMS 261 Contributo alla revisione dei Lumbricidae. Arch. Zool. Italiano, Al: 129-212. La reserve naturelle integrale du Mont Nimba. I. Oligochetes. Mem. Inst. Afrique Noire, 53: 9-109. PICKFORD, G. E. 1937. 1945. Pops Vi. 1950. SAVIGNY, J. 1826. Smiru, F. 1924. A monograph of the Acanthodriline earthworms of South Africa. Cambridge, 1-612. Additional observations on the oligochaete genus Syngenodrilus. Jour. Washington Acad, Sci., 35: 397-399. Lumbricidele din Romania. An. Acad. Romane, Sect. Sti. Geol. Geogr. Biol., (A) 1 (9): 1-129. C. In: Cuvier, G. Analyse des travaux de 1’Academie royale des Sciences, pendant l’annee 1821. Partie physique. Mem. Acad. Roy. Sci. Inst. France, 5: 153-203. The caleiferous glands of Lumbricidae and Diplocardia. Illinois Biol. Mon., 9: 1-76. STEPHENSON, J. 1921. Contributions to the morphology, classification and zoogeography of the Indian Oligochaeta. Proc. Zool. Soc. London, 1921: 103- 142. Oligochaeta. In: Fauna of British India including Ceylon and Burma, London, xxiv +518 pp. The Oligochaeta. Oxford, xiii +978 pp. STEPHENSON, J. and H. Ram OM: TAYLOR, A. 1949, The prostate glands of the earthworms of the family Megascole- cidae. Trans. Roy. Soc. Edinburgh, 52: 435-453. G. A West African earthworm, Hippopera nigeriae, belonging to a new family Hippoperidae. Proe. Zool. Soc. London, 119: 703-710. a wale AN mage aa) Lone sori. "| 7 "hd as 4 Pon ila adalat a ie owes deal aa Ge ipiaoinmabias oc or). a Mae Me Ty, Git 4, qyrrcd wh to rit Toe een. / mee re jee’ yih te hit ale gi AS a 8 ba : : ; de f An . OE yhit " ° iy bie t 7 7 — y ¥ a : ~ a Tp i . ( sp mr Ls igvesa i 2 ata rh ae Wen! ti as roe ve ee Ty hans ate om iia i rr’, 5 nc nL Aligny aed , ae Ail 8 a > ah dined Fy! wie , uw re ONE i" a: AM i Fi "nad , ie el ae ip Afi ¥ = 1 i alg ft AT iri . one - iw ee er | ri i ig Bulletin of the Museum of Comparative Zoology AGT H-ASR V ALRID:- CO bash iG Vol. 121, No. 7 BATHONIAN FORAMINIFERA OF ENGLAND By RicHArpD CIFELLI WitH SEVEN PLATES CAMBRIDGE, MASS., U.S.A. PRINTED FOR THE MUSEUM SEPTEMBER, 1959 PUBLICATIONS ISSUED BY OR IN CONNECTION WITH THE MUSEUM OF COMPARATIVE ZOOLOGY AT HARVARD COLLEGE BULLETIN (octavo) 1863 — The current volume is Vol. 121. BREVIORA (octavo) 1952 — No. 111 is current. Memorrs (quarto) 1864-1938 — Publication was terminated with Volkan, JOHNSONIA (quarto) 1941 — A publheation of the Department of Mollusks. Vol. 5, no. 38 is current. OccCASIONAL PAPERS OF THE DEPARTMENT OF MOLLUSKS (octavo) 1945 — Vol. 2, no. 22 is current. PROCEEDINGS OF THE Nrw ENGLAND ZoouoaicAL CuuB (octavo) 1899-1948 — Published in connection with the Museum. Publication terminated with Vol. 24. The continuing publications are issued at irregular intervals in num- bers which may be purchased separately. Prices and lists may be obtained on application to the Director of the Museum of Comparative Zoology, Cambridge 38, Massachusetts. Of the Peters ‘‘Cheek List of Birds of the World,’’ volumes 1-3 are out of print; volumes 4 and 6 may be obtained from the Harvard Uni- versity Press; volumes 5 and 7 are sold by the Museum, and future volumes will be published under Museum auspices. Bulletin of the Museum of Comparative Zoclogy AT HARVARD COLLEGE Volo 121, No.7 BATHONIAN FORAMINIFERA OF ENGLAND By RicHArpD CIFELLI WitH SEVEN PLATES CAMBRIDGE, MASS., U.S.A. PRN Le De eOM a ler Hes Mas SE: eM SEPTEMBER, 1959 No. 7 — Bathonian Foraminifera of England By RicHARD CIFELLI TABLE OF CONTENTS LIN TORI EO Bi GA Gil td 0G Li NRE AORN Lae ih int ee Ae 8 265 UIT aKOYG GPE Eo een as Se eT SSPE PON EMME, pT ites MTOR fe 2 267 PCa WIEUMCMEMILS uno os So hs she oe a ee ee ee 267 peu Ae GUE ASP TRAY soya 1A sy ccgeree th, oi 20 mies turk bs bres yah kaenden dl anaeay 268 MherBathonian staveunm, Hneland ie... fs... 2s 2s. een 268 WorseteC Caste saan Sevacltn Mey atsis hk dele ace eens ae ee 271 IB AGMVARCA Mest U ys cae. ahaa tere eS ee Marat ete Ree alates 275 Northern Cotswolds and Oxfordshire .................. 278 Northamptonshire and Tuutlamd? 3... nase... gees oe ce 279 DISTRIBUTION OF FORAMINIP HRA (i000. 2508 262 280 SSL RMA TIC DESCRIP TIONS) 0.0. asco 0 teeta le oe sss 282 Be 1 @GIRCACP ERO LIE), SEI IE, RIOR MR LT Ae SU EAN OT INTRODUCTION Whereas the Jurassic megafauna of England has been inten- sively studied for over a century, the Foraminifera and other miroscopic fossils have, until recently, received but scant atten- tion. Before Macfadyen (1941) published his excellent account of the Foraminifera from the Green Ammonite Beds in the Lower lias of the Dorset Coast there were perhaps two dozen papers in the literature dealing with the Jurassic Foraminifera of England. All of these were published before the turn of the century, and most of them contain but few descriptions or illustrations. In almost all instances they are concerned with the faunas of the Lias. Since World War II there has been a renewed interest in Foraminifera, largely through the efforts of 266 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY Barnard and his students. A number of important papers have appeared (Barnard, 1950a, 1952, 1953, 1956, 1957; Adams, 1957), but again, almost all of them thus far have dealt only with the Lias. No Foraminifera have ever been described from the English Bathonian and the only previous records of Fora- minifera in this stage are a few lists of species and genera in the older literature. These are of little value because of the un- certainties in identification and nomenclature. In this paper 104 species, subspecies and varieties (exclusive of attached forms) are described and their known stratigraphic occurrences are recorded. These include one species (Massilina dorsetensis) and one subspecies (Vaginulina clathrata eypensa) which are new. There are, in addition, several previously un- described varieties, but these are not given formal taxonomic status here. No claim is made that this represents the entirety of the Bathonian foraminiferal fauna. The strata of the Bathonian are generally poorly exposed in England, and parts of them are completely concealed. As new road cuts and quarries make addi- tional exposures available there will undoubtedly be descriptions of new and previously unrecorded species as well as additional records of stratigraphical occurrences of known forms. There are four foraminiferal faunules that can be distinguished, which should provide a framework for future faunal divisions of this stage. The faunules are provisional units and can not be used to establish zones at present, because their stratal limits are not known, and their relationships to larger Middle Jurassic faunal units within the European province are not yet known. How- ever, the faunules can be recognized in Dorset and the Bath areas, where they occur in the same order of superposition. Con- sequently, they probably contain stratigraphically restricted species which should later prove useful in establishing zones based on Foraminifera. The Bathonian fauna, like other Jurassic faunas, is character- ized by the dominance of the Lagenidae. Representatives of this family oceur in a large variety of sediments, and in practically all instances they are the most common forms in the foraminiferal assemblages. Other families are represented by few, unspecial- ized, genera and species. CIFELLI: BATHONIAN FORAMINIFERA OF ENGLAND 267 It is well known that many species of Jurassic Lagenidae are highly variable. Only recently, however, have there been any attempts to describe the variation of species (Barnard, 1950a, b ; Adams, 1957). Classification of specimens is difficult because species and genera do not have well defined morphological limits. Nevertheless, it is believed that recognition of variation is vital to an understanding of the evolution of the family ; consequently many lagenid species are conceived here quite broadly. Methods Field work for this investigation was conducted during the summer of 1955. All of the samples are from surface outcrops and each of them weighed about 2 pounds before washing. The samples were prepared in the conventional manner of disaggre- gating in water, sometimes with the assistance of boiling, or adding hydrogen peroxide, and passing them through a 200-mesh screen. In some instances the sample was entirely picked clean, but most of the residues were so large that they were first quartered with a micro-splitter. In the records of species the following scale of relative abundance was used. In each sample, rare corresponds to 1- 3 specimens few co oe AZ Wf ce common a ‘“* 8-19 ae abundant a ‘* over 20 specimens During the spring of 1956 I examined a number of collections containing species previously described from the Jurassic of Europe. Wherever specimens from these collections were used in the identifications of species they are noted in the discussion following the synonymies and descriptions of the species. The types and figured specimens are deposited at the Museum of Comparative Zoology, Harvard University. Acknowledgements This investigation was sponsored by Professor H. B. Whitting- ton, of the Department of Geology, Harvard University, and his advice and helpful criticism is gratefully acknowledged. Pro- fessor B. Kummel of Harvard offered helpful suggestions. The 268 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY late Dr. W. J. Arkell of Cambridge University offered advice on collecting areas and general problems of Bathonian stratigra- phy during the early stages of the investigation. Mr. P. C. Sylvester-Bradley of Sheffield University and Dr. D. T. Donovan of Bristol University provided additional advice on important matters of stratigraphy. To Mr. R. V. Melville of the British Geological Survey I am grateful for details of Bathonian sections in Northamptonshire and Lincolnshire. I also wish to thank the following persons who have helped in many ways: Dr. C. G. Adams of the British Museum of Natural History, Dr. J. Roger of the Laboratory of Paleontology, Muséum National d’Histoire Naturelle, Paris, and Dr. H. Malz of the Senckenberg Museum, Frankfort. Mr. Stuart McNichol, a graduate student at University College, London, collaborated in some of the field work in the Bath and Dorset areas. Finally, I am indebted to Dr. T. Barnard for assistance and the facilities provided at University College during my stay in London. STRATIGRAPHY The Bathonian Stage in England The Enelish strata included in the Bathonian Stage are shown in Figure 1, following Arkell’s (1951, pp. 16-22) interpretation of d’Orbigny’s definition of this stage. In general, the Bathonian in England corresponds to the Great Oolite Series, a lithogenetic unit composed of a highly variable sequence of rocks, chiefly limestones, clays and marls between the Inferior Oolite below and the Oxford Clay and Kellaway Beds above. To Americans unaccustomed to British stratigraphic nomenclature, the term Series is apt to be con- fusing, as in America that term has been standardized in a time- rock sense, denoting a unit larger than a Stage, but smaller than a System. In Great Britain, however, the term is lithogenetic, and is used to unite distinct, but related, rock units without any time connotations. It is similar to the term Group, except that whereas Group combines two or more rock units of formational rank, Series combines rock units of all magnitudes, regardless of rank. CIFELLI: BATHONIAN FORAMINIFERA OF ENGLAND 269 In general, the major units of the Great Oolite Series (often, but not always designated as formations) are as follows, in ascending order: Fuller’s Earth Clay, Great Oolite Limestones, STAGE casnuves (DORSET COAST] BATH AREA LOWER CORNBRASH NORTHERN COTSWOLDS] NORTHAMPTONSH!RE AND OXFORDSHIRE AND RUTLAND LOWER CORN BRASH CORNBRASH | CORNBRASH OBTAINED WYCHWOOD | — — — — — BEDS a en ee FOREST MARBLE BRADEORD | SSe——= a) ae GREAT OOLITE Ls FOREST MARBLE 9 BOUETI BED KEMBLE BEDS FULLER'S - EARTH ROCK LOWER FULLER'S EARTH CLAY BEDS oc © | BEDS SHARPS HILL BEDS (=MAINLY HAMPEN MARLY BEDS ? ARKELL, I95!, p.14) ACUMINATA BEDS LOWER FULLERS|" EARTH CLAY KNORRI_ BEDS NORTON ZIG-ZAG Ls BEDS HOOKNORTON Ls Figure 1. Inferred relationships of English Bathonian strata. LOWER BoA a rhOsNReAGN MIDDLE UPPER 5 Forest Marble and Cornbrash. All of these strata are Bathonian except the Upper Cornbrash, which is Callovian. In addition, the 270 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY base of the Bathonian includes the Zigzag Beds, which comprise the upper few feet of the Inferior Oolite. No Foraminifera were collected from the Zigzag Beds or the Cornbrash. The Great Oolite Series is highly variable and is characterized by great facies changes, particularly below the Forest Marble. In southern England, along the Dorset Coast, the Forest Marble directly overlies the Fuller’s Earth Clay. In the Bath District the Great Oolite Limestone intervenes between the two forma- tions, and farther north in the Cotswolds and Oxfordshire the Fuller’s Earth Clay is mostly replaced by the Great Oolite Lime- stones. These pass northward into the Yorkshire basin, where the Bathonian consists mostly of sands and clays of nonmarine, deltaic origin. The maximum thickness of the Bathonian is in south Somerset, where it reaches a thickness of 500 feet (Arkell, 1951, p. 7). In Northamptonshire the thickness is only about 100 feet. Ammonites, which are generally common throughout the Eng- lish Jurassic and provide accurate guides for subdivisions of this system, are scarce and unevenly distributed in the Bath- onian. They are common only in the Zigzag Beds and in the Cornbrash. In the Fuller’s Earth Clays and much of the Great Oolite Limestones they are very rare or absent. It is only recently that the Bathonian has been divided into zones based on ammonites (Arkell, 1951, p. 21). It is doubtful, however, that these zones are as reliably established as others in the Jurassic. At Whatley in Somerset, for example, a recent large collection of ammonites from the Rugitella Beds, a few feet above typical Subcontractus Zone, yielded a mixture of Lower, Middle and Upper Bathonian species. Arkell (1957, p. 824) states, ‘‘The assemblage from the Rugitella Beds at Whatley is highly ano- malous. It does not contain any of the zonal index species or eenera of the Lower, Middle or Upper Bathonian (Zigzagiceras, Morphoceras, Tulites, Morrisiceras, Clydoniceras) but consists of a mixture of accessory ammonites individually considered charac- teristic of all 3 subdivisions of the Bathonian, but never previ- ously found together.’’ It appears that even these most honored of all index fossils may sometimes be susceptible to facies changes, as are other fossils. CIFELLI: BATHONIAN FORAMINIFERA OF ENGLAND 271 Because of this lack of ammonites, correlations within the Bathonian are difficult, and the relationships of many of the strata are uncertain. Brachiopods are locally abundant, but their distribution is uneven, and they have been useful, thus far, only for local correlations. Foraminifera are abundant throughout most of the Bathonian, but this present effort repre- sents but a beginning in the establishment of the Bathonian foraminiferal succession. The stratigraphy of the English Bathonian has already been summarized (Woodward, 1894; Arkell, 1933, 1951, 1956; Arkell and Donovan, 1952), and the present account is limited to those details necessary to understanding the succession of Foramini- fera. The major relationships of the strata are summarized in Figure 1, which has been compiled from many sources in the literature, but conforms mostly to the authority of Arkell. Dorset Coast The best English Bathonian exposures are found along the Dorset Coast between Bridport and Weymouth. A composite section showing the stratigraphic positions of the foraminiferal samples is given in Figure 2 and the stratigraphic occurrences of the foraminiferal species are shown in Table 1. The strata are nowhere exposed in their entirety and the section has been pieced together from a number of localities (Arkell, 1933, p. 200). The Lower Fuller’s Earth Clay is poorly exposed and most of it is never seen. Some of the clay occurs in the eastern part of Watton Cliff, but the beds there are faulted and the stratal relations are uncertain. The lower few feet of the Lower Fuller’s Earth Clay are exposed at the top of Burton Chiff, between the Bredy River and Burton Bradstock. The cliff is not readily accessible and the present samples were obtained from a large boulder at the bottom of the cliff which had toppled from the top. The block contained 6 feet of Lower Fuller’s Earth Clay, including a few inches of Seroff at its base, underlain by the Zigzag Bed and the Inferior Oolite. The Fuller’s Earth Roek does not extend into the Dorset Coast and overlying the Lower Fuller’s Earth Clay in this area are the Wattonensis Beds (Kellaway and Wilson, 1941, p. 160), DAP BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY FORMATION LITHOLOGY ARGILLACEOUS, PLATY LIMESTONES GRAY, PLASTIC CLAY WITH OCCASIONAL THIN, | PLATY LIMESTONES MASSIVE ,SHELLY LIMESTONE GRAY OR GREENISH PLASTIC CLAY, SOMETIMES CALCAREOUS, WELL INDURATED FOREST MARBLE BOUETI BED MASSIVE GRAY CLAY, WITH CONCHOIDAL FRACTURE 35 = joa tt WwW Nn (Ve lJ = —J > re joa WwW a (oi =) ag OSTREA HEBRIDICA VAR. ELONGATA 13 BLUE-GRAY CLAY WITH THIN FOSSILIFEROUS BEDS OF ARGILLACEOUS LIMESTONES BLUE CLAYS, PARTIALLY EXPOSED IN FAULTED AREA OF WATTON CLIFF MOSTLY UNEXPOSED 5 GRAY CLAY WITH LIMONITIC STAINS Figure 2. Composite section of Bathonian strata of the Dorset Coast. CIFELLI: BATHONIAN FORAMINIFERA OF ENGLAND 23 a series of dark, argillaceous limestone bands with intervening clays. They occur along the beach at the base of Watton Cliff, just east of Eype Mouth. The beds strike NNE, parallel to the fault which extends through Eype Mouth, and dip sharply to the east. There are about eight limestone bands, each about one-half to one foot thick and as many intervening clays. The total thickness is estimated to be twenty-five feet (Buckman, 1922, p. 381), but the outcrop is mostly covered by the shingle and can be observed at the foreshore only at unusually low tides. Fora- minifera were collected from the clay bands, immediately below each limestone. At Watton Cliff there are approximately 100 feet of the Upper Fuller’s Earth Clay exposed in the vertical cliff wall. The upper contact is clearly defined in the brow of the cliff, where the Upper Fuller’s Earth Clay is overlain by the Forest Marble, with the Boucti Bed at the base. A much more accessible section is exposed 3 miles to the east at Cliff’s End, Burton Brad- stock, where there are about 60 feet exposed and overlain by the Boueti Bed (Arkell, 1933, p. 253). The clay is arenaceous and contains abundant fragments of quartz, chert and mica. The lower portion of the Upper Fuller’s Earth Clay is concealed beneath the beach and is not visible in the vicinity of Bridport, but the lower beds are exposed 12 miles to the east, at Langton Herring. They consist of tough, dark gray clays containing masses of shells belonging mostly to Ostrea hebridica var. elon- gata Dutertre. This is the Llongata Bed, which is 12 feet thick at Laneton Herring and is underlain by the Wattonensis Beds (Arkell, 1933, p. 252). The base of the Forest Marble is represented by the Boueti Bed, a remarkably persistent fossiliferous marl, only one foot thick and characterized chiefly by the abundance of Goniorhyn- chia boueti (Davidson). This bed is best seen at Herbury along the West Fleet, but persists as far inland as Wincanton and Sherborne (Kellaway and Wilson, 1941, p. 162). In the past it has been customary to correlate the Boueti Bed with the Brad- ford Clay, as is shown in the chart in Figure 1. Recently, how- ever, Sylvester-Bradley (1957, p. 27) has presented evidence to suggest that the Boueti Bed is older than the Bradford Clay and is the equivalent of the Bath Freestone in the Bath area. 274 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY The older, conventional correlation is retained here, however, pending a more complete statement on the problem. Practically the entire Forest Marble is exposed at Watton Cliff, and is accessible at Fault Corner. The following section was obtained (measured by Cifelli and McNichol, 1955) : feet Platy, argillaceous limestone, gray on fresh surface, weathering rust brown; interbedded with thin marl bands; Srades imborelaysbelOwe rian aie cece occ ce eee eee i Gray or green soft, plastic clay, sandy, micaceous, with thin bands of argillaceous limestone becoming more numer- OUS COWATUS: LODE c eee Wits che cs ais Spe sicrs ore eter 21 Massive, oolitic limestone, gray on fresh surface, weather- ing buff, crossbedded ; abundant broken shells, pectenids, oysters and Wwew brachiopods. 72.9 ts et ene eee es oe 10 Gray or green or brown soft clays, weathering to rust brown; sandy, micaceous; interbedded with thin, calcareous, well indurated clays containing shell fragments. .......... 47 Total 85 Localities of foraminiferal samples: Samples 1-5 were all collected from a single boulder at the base of Burton Cliff, 1.2 miles southeast of Bridport Harbour. The total stratigraphic interval represented in the boulder is about 6 feet. Samples 6-13 were collected from the foreshore of Watton Cliff, at low tide level. Each sample was taken from the clay immedi- ately underlying each of the argillaceous bands. The most westerly and stratigraphically lowest of these was located about 30 yards east of Eype Mouth. Samples 14-15 were collected along the West Fleet, about 600 yards west of the Coast Guard Station at Langton Herring. Samples 16-17 are from the Hlongata Beds, about 200 yards west of the ferry at Langton Herring. Sample 18 was taken from the vertical cliff wall of Watton Cliff, about midway between Eype Mouth and West Bay, about 10 feet above the level of the beach. Samples 19-27 are from Cliff’s End, in the cove about 300 yards WNW of the Coast Guard Station. CIFELLI: BATHONIAN FORAMINIFERA OF ENGLAND atic —) Samples 28-31 were collected from Herbury, about 1 mile south of Langton Herring. Samples 32-40 were collected from the western part of Watton Cliff, at Fault Corner, near EKype Mouth. Bath Area Stratigraphic imvestigations in the Bath area date back to the mapping of William Smith. Exposures at the present time are poor, however, and most of the quarries and railroad cut- tings from which the succession was established are now covered. Only isolated portions of the sequence are revealed and the exact horizous of some of the exposures are uncertain. For this reason, Foraminifera collected from the Bath area have not been com- piled into a single, composite check list. They are shown in indi- vidual tables for each formation, at the localities from which they were collected (Tables 2-11). The most complete section of the Lower Fuller’s Earth Clay in the Bath area was obtained by Richardson (1910, p. 426) from a railway cutting at Combe Hay, about 3 miles south of Bath. Richardson recorded 35 feet of the clay, containing Zigzagiceras at the base, resting on the Inferior Oolite. There was no trace of the Fuller’s Earth Rock, but the occurrence of Ostrea acwmi- nata in the upper layers suggested that almost the entire thick- ness of Lower Fuller’s Earth Clay was represented (Arkell, 1933, p. 280). The cutting is now obscured, but 2 miles east of Combe Hay in the railway embankment at Midford there are 10-12 feet of the clay resting on the Inferior Oolite. The weathered, yellow clay which can be reached within a couple of feet of the grass covered surface yields numerous, moderately well preserved Foraminifera. The Fuller’s Earth Rock is well developed in the Bath area. It is still exposed in the road cut between Maperton and Charle- ton-Horetorne, where it rests on the Lower Fuller’s Earth Clay. The section was described by Richardson (1909, p. 213) who recorded 2214 feet of Fuller’s Earth Rock and 15 feet of Lower Fuller’s Earth Clay. In the town of Box, 5 miles northeast of Bath, the upper part of the Lower Fuller’s Earth Clay appears to be overlain by the Fuller’s Earth Rock. The area has not 276 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY been mapped in detail and the stratal relations are not certain. However, in the summer of 1955 a drainage ditch was dug at the road intersection about 50 yards down hill from Rose Cottage in Box. It revealed a rubbly, oolitie limestone underlain by 6 feet of dark brown clay. Another ditch, about 10 feet below the first, exposed an additional few feet of clay. In view of the Old Series 1 inch map showing the Great Oolite Series at Box and Lycett’s record of 148 feet of Fuller’s Earth, including Fuller’s Earth Rock at Box Tunnel (Arkell, 1933, p. 281), it appears lkely that the upper part of the Lower Fuller’s Earth Clay and the Fuller’s Earth Rock were represented in the ditches. Foramini- fera were collected from the clays (Table 4). At Cross Ways Inn, a few miles south of Bath, at the inter- section of the roads to Radstoeck and Timsbury, Cox (1941, p. 20) described the following section: feet Marly clay (Upper Pullerss! Barth); (js2.ce)ascer 4: 2—0 tubbly limestone (Fuller’s Earth Rock proper), with peleeypods and brachiopods including species of Rugitella, Ormthella, Rhynchonelloidea, Wat- LOTALRULIS DO nn tuibt leat bexregoee ara AM a -tatn eter Dbee 50) Mottled: telayigy:n asad canoe ods took yak then A ces ae 5—0 LIMESTONE; wehctosy elon: Ades Pactacena zie ceesisrel weoseea eee 0—6 Clay with Rhynchonelloidea turchert .............. 1—6 Areillaceous JimestOne wets. ange 36 eis sctece de eles 0—6 Mottled felatyaj Si) tcctearaetars fier nt red Uni teas tet ee 5—0 Clay with abundant Ostrea acuminata .............. 10) Clay ng aidtine « beduntaophoved wlisieletswsle oagtteacnioe 3—0 Estimated thickness of beds underlying grass slope to top of Inferior Oolite:.... ss. a.4uehirsseaee noe 15—0 The section is unusual in that there are only 5 feet of rubbly limestone (‘‘Fuller’s Earth Rock proper’’) represented and 121% feet of clay intervene between the limestone and the clay containing Ostrea acuminata. Normally the Acwminata Beds occur immediately below the Fuller’s Earth Rock and tradi- tionally they have been regarded as a constant paleontological horizon marking the top of the Lower Fuller’s Earth Clay. How- ever, this oyster was found to straddle the boundary and occur o7 CIFELLI: BATHONIAN FORAMINIFERA OF ENGLAND PAT (| in the Fuller’s Earth Rock at Whatley (Sylvester-Bradley and Hodson, 1957, p. 315). The section at Cross Ways Inn is now covered with grass, but one sample was recovered about 4 feet below the surface which yielded a rich assemblage of Foramini- fera (Table 6). North of Bath the Fuller’s Earth Rock splits into 2 separate limestone bands, an upper Tresham Rock and a lower Cross Hands Rock, separated by a clay bed (Hawkesbury Clay). No Foraminifera were obtained from these units. The best known part of the Upper Fuller’s Earth Clay is the commercial clay which occurs in a 5 or 6 foot bed near the top. It is a blue or green clay which weathers to a buff or yellow color, lacks plasticity and disintegrates completely in water. It has been mined for many years in the vicinity of Bath. At Midford it is separated from the overlying Great Oolite Lime- stones by 1714 feet of unworkable clay, while at Combe Grove there are 1114 feet of intervening clay (Woodward, 1894, p. 242). The commercial clay is presently being worked at the Combe Hay Fuller’s Earth Works on Fosse Way, about 3 miles south of Bath. The Foraminifera collected there are shown in Table 7. The top beds of the Upper Fuller’s Earth Clay are exposed at Vernham Wood, about 1144 miles west of the Combe Hay Fuller’s Earth Works, where the commercial clay is worked oceasionally from small, open cast pits. The following section was obtained from one of these pits (measured by MeNichols and Cifelli, 1955) : feet a— Rubbly, white oolitic limestone (Base of Great Oolite Lamestones eins os oot). ees eee «en i) b — Green, plastic clay with thin, oolitic marl at the GOP | % Adie Sis wo Moh Bist GlE 2s, Segui aes Shep ena tae Bape 2—6 e — Mottled, carbonaceous, dark gray and black plastic GTA. Sag ther idesiteice oer pears ts kat Setanta Se erate 1—3 d — Buff, oolitie limestone with 2” clay band in middle = 1—9 e— Green clay, weathering buff, calcareous, non- plastic (probably the top of the commercial BullerissMarth)) (Ol heme Sada: eed deere oe 2—4 278 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY The Foraminifera are shown in Table 8. Additional Foramini- fera from the top part of the Upper Fuller’s Earth Clay were obtained on Henley Hill, 1 mile south of Box, where the clay is overlain by the Great Oolite Limestones (Table 9). The Forest Marble overlies the Great Oolite Limestones (which separate it from the underlying Upper Fuller’s Earth Clay) and consists mostly of shelly, oolitic limestones and clays. At Brad- ford-on-Avon, 5 miles east of Bath, the base of the Forest Marble is represented by the Bradford Clay, consisting of 10 feet of clay with thin layers of argillaceous limestone. At the bottom is a distinctive fossil bed which occurs intermittently at the same horizon from Somerset to Oxfordshire (Arkell, 1933, p. 269). It has been customary to correlate this fossil bed with the Boueti Bed of Dorset, although Sylvester-Bradley (1957, p. 27) has suggested a correlation of the Bradford Clay with the Digona Beds, a fossiliferous horizon somewhat higher in the Forest Marble of Dorset. The Foraminifera from the Bradford Clay are shown in Table 10. The Forest Marble is revealed below the surface in the Cor- sham Monk Quarry, on Monk Lane near Gastard, where the Bath Freestone is mined for building stone. At the bottom of the quarry there are 28 feet of Bath Freestone overlain by 3 feet of the Upper Rag. Above this there are approximately 70 feet of Forest Marble in the shaft of the quarry, consisting of lime- stones, clays and marls. The clays are wet and weathered, yielding relatively few, poorly preserved Foraminifera (Table dal yi: Northern Cotswolds and Oxfordshire In this region the Great Oolite Series is dominantly of a caleareous facies, composed chiefly of limestones with lesser amounts of clay and marl. The relationships of the strata have been described by Arkell (1951) and Arkell and Donovan (1952): An almost complete section of the Great Oolite Series is ex- posed along the railway between Chedworth and Cirencester in East Gloucestershire. The cuttings extend a distance of 8 miles and were deseribed by Richardson (1911la; 1933). The succession includes beds ranging from the Lower Fuller’s Earth Clay CIFELLI: BATHONIAN FORAMINIFERA OF ENGLAND 279 at the base to the Kemble Beds (Lower Forest Marble) at the top. Although the railway embankments are now partially covered, many of the beds may still be observed. The Hampen Marly Beds and Taynton Stone were not included in Richard- son’s original description of the section, but were later identified as beds 33-35 and 36, respectively, of Richardson by Arkell and Donovan (1952, p. 246) and Green and Melville (1956, pp. 10-11). The Stonesfield Slate Beds have never been positively identified along the railroad, but Green and Melville (1956, p. 3) believe that they may be present below the Chedworth Station. Most of the Great Oolite Limestones are exposed in the Hampen railway cuttings, about 5 miles east of Cheltenham. The section was described by Richardson (1929, pp. 102-106), and includes beds ranging from the Lower Fuller’s Earth Clay to the White Limestone. The Taynton Stone and the Hampen Marly Beds are particularly well represented in this sequence. In Oxfordshire the upper part of the Bathonian is represented in the quarry of Kirtlington Cement Works, Kirtlington. The section extends from the White Limestone at the base to the Cornbrash at the top. The lower part of the sequence is exposed in the Sharp’s Hill Quarry near Hook Norton, northern Ox- fordshire. The section includes the Hook Norton and Chipping Norton Limestones, which replace the Lower Fuller’s Earth Clay at the base, and the Lower and Upper Sharp’s Hill Beds. Foraminifera collected from the northern Cotswolds and Oxfordshire are shown in Tables 12-16. Because of the dom- inantly calcareous nature of the Bathonian in this region suitable foraminiferal samples are difficult to obtain, and portions of the sequence have been but sparsely sampled. No Foraminifera were obtained from the Lower Fuller’s Earth Clay or the Cornbrash. Northamptonshire and Rutland Foraminifera were collected only from the Upper Estuarine Beds, a series of various colored clays with some sandy shelly beds and limestones. They include both freshwater and marine deposits. Foraminifera are not common (Tables 17-18), but there are numerous ostracods and charophytes. 280 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY DISTRIBUTION OF FORAMINIFERA The present records of species are still too meager to establish the succession of Bathonian Foraminifera in England. More- over, many of the Jurassic Foraminifera are yet to be described, so that the total ranges of the Bathonian species and their dis- tribution throughout the European province cannot as yet be determined. Consequently, formal subdivision of Bathonian strata based on Foraminifera would hardly be justified at this time. The present records of occurrences of species (Tables 1-18) do reveal that there are 4 faunules that ean be recognized in the Dorset and Bath areas. In both of these areas the faunules occur in the same order of superposition and therefore probably contain elements diagnostic of faunizones. North of Bath, where the strata are dominantly of a calcareous facies, the foraminiferal assemblages are generally less rich, containing fewer species. The preservation is often poor and the assemblages are much less distinctive; the faunules cannot be recognized north of Bath. Faunule A This faunule occurs in the basal few feet of the Lower Fuller’s Earth Clay in Dorset and Bath. The species which have been observed to occur only in this faunule are: Vaginulina macilenta, Frondicularia nodosaria var. A and Lenticulina quenstedti var. A. Although not restricted, Lenticulina galeata and Planularia eugent are particularly common here. The following species occur in faunule A and succeeding faunules but have not been recorded from strata older than Bathonian in England: Denta- lina oolithica, Lenticulina quenstedti (sensu lato), L. tricarinella, Nodosaria opalini and Trochammina haeuslert. Species of fau- nule A, not recorded from the sueceeding faunules, but charac- teristic of the Lias are: Vaginulina clathrata, Frondicularia lignaria and Nodosaria liassica. Faunule B This faunule has been recognized in the upper beds of the Lower Fuller’s Earth Clay, the Fuller’s Earth Rock and the Wattonensis Beds. Common and characteristic of this faunule, CIFELLI: BATHONIAN FORAMINIFERA OF ENGLAND 281 though not restricted to it, are: Planularia beierana, Lenticulina tricarinella and L. munsteri. Hold-overs from the previous faunule which have not been observed to range above this one are: Citharina colliezi, Nodosaria aff. N. prima, Lenticulina quenstedti var. B, L. tricarinella, and Planularia eugenii. Species not occurring in faunule A, but occurring in faunule B and succeeding faunules are: Dentalina propinqua, Nodosaria pecti- nata, Frondicularia spissa and Vaginulina clathrata eypensa n. subsp. Faunule C This faunule is characteristic of the Upper Fuller’s Earth Clay in the Dorset and Bath areas. Common and characteristic, though not restricted to this faunule, are: Dentalina intorta, Nodosaria opalim, Frondicularia nodosaria, Epistomina. stelli- gera and Lenticulina subalata. Species occurring in faunules C and D but not observed in previous faunules are: Citharina heteropleura and Dentalina aft. communis. Species occurring in this and previous faunules but not faunule D are Vaginulina harpa and Nodosaria pectinata. Faunule D This faunule characterizes the Forest Marble, including the Boueti Bed and the Bradford Clay. It is very similar to faunule C, but species of Lenticulina are decidedly less common here. The most characteristic aspect of Faunule D is the common, almost restricted occurrence of Massilina dorsetensis n. sp. The lowest observed occurrence of this species was in the beds im- mediately underlying the Boueti Bed at Herbury (Sample 30). Nodosaria clavula and Frondicularia intumescens have not been observed in other faunules, but both of these species are rare. Vaginulina clathrata eypensa n. subsp. and Dentalina subplana make their last appearances in the Boueti Bed and Bradford Clay, the basal beds in which this faunule oceurs. 282 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY SYSTEMATIC DESCRIPTIONS Phylum PROTOZOA Order FORAMINIFERA Family SACCAMMINIDAE Genus THURAMMINA Brady, 1879 THURAMMINA TUBEROSA Haeusler Plate 1, figure 29 Thurammina tuberosa Haeusler, 1890, p. 49, pl. 6, fig. 24; pl. 7, figs. 6-9. Thuramminopsis canaliculata Haeusler, 1890, p. 50, pl. 7, figs. 1-5; pl. 8, figs. 4-6. Thurammina papillata, Frentzen (non Brady), 1944, p. 322, pl. 17, figs. 1-9. The present specimens are large, globular and little com- pressed. The walls are composed of very fine grains, with abun- dant cement, so that they appear almost calcareous. There are a number of tiny apertures on various parts of the test which have the appearance of pin holes. The shapes are irregular, but the specimens agree with the figures cited. Family REOPHACIDAE Genus REOPHAX Montfort, 1808 REOPHAX MULTILOCULARIS Haeusler Plate 1, figures 6-7 Reophax multilocularis Haeusler, 1890, p. 28, pl. 3, figs. 9-11, 26. Reophax multilocularis, Bartenstein and Brand, 1937, p. 133, pl. 5, fig. 8; DEVS) figss8a-bi ple LO tio 9s pla vldlas ction o5) el mloar tions ee pheloce fig, 2. Reophax scorpiurus, Bartenstein and Brand (non Montfort), 1937, p. 134, Diaries) plevos digs 19. This species is very rare, and generally poorly preserved in the present material. The arenaceous wall is moderately coarse, and there are four or five chambers which are slightly higher than broad. The aperture is terminal and rounded. In one specimen the test is shghtly curved at the base, but there is no indication of a coil. eal at CIFELLI: BATHONIAN FORAMINIFERA OF ENGLAND 283 Family LITUOLIDAE Genus FLABELLAMINA Cushman, 1928 FLABELLAMINA ALTHOFFI Bartenstein Plate 1, figures 25-26 Flabeliamina althoffi Bartenstein in Bartenstein and Brand, 1937, p. 187, pl. 14a, fig. 8a-b; pl. 15a, fig. 41a-b. The specimens are higher in relation to breadth, but otherwise compare favorably with the types of the species. The arenaceous wall consists of large grains mounted in abundant calcareous cement, which gives the test a smooth appearance. The grains are generally quartz, but oolites and occasional shell fragments also occur. This species superficially resembles Triplasia bartensteim, and may possibly represent a flattened variant of the latter. No actual transitions between the two forms, however, were ob- served. Genus AMMOBACULITES Cushman, 1910 AMMOBACULITES AGGLUTINANS (d’Orbigny ) Plate 1, figure 1 Spirolina agglutinans d’Orbigny, 1846, p. 187, pl. 7, figs, 10-12. Haplophragmiwm coprolithiforme, Deeke (non Schwager), 1884, p. 20, ple deetiga io. Haplophragmium coprolithiforme, Haeusler, 1890, p. 33, pl. 4, figs. 7, 20. Ammobaculites agglutinans, Franke (non Schwager), 1936, p. 127, pl. 12, fig. 25. Ammobaculites agglutinans, Bartenstein and Brand, 1937, p. 186, pl. 4, fies id) ple 5, Aes 185. ple Gj fig. wta-bs pli 8S satieesS8a-c opi. 0; fig. 45a-b; pl. lla, fig. 19a-b; pl. 11b, fig. 28a-b; pl. 12a, fig. 22; pl. 13, fie 235, pls l4b; fig. 19: Bartenstein and Brand (19387, p. 186) state that the wall material is variable in this species. Among the Bathonian forms the wall is remarkably uniform. It is composed of very fine arenaceous particles, and usually contains a large amount of pyrite. The tests are small, and mostly uncoiled. The species occurs throughout the Bathonian, but individuals are few. 284 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY AMMOBACULITES FONTINENSIS (Terquem) Plate 1, figures 2-5 Haplophragmium fontinense Terquem, 1870b, p. 337, pl. 24, figs. 29, 30. Ammobaculites fontinensis, Franke, 1936, p. 127, pl. 12, fig. 25. Ammobaculites fontinensis, Bartenstein and Brand, 1937, p. 186, pl. 5, fig. 79a-b; pl. 6, fig. 43; pl. 8, fig. 37a-d; pl. 10, fig. 44; pl. 11b, fig. 27; pl. 12a, fig. 2la-b; pl. 13, fig. 22. Ammobaculites fontinensis, Barnard, 1950b, p. 4, pl. 1, fig. 2. This species shows little selection in the nature of the wall material, which appears to be determined by the nature of the enclosed sediment. The wall is finely to coarsely arenaceous, consisting mostly of quartz grains, but occasionally oolites as well. The degree of coiling is highly variable, and some speci- mens are practically linear with a minute spire, while others are completely coiled. Family TEXTULARIDAE Genus TEXTULARIA Defrance, 1824 TEXTULARIA JURASSICA Gumbel Textularia jurassica Gumbel, 1862, p. 228, pl. 4, fig. 17a-b. Textularia franconica Gumbel, 1862, p. 229, pl. 4, fig. 18a-b. Textularia agglutinans, Blake (non d’Orbigny), 1876, p. 472, pl. 17, fig. 37. Textularia jurassica Paalzow, 1932, p. 94, pl. 4, figs. 21-23. Textularia agglutinans, Bartenstein and Brand, 1937, p. 182, pl. l4a, fig. 5a-b; pl. 14e, fig. 16; pl. 15a, fig. 40a-c; pl. 15b, fig. 3a-c; pl. 15e, fig. 2la-b. Textularia jurassica, Seibold and Seibold, 1955, p. 98, pl. 13, fig. 1; text fig. 2a. Specimens are rare, and agree well with topotypes from the Gumbel locality. Material deposited at M.C.Z. No. 3306. Family VERNEUILINIDAE Genus VERNEUILINA d’Orbigny, 1840 VERNEUILINA MAURITI Terquem Verneuilina mauritii Terquem, 1866a, p. 448, pl. 18, fig. 18a-b. Verneuilina georgiae Terquem, 1866a, p. 448, pl. 18, fig. 19a-b. Verneuilina mauritii, Franke, 1936, p. 126, pl. 12, figs. 22, 23. CIFELLI: BATHONIAN FORAMINIFERA OF ENGLAND 285 Verneuilina mauritii, Bartenstein and Brand, 1937, p. 1838, pl. la, fig. 22. Verneuilina mauritii, Cushman, 1946, p. 6, pl. 1, figs. 1-2. Verneuilina mauritii, Usbeck, 1952, p. 385, pl. 14, fig. 1D. There is only one specimen and it agrees well with other described forms of this species except that the early part of the test is shghtly arched. Material deposited at M.C.Z. No. 3307. Family SILICINIDAE Genus PROBLEMATINA Borneman, 1874 ?PROBLEMATINA ef. P. LiAssIcaA (Jones) Plate 1, figure 8 One specimen recovered from the Elongata Beds at Langton Herring superficially resembles Problematina liassica as figured by Macfadyen (1941, p. 19, pl. 1, figs. 9-11). The pillars are well seen on the ventral surface, but the specimen is otherwise poorly preserved and largely replaced by pyrite. The composi- tion of the wall is therefore uncertain; but the granular appear- ance suggests that it is arenaceous, in which case it would differ from the forms analyzed by Macfadyen. This species has been previously reported from the Lias only. Genus HAPLOPHRAGMIUM Reuss, 1860 HAPLOPHRAGMIUM SUPRAJURASSICUM Schwager Plate 1, figures 9-10 Haplophragmium suprajurassicum Schwager, 1865, p. 92, pl. 2, fig. 1. Haplophragmium aequale, Bartenstein and Brand (non Roemer), 1937, p. 187, pl. 12a, fig. 24; pl. 13, fig. 25a-b; pl. 14a, fig. 9a-b; pl. 14b, fig. 20a-b; pl. 15a, fig. 42. The walls are generally coarsely arenaceous, but the size of the erains is variable, and occasional specimens are fine grained. There is a sheht tendency towards uncoiling, with only one or two chambers beyond the coil. Genus TRIPLASIA Reuss, 1853 TRIPLASIA BARTENSTEINI Loeblich and Tappan Plate 1, figure 11 Triplasia variabilis, Bartenstein and Brand (non Brady), 1937, p. 185, pl. 14a, fig. 6. 286 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY Triplasia bartensteini Loeblich and Tappan, 1952, p. 8, pl. 1, fig. 4. The cross-section of this species is highly variable, which is also characteristic of other species of Triplasia. Triradiate forms intergrade imperceptibly into quadrate forms, and I agree with Loeblich and Tappan (1952) in suppressing the genera Tetraplasia Bartenstein and Centenarina Majzon which are distinguished from Triplasia on the quadrate form of the eross- section. Some of the specimens are somewhat compressed, but no transition to the specimens included here in Flabellamina althoffi was observed. The height of the spire is also variable, and in some specimens it comprises about half of the test, while in others it is minute, though always conspicuous. This species is common is certain beds of the Upper Fuller’s Earth Clay at Dorset and Vernham Wood, near Bath. Family MILIOLIDAE Genus MASSILINA Schlumberger, 1893 In redeseribing some of the species of d’Orbigny, Schlum- berger (1893) erected the genus Massilina for those species of Spiroloculina d’Orbigny in which quinquiloculine growth of the test was highly developed. It differs, therefore, from Spirolocu- lina in degree only, and does not possess any distinctive morpho- logical features. The two genera overlap considerably, and as mentioned by Wood and Barnard (1946, p. 83), there would be great difficulty in producing natural groups from specimens involved in these genera. The evolutionary significance of the degree of development of the quinqueloculine growth stage has yet to be demonstrated, and perhaps there is no adequate way of distinguishing the two genera. Pending further study, however, the genus Massilina is retained here for those specimens in which the major part of the test is quinqueloculine and the latter part is planar. MASSILINA DORSETENSIS Nl. sp. Plate 1, figures 15-17 Diagnosis. Test small, suboval in outline, slightly biconvex in cross-section, with thin, somewhat transparent wall; eight CIFELLI: BATHONIAN FORAMINIFERA OF ENGLAND 287 chambers, each half a whorl] in length and increasing in size as added, the first six arranged in a quinqueloculine series, the last two opposite; sutures flush, producing a smooth surface on both sides of test; proloculus minute, spherical; aperture simple, without tooth. Discussion. All of the proloculi examined were minute, show- ing no appreciable differences in size, so that microspherie and megalospheric generations are not distinguishable in the present material. The species is distinctive, although it varies consid- erably in size, and to a lesser extent in thickness and ratio of width to length. This species most closely resembles Spiroloculina lanceolata (Terquem and Berthelin), from which it may be directly de- scended. It is separated from the latter by the much greater development of the quinqueloculine stage and the lack of a long neck on the last chamber. This species is common in the Upper Fuller’s Karth Clay and Forest Marble, but was not observed at lower horizons. Greatest length of holotype: .85 mm. Greatest width of holotype: .18 mm. Greatest thickness of holotype: .06 mm. Material deposited at M.C.Z. No. 3311C1. Genus SPIROLOCULINA d’Orbigny, 1826 SPIROLOCULINA LANCEOLATA (Terquem and Berthelin) Plate 1, figures 18-19 Quinqueloculina lanceolata Terquem and Berthelin, 1875, p. 84, pl. 7, fig. 7a-b. Quinqueloculina ovula Terquem and Berthelin, 1875, p. 85, pl. 7 Quinqueloculina rotundata Terquem and Berthelin, 1875, p. 85 10a-b. Quinqueloculina compressa Terquem and Berthelin, 1875, p. 85, pl. 7, fig. 1la-b. Quinqueloculina contraria Terquem and Berthelin, 1875, p. 85, pl. 7, fig. 12a-b. The early portion of the test is very small, and consists of a variable number of chambers arranged in a quinqueloculine fashion. The last two chambers are opposite. The final one is characterized by a rather long neck, but this is found only on ; fig: 8a-b: 5 Tolle Ze ane, 288 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY the better preserved specimens. The resemblances and differ- ences with Massilina dorsetensis have already been noted above: it might be added here that Spiroloculina lanceolata is generally smaller and slimmer, and the quinqueloculine portion more con- densed. This species is fairly common throughout the English Bath- onian. Family OPHTHALMIDIIDAE Genus CORNUSPIRA Schultze, 1854 CORNUSPIRA LIASINA Terquem Plate 1, figures 27-28 Cornuspira liasina Terquem, 1866b, p. 474, pl. 19, fig. 4a-b. Cornuspira orbicula Terquem and Berthelin, 1875, p. 17, pl. 1, fig. 12a-e. Comusptra orbicula, Bartenstein and Brand, 1937, p. 137, pl. la, fig. 3a-b; pl wh; fies 359 pliv2a figs 3a-bs) pl. abs hige bo plata, aie. a4 lends figs Woe ple eo, aigees Upl) 6 te’ GCa-bise ply ace etigs as bit a olsamnOs fig. 6a-b; pl. 12a, fig. 2a-b; pl. 12b, fig. 1; pl. 14¢, fig. 2. Cornuspira liasina Bartenstein and Brand, 1937, p. 131, pl. 4, fig. 8. Coiled, tubular forms are abundant in the English Bathonian, but their relationships are difficult to ascertain because of the minuteness of the test and the imperfect preservation of the wall. Many specimens included here may actually belong to Spirillina or Ammodiscus. The forms are being investigated at present by Mr. Stuart McNichol of University College, London. Mr. Me- Nichol is using very high magnification, and is studying the wall structure and growth plan of the early part of the test in great detail. His results will undoubtedly shed new light on the relationships of these forms, and the distributional patterns of species of Cornuspira, Spirillina and Ammodiscus will probably require some revision. Genus SPIROPHTHALMIDIUM Cushman, 1927 SPIROPHTHALMIDIUM CONCENTRICUM (Terquem and Berthelin) Plate 1, figures 12-14 id Spiroloculina concentricum Terquem and Berthelin, 1875, p. 80, pl. 7, figs. 1-4. CIFELLI: BATHONIAN FORAMINIFERA OF ENGLAND 289 Spirophthalmidium tenuissimum Paalzow, 1932, p. 100, pl. 5, figs. 11-13. Spirophthalmidium concentricum, Franke, 1936, p. 123, pl, 12, figs. 15, 17. Spirophthalmidium concentricum, Bartenstein and Brand, 1937, p. 181, pl. 2b, figs. 37, 38; pl. 4, fig. 16; pl. 5, fig. 7la-b; pl. 8, fig. 36; pl. 13, fig. 21; pl. 15a, fig. 39a-b. The proloculus and early spiral are minute, but under ordinary magnifications the specimens can be seen to lack clearly defined ophthalmid structure. The validity of the genus has been ques- tioned, but I agree with Wood and Barnard (1946, p. 87) that the name should be retained until the relationships of the Ophthalmidiidae are better understood pending further study of the living representatives. Family TROCHAMMINIDAE Genus TROCHAMMINA Parker and Jones, 1859 TROCHAMMINA HAEUSLERI (Galloway) Plate 1, figures 21-22 Valvulina triangularis, Haeusler (non d’Orbigny), 1890, p. 75, pl. 12, figs. 23, 24. Valwulina conica, Haeusler (non Parker and Jones), 1890, p. 76, pl. 12, figs. 27-35. Tritaxis haeusleri Galloway, 1933, pl. 19, fig. 2a-b. Valvulina (2?) haeusleri, Cushman, 1937, p. 4, pl. 1, fig. 1. Valvulina (?) sp. Cushman, 1937, pl. 1, figs. 2-4. Test small, finely arenaceous with much cement, conical, chambers arranged in a trochoid spiral, about four to a whorl in the early portion, later only three; sutures flush; proloculus tiny, globular, alternate generations not apparent; aperture a poorly defined slit at base of last chamber in the umbilical region. Haeusler did not figure a side view of this form which Gallo- way later used as the type of the species. Galloway interpreted the test as being flat, and apparently on this basis referred the species to Tritaxis. In the present specimens the height is variable, but the majority of tests are distinctly conical and only occasional ones are flattened. They agree very well with Haeusler’s figures, and on the basis of growth plan and aperture it seems best to refer the species to Trochammina. The species clearly differs from Valvulina triangularis Parker and Jones, a recent form, and Galloway’s usage of the name is otherwise adhered to. 290 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY TROCHAMMINA GLOBIGERINIFORMIS (Parker and Jones) 1863 Plate 1, figures 23-24 Haplophragmium globigeriniformis, Haeusler (non Parker and Jones), 1890, p. 36, figs. 193-195. Trochammina globigeriniformis, Cushman, 1920, p. 78, pl. 16, figs. 5-6 (1918-31). Trochammina globigeriniformis, Seibold and Seibold, 1953, p. 46, text figs. 5-6. There are three or four chambers in the last whorl and the aperture is not visible. The amount of cement is variable, and some specimens are distinctly arenaceous while others are almost completely calcareous. The chambers are generally bulbous, but occasionally they are compressed, probably due to squashing. The Jurassic forms are generally smaller than the recent counterparts, but otherwise they are morphologically identical. This appears to be an example of a very long-ranged species. Specimens are rare, but occur sporadically throughout the Bathonian. TROCHAMMINA sp. Plate 1, figure 20 The material consists of two specimens which resemble T'ro- chammina globigeriniformis, but are less trochoid and are flatter. They are also similar to Trochammina canariense (d’Orbigny ) figured by Haeusler (1890, p. 34, pl. 4, figs. 1-3), but these are broader and have fewer chambers in the last whorl. Family LAGENIDAE Genus LENTICULINA Lamarck, 1804 LENTICULINA GALEATA (Terquem) Plate 2, figures 15-17 Cristellaria galeata Terquem, 1870a, p. 444, pl. 16, fig. 15a-b. Cristellaria subangulata, Bruckman, (non Reuss), 1904, p. 22, pl. 3, fig. 9. Test smooth, finely perforate, slightly involute, generally bi- convex with slight depression in umbilical area, some specimens triangular in cross-section; periphery rounded, mostly close coiled, but occasionally uncoiled in later portion; chambers CIFELLI: BATHONIAN FORAMINIFERA OF ENGLAND 291 numerous, six to ten in the last whorl, increasing in size as added, last ones inflated; sutures deeply depressed, with moderately strong ribs along the margins which extend to the keel; well developed keel throughout but indented at the sutures; aperture radiate, terminal, projecting slightly at peripheral margin. Most of the tests are biconvex, but some have practically parallel sides, while others are triangular in cross-section, as in the genus Saracenaria. The ribbing along the margins of the sutures also exhibits some modification. Most often the ribs are quite strong, but occasionally they are weakly developed, and on one specimen observed they were completely inconspicuous. Oc- casionally a weak cireular rib is developed, as in L. quenstedti, and except for the depression of the sutures such specimens are difficult to distinguish from the latter species . Terquem’s figure of this species does not show the ribs along the margins of the depressed sutures. At the Muséum d’Histoire Naturelle, Paris there are two specimens of this species in the Terquem collection, and ribs are well developed in both. This species is common in the Lower Fuller’s Earth Clay, but isolated specimens appear throughout the remainder of the Bathonian. LENTICULINA MULTANGULOSA (Schwager ) Plate 2, figure 14 Cristellaria multangulosa Sehwager, 1865, p. 126, pl. 6, fig. 3. Lenticulina (Astacolus) multangulosa, Seibold and Seibold, 1953, p. 56, OLS ye aaah) DY The angular periphery of this species is characteristic. Schwa- ger’s figure shows six chambers, and the last two are decidedly uncoiled, arranged in a linear series. Otherwise the present specimens agree well with the original figure. LENTICULINA MUNSTERI (Roemer) Plate 2, figures 3-5 Robulina munsteri Roemer, 1839, p. 48, pl. 20, fig. 29. Cristellaria inflata Schwager, 1865, p. 125, pl. 6, fig. 16. 292 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY Cristellaria (Lenticulina) munsteri, Bartenstein and Brand, 1937, p. 174, pl. 3, fig. 30a-b; pl. 4, fig. 69a-e; pl. 6, fig. 34a-d; pl. 9, fig. 49a-e; pls WO; fies (o8a-b;) spl lllat tice 3a-di) ple dll, fie. 9 a-dis splyvdligar fig. 16a-b; pl. 12b, fig. 15a-e; pl. 13, fig. 36; pl. 14b, fig. 14a-e; pl. 14e, fig. 13a-b; pl. 15a, fig. 34a-c; pl. 15c, fig. 19a-c. Cristellaria (Astacolus) matutina, Bartenstein and Brand (non d’Orbigny ) 1937, p. 172 (pars), pl. 4, fig. 79; pl. 5, fig. 53; pl. 6, fig. 33. Cristellaria munsteri, Macfadyen, 1941, p. 31, pl. 2, fig. 23a-b. Cristellaria matutina, Macfadyen, 1941, p. 30, pl. 2, fig. 22. Lenticulina munsteri, Barnard, 1950b, p. 7, pl. 2, fig. 1. Lenticulina (Lenticulina) munsteri, Seibold and Seibold, 1955, p. 104, text fig. 4a-c. Lenticulina (Astacolus) matutina, Seibold and Seibold, 1955, p. 108, text fig. 4k-]. Many specimens have a slight keel, the presence of which appears to be determined partially by preservation. Possibly the Jurassic forms referred to as Lenticulina cultrata (Montfort) should be included here, as this species is distinguished from L. munsteri only by the presence of the keel. However, the keels on the English Bathonian specimens are always small, and never highly developed as in the described forms of L. cultrata. This species occasionally exhibits a marked tendency to uncoil, and in some assemblages there occur transitions from the typical, tightly coiled types to linear forms with a small spire. In the uncoiled individuals the sutures tend to become de- pressed in the last stages, and the last chambers are sometimes bulbous. The uncoiled individuals are identical with some of those of Lenticulina matutina (d’Orbigny), and probably many specimens identified as this latter species are actually variants of L. munsteri. , LENTICULINA QUENSTEDTI (Gumbel) Plate 2, figures 6-7 Cristellaria quenstedti Gumbel, 1862, p. 226, pl. 4, fig. 2a-b. Jristellaria polonica Wisniowski, 1890, p. 222, pl. 10, fig. 3. Cristellaria quenstedti, Klahn, 1921, p. 49, pl. 2, figs. 16, 18-25. Cristellaria quenstedti, Paalzow, 1932, p. 102, pl. 6, figs. 3-5. Cristellaria (Lenticulina) quenstedti, Bartenstein and Brand, 1937, p. 177, pl. lla, fig. 16a-c; pl. 11b, fig. 23a-c; pl. 12a; fig. 19a-d; pl. 12b, fig. 7a-c; pl. 13, fig. 39a-c; pl. 14, fig. 17a-b; pl. 15a, fig. 36a-c; pl. 15e, fig. 20a-b. CIFELL1: BATHONIAN FORAMINIFERA OF ENGLAND 293 Lenticulina quenstedti Barnard, 1952, p. 339, text fig. A-6. Lenticulina (Lenticulina) quenstedti Seibold and Seibold, 1955, p. 105, pls W3y figs 3. This common Jurassic species has been deseribed many times. Most characteristic are the raised sutural ribs which converge at the umbilical area and are connected by a circular rib. The speci- mens figured by Bartenstein and Brand have ribs which are beaded, and differ in this respect from those recorded here and most other described forms of this species. The difference is not considered taxonomically significant. The ribbing, however, is variable and Bartenstein and Brand (1937, p. 177) note that in the Middle Dogger of Germany this species appears to be transi- tional to Lenticulina tricarinella (Reuss). In the present ma- terial some specimens lack a clearly defined cireular rib, and have sharp sutural ribs extending to the periphery. Such specimens are very similar to L. tricarinella, but no actual transi- tion was observed, as the forms included here consistently lack the lateral keels of L. tricarinella. This species is common in the English Bathonian, appearing at the base of the Lower Fuller’s Earth Clay, which is the earliest recorded occurrence in England. LENTICULINA QUENSTEDTI (Reuss) var. A n.var. Plate 2, figures 9-10 Test large, compressed, with rugose surface; periphery rounded, with distinct keel; chambers numerous, seven or eight visible in the last whorl, inereasing in size as added; sutures gently curved, raised along the greater part of their length, and meeting at umbonal area; oblique costae weak and poorly de- veloped near peripheral margin in last part of test; aperture peripheral, radiate, slightly projecting. The test is larger, and the surface is more rugose than in the typical forms. The circular rib is not as well developed, and oblique costae are generally present in the later chambers, which was not observed in Lenticulina quenstedtt. The distribution of this variety appears to be restricted. It was recovered only from the Lower Fuller’s Earth Clay at Dorset and at Midford, in the Bath area. 294 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY LENTICULINA QUENSTEDTI (Reuss) var. B n.var. Plate 2, figure 8 Test stout, shghtly biconvex, involute, slightly longer than broad; periphery sub-rounded ; chambers numerous, six to eight visible in the last whorl, increasing in size as added; sutures curved, strongly limbate, extending from periphery to umbilical area; thin keel, not visible on all specimens; umbilical area depressed, partially filled with beads of calcite, sometimes with partially developed circular rib connecting sutures; aperture radiate, projecting slightly from peripheral margin. This variety differs from the typical forms in being more involute and robust. The circular rib connecting the sutures is much less developed, and the sutures are generally more limbate. The variety occurs sporadically throughout much of the Bath- onian. It is most common in the Wattonensis Beds at Dorset. LENTICULINA SUBALATA (Reuss) Plate 2, figures 1-2 Cristellaria subalata Reuss, 1854, p. 68, pl. 25, fig. 13. Cristellaria casis, Jones and Parker (non Fitchel and Moll), 1860, p. 457, pl. 20, fig. 41. Cristellaria helios Terquem, 1870a, p. 445, pl. 16, figs. 19-21. Cristellaria subalata, Franke, 1936, p. 115, pl. 11, fig. 19. Cristellaria (Lenticulina) subalata, Bartenstein and Brand, 1937, p. 176, pl. 6, fig. 35a-c; pl. 9, fig. 54a-e; pl. 10, fig. 41a-b; pl. lla, fig. 15a-b; pl. 11b, fig. 22a-b; pl. 12a, fig. 18a-c; pl. 12b; fig. 16; pl. 18, fig. 38a-c; pl. 14b, fig. 16a-b; pl. 15a, fig. 35a-b. Cristellaria (Lenticulina) subalata form A Bartenstein and Brand, 1937, p. 177, pl. 9, fig. 55a-f; pl. 10, fig. 42a-b. This species is similar to LZ. munsteri, but has raised and generally broader sutures. It also resembles Lenticulina varians (Borneman), but besides having the raised sutures, is more biconvex and rounder in cross-section than is this latter species. Specimens of Cristellaria helios in the Terquem collection are identical with the present ones. However, other identical speci- mens in the collection are scattered among various dissimilar species of Cristellaria (sensu lato) type, some bearing little CIFELLI: BATHONIAN FORAMINIFERA OF ENGLAND 295 resemblance to the figures which are supposed to illustrate them. Consequently, a comprehensive synonymv is difficult. The type of this species is from the Cretaceous of the eastern Alps, but the Jurassic forms appear to be morphologically iden- tical. It oceurs commonly throughout the English Bathonian. LENTICULINA SOWERBYI (Schwager) Cristellaria sowerbyi Schwager, 1867, p. 660, pl. 34, fig. 18. Cristellaria sowerbyi Deeke, 1886, p. 322, pl. 2, fig. 34. In some specimens the later chambers overlap on one side, producing a test that is trochoid, and similar to the growth plan that is found in Darbyella. The sutures are most generally flush, but they tend to become depressed in the later chambers, a tendency noted in many lagenid species. LENTICULINA TRICARINELLA (Reuss) Plate 2, figures 20-23 Cristellaria (Cristellaria) tricarinella Reuss, 1863a, p. 68, pl. 7, fig. 9; pl. 12, figs. 2-4, Cristellaria polymorpha Terquem, 1870a, p. 454, pl. 19, figs. 1-30; pl. 21, figs. 1-30. Cristellaria tricarinella, Paalzow, 1917, p. 240, pl. 56, fig. 6. Cristellaria tricarinella, Klahn, 1921, p. 50, pl. 21, figs. 7-10. Planularia feifeli Paalzow, 1932, p. 105, pl. 6, figs. 11, 12. Cristellaria tricarinella, Macfadyen, 1935, p. 15, pl. 1, fig. 18a-b. Cristellaria (Astacolus) tricarinella, Bartenstein and Brand, 1937, p. 173, pl. 13, fig. 35a-b; pl. 14b, fig. 13a-b; pl. 15a, fig. 33a-b; pl. 15e, fig. 18. Cristellaria tricarinella, Frentzen, 1941, p. 353, pl. 5, figs. 13, 14. Lenticulina (Planularia) tricarinella, Seibold and Seibold, 1958, p. 54, pl. 4; fig. 5: This common and distinctive species has been either described or recorded many times from the Jurassic. The sharp, sutural ribs joining the lateral keels are characteristic. Occasionally, the lateral keels are incompletely developed, appearing instead as impersistent costae along the margins of the test. Typically, the tests are tightly coiled, with only one or two chambers ex- tending beyond the spire. In occasional assemblages, however, all degrees of coiling occur, and in some tests the chambers are arranged in linear or curved series, with the spiral part forming 296 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY a small, minor portion of the test. Some of the linear types tend to become triangular in eross-section and except for their mutual relationships with the rest of the specimens could be confused with Saracenaria. On some specimens, faint, oblique costae are present between the sutural ribs of the last two chambers. The number and strength of these are very variable, and they most generally occur on those specimens which lack the clearly defined lateral keels. I have examined a large number of specimens of Cristellaria polymorpha Terquem in the Terquem collection, and agree with Macfadyen (1935, p. 15) that this species is a synonym of Lentt- culina tricarinella (Reuss). The intricate scroll pattern illus- trated by Terquem was not observed on any of the specimens. It was observed, however, that when the slides of the Terquem collection were viewed from the underside, irregular streaks in the mounting medium appeared to cover many of the specimens in intricate, scroll-like patterns. Possibly Terquem confused these patterns with ornamentation. Franke (1936) illustrated a specimen of Cristellaria poly- morpha Terquem with peculiar markings on the wall which, though less elaborate, are similar to Terquem’s scroll-like pat- terns. His figure otherwise appears to be identical with Lenti- culina tricarinella (Reuss). The markings may be due to foreign matter on the test, or possibly they may be indistinct, oblique costae. This species has not been previously described from England. It occurs in the lowermost Fuller’s Earth Clay, and the Lower Bathonian is the earliest known Enelish occurrence. This species is a common and characteristic form in the Middle Jurassic of Europe, and Bartenstein and Brand (1937, p. 173) record the earliest occurrence in northwest Germany from the Parkinsoni Schichten (Upper Bajocian). Jf Franke’s Cristellaria poly- morpha (recorded from the Lias Gamma) is a synonym, the range in Germany would extend into the Lower Jurassic. Barten- stein and Brand, however, do not consider Franke’s form a synonym. CIFELLI: BATHONIAN FORAMINIFERA OF ENGLAND 297 LENTICULINA TRICARINELLA (Reuss) var. A n.var. Plate 2, figure 24 Test small, compressed, finely perforate, incompletely coiled, three to five chambers arranged in gently curved series, increas- ing in size as added; keel single, well developed, extending with- out interruption over entire peripheral margin; striae absent or faintly developed in last chambers; sutures raised in sharp ridges, connecting with keel; aperture radiate. This variety lacks the marginal keel and is generally smaller than typical members of the species. It occurs rarely in the Upper Fuller’s Earth Clay and Wattonensis Beds of the Dorset Coast. LENTICULINA TURGIDA (Schwager) Plate 2, figures 18-19 Cristellaria turgida Schwager, 1865, p. 127, pl. 6, fig. 4. Cristellaria informis Schwager, 1865, p. 128, pl. 6, fig. 8. Cristellaria inflata, Wisniowski (non Schwager), 1890, p. 227, pl. 10, fig. 13a-b. Lenticulina (Astacolus) matutina informis, Seibold and Seibold, 1956, p. 119, pl. 7, fig. 13; text fig. 4e-g. There is one microsphere in the present material, and the specimen is larger than the others. It consists of eight chambers, and the last one overlaps on one side, as in Darbyella. There is good agreement between specimens included here and the figures of Seibold and Seibold who studied topotype material from which Schwager originally described the species. The English Bathonian forms, however, show less tendency to uncoil. LENTICULINA VARIANS (Borneman) Plate 2, figures 11-13 Cristellaria varians Borneman, 1854, p. 41, pl. 4, figs. 32-34. Cristellaria inquisita Terquem, 1870a, p. 444, pl. 16, figs. 16-18. Cristellaria (Lenticulina) varians Borneman var. recta Franke, 1936, p. 113, pia fies W2: Cristellaria (Lenticulina) varians, Bartenstein and Brand, 1937, p. 176, pl. Tayo US epla 2a nies) NG 20s placse fe wole-s pla oe Tos GOpeplakss fig. 53a-b. 298 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY Cristellaria (Lenticulina) varians form A Bartenstein and Brand, 1937, p. 176, pl. 10, fig. 40a-d; pl. lla, fig. 14a-d; pl. 11b, fig. 2la-b. This species most closely resembles Lenticulina munsteri (Roemer), and is distinguished by the more compressed nature of the test and the narrower, shghtly depressed sutures. Borneman’s figures are not too clear, nor do they indicate the possible variation of this species. Consequently, various interpre- tations are possible and there are a variety of forms in the litera- ture bearing this name, many of which are distinct from those included here. Synonymies are uncertain and the types need to be re-studied. Terquem showed only outline drawings of Cristellaria in- quisita, but the specimens in the Terquem collection are identical with the ones recorded here. Genus PLANULARIA Defrance, 1824 PLANULARIA ANCEPS (Terquem) Plate 3, figures 14-15 Cristellaria anceps Terquem, 1870a, p. 428, pl. 9, figs. 11-21. Cristellaria dolium Terquem, 1870a, p. 431, pl. 9, fig. 29a-b. There is one microsphere in the present material, and it is identical with the megalospheres, except that the large proloculus is represented by several small chambers which form an incom- plete coil. Terquem illustrated a variety of specimens of this species, including forms with depressed sutures and a globular prolocu- lus. He later extended the species (Terquem, 1883, p. 350, pl. 38, fies. 18-28; pl. 39, figs. 1-2) to include a variety of forms which bear very little resemblance to the earlier figures. Many of the later figures seem to represent aberrant or immature specimens, and probably belong to other species. Even as originally de- scribed, the species probably consists largely of an unrelated eroup of specimens. However, there is good agreement between my specimens and some of Terquem’s figures (particularly pl. 9, fig. 12), as well as the specimens from the Terquem collection, so that there is at least partial identity. CIFELLI: BATHONIAN FORAMINIFERA OF ENGLAND 299 PLANULARIA BEIERANA (Gumbel) Plate 3, figures 1-10; text figure 3. Marginulina beierana Gumbel, 1862, p. 221, pl. 3, fig. 20a-b. Cristellaria jurassica Gumbel, 1862, p. 224, pl. 3, fig. 25a-e. Cristellaria semiinvoluta Terquem, 1870a, p. 435, pl. 11, figs. 1-30; pl. 12, figs. 1-30. Cristellaria suturalis Terquem, 1870a (non Terquem, 1866a), p. 434, pl. 10, figs. 19-24, Cristellaria semiinvoluta var. plana Deeke, 1886, p. 317, pl. 2, fig. 35. Planularia semiinvoluta, Paalzow, 1932, p. 105, pl. 6, figs. 18, 14, 20, 21. Cristellaria (Planularia) cordiformis, Bartenstein and Brand (non Ter- quem), 1937, p. 169, pl. 6, fig. 30a-b; pl. 9, fig. 48a-e; pl. 10, fig. 30a-d; pl. 12a, fig. 10; pl. 12b, fig. lla-b; pl. 13, fig. 29a-c; pl. 14b, fig. 10; pl. 15a, fig. 28a-b. Cristellaria (Planularia) crepidula, Bartenstein and Brand (non Fitchel and Moll), 1937, p. 169, pl. 2a, fig. 18a-b; pl. 3, fig. 48; pl. 4, fig. 76; pl. Hy sales VO jolly Oy ike Cope soll AlOy item Gul joe Ill sae IAT ol eaeloy fig. 16a-b; pl. 12a, fig. 12a-d; pl. 12b, fig. 12a-b; pl. 18, fig. 31; pl. 14b, fig. 11; pl. 14e, fig. 12; pl. 15a, fig. 30; pl. 15c, fig. 1da-b. Cristellaria (Planularia) filosa, Bartenstein and Brand (non Terquem), 1937, p. 169, pl. 4, fig. 77; pl. 9, fig. 44; pl. lla, fig. 15a-b; pl. 12a, fig. 1; pl. 113, fig. 315) pl. 15a, fig. 29a-cs pl. Use; fig. 4a-b- Lenticulina (Planularia) cordiformis, Bartenstein, 1948, pl. 2, figs. 13-15. Lenticulina (Planularia) crepidula, Bartenstein, 1948, pl. 2, figs. 20-21. Falsopalmula deslongchampsi, Bartenstein (non Terquem), 1948, pl. 1, figs. 6-H; pl. 9258 figs 10, Falsopalmula obliqua, Bartenstein (non Terquem), 1948, pl. 2, figs. 11-12. Lenticulina (Planularia) beierana, Seibold and Seibold, 1955, pl. 13, fig. 7; text fig. 4e, f. Test small, much compressed, with thin, transparent wall; chambers, numerous, five to eleven in number, first four or five arranged in a loose coil, remainder arranged in a linear or gently curved series, last ones sometimes equitant; proloculus in megalosphere large, suboval, generally subparallel to longi- tudinal axis of test; sutures flush, depressed or sometimes raised in form of marginal ribs; aperture terminal, radiate. Microspheres are rare, occurring in about one out of eight specimens. They do not differ from the megalospheres, except that the proloculus is smaller, and there are a greater number of chambers. The size of the proloculus is quite variable, but is always large in relation to the size of the test. There is no 300 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY evidence to suggest trimorphism within this species, as the prolocular size differences are transitional among the mega- lospheres. The shape of the test is highly variable, and is related to the degree of overlap of the chambers on the prolocular area. This relationship has been previously deseribed in Planularia pau- perata Jones and Parker by Barnard (1950b, p. 11). The raised sutures were observed only on specimens from the Wattonensis Beds, where this species exhibits an extremely high degree of variability. All gradations from depressed to flush, to limbate, can be observed. Not only is this character variable from specimen to specimen, but in a single individual all three types of suture may be found. The transitional nature of the typical uncoiled growth plan to the equitant or flabelline chamber form is illustrated in Figure 3A-N. All specimens were recovered from a single sample from the Wattonensis Beds on the Dorset Coast. The equitant cham- bers vary in number, size and degree of symmetry, reflecting a high instability of the character. In some instances the change in growth plan is marked (Figure 31), while in others it is accomplished gradually, by progressive overlap of the later chambers on the oral margin (Figure 3G). Growth is sometimes irregular, and in Figure 3K the ninth chamber partially reverts to the lnear arrangement by failing to overlap on the oral margin. In Figure 3N the equitant chambers have developed before the test has unecoiled, and the longitudinal axis of the test 1s shifted towards the adoral margin. This specimen is also unusual in that the proloculus is elongated in a direction approxi- mately perpendicular to the longitudinal axis of the test, con- trary to other specimens in which the proloculus is elongate parallel to the long axis of the test. This species is highly variable in almost all observable mor- phological characters, and the problem of deciding which specific name to use is difficult. All of the diverse types illustrated here are connected by integradine forms, and there appears to be no logical way of separating them on the basis of morphology or distribution. They are therefore regarded here as a single, highly variable species. I have examined topotypes of Planularia beierana, and there is good agreement with these specimens, BATHONIAN FORAMINIFERA OF ENGLAND 301 CIFELLI : W ‘yosiog ‘speq sisuauojjn yy ‘Furgpod e[duvs wo1Z 318 suoutoods [Ty ‘“(jequiny) vuUnsLa12q DiWojNUD)T JO WOTZPVIIBA “FE dINSLT i a d y 302 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY except that the topotypes tend to be broader, and the later chambers generally overlap more on the prolocular area. They may possibly form a link with Planularia pauperata Jones and Parker. The specimens of Planularia pauperata from Chelaston, Derby, in the Jones and Parker collection at the British Museum (Natural History) appear to be distinct from the present ones. They are generally more robust, broader, and the chambers show more overlap. There are only three specimens in the collection, and possibly a more numerous suite would show transitions to the present ones. However, the specimens of P. pauperata from the Lias of Byfield described by Barnard (1950b, p. 10, pl. 2, fig. 7) are identical with those from Chelaston, Derby, and are distinct from the Bathonian forms. Cristellaria semiunvoluta Terquem appears to represent a heterogeneous group of specimens. Unfortunately, many of the specimens of this species in the Terquem collection do not agree well with the figures that presumably illustrate them, and with- out the original types it is not possible to properly analyze this species. There is no doubt, however, that in part, at least, this species is a synonym of Planularia beierana. The striate forms do not seem to belong here, but this cannot be determined until the types have been recovered, or the type locality re-examined. The large number of forms illustrated by Bartenstein and Brand (and included here in this species) do not appear distinct to me, even on the basis of those authors’ material. The variational pat- terns of P. beierana in northwest Germany are similar to the ones encountered in the English Bathonian. However, the form described as Cristellaria (Planularia) cordiformis Terquem may possibly be transitional to P. pauperata. Forms identical with the ones described by Bartenstein as Falsopalmula deslong- champsi (Terquem) and Falsopalmula obliqua (Terquem), are included here because of the transitional nature and intimate association of these identical forms with typical forms of Planularia beierana, in the Enelish Bathonian. This species is common in the English Bathonian, and indi- viduals are generally numerous. The variational patterns are sporadic, but by far the greatest ‘‘bursts’’ occur in the Watton- ensis Beds at Dorset. Here the sutures show all gradations from distinetly depressed to rather prominent marginal ribs. The CIFELLI: BATHONIAN FORAMINIFERA OF ENGLAND 303 shapes are highly variable, and equitant chambers are more common than at any other locality. Equitant chambers were also observed in specimens from the Lower Fuller’s Earth Clay at Dorset and Midford, and from the Bradford Clay in the Monk quarry at Corsham. PLANULARIA BREONI (Terquem) Plate 3, figure 11 Cristellaria breoni Terquem, 1864a, p. 420, pl. 9, figs. 4a, b. Cristellaria (Astacolus) breoni, Bartenstein and Brand, 1937, p. 173, pl. 4, figs Moist pls jos atle. (06; Cristellaria (Astacolus) radiata, Bartenstein and Brand (non Terquem), 1937, p. 172, pl. 5, fig. 58; pl. 12b, figs. 14a-c; pl. 13, figs. 34a-b. Cristellaria (Astacolus) quadricostata, Bartenstein and Brand (non Ter- quem), 1937p. 173, pl. 3) fig. 47; pl. 4, fig. 715 ple 5, tes of. This species resembles Planularia eugeni (Terquem) in its numerous striations which completely cover the sides of the test. However, P. breoni is more rounded in cross-section, has broader chambers and the sutures are more depressed. PLANULARIA EUGENIL (Terquem) Plate 3, figures 16-17 Cristellaria eugenti Terquem, 1864a, p. 414, pl. 9, fig. 16a-b. Cristellaria arictis Issler, 1908, p. 81, pl. 5, figs. 255-260, Cristellaria crepidula (Fitchel and Moll) var. striata Issler, 1908, p. 82, pl. 5, figs. 255-266; pl. 6, figs. 266-268. Cristellaria (Planularia) eugenii, Bartenstein and Brand, 1937, p. 170, pl. 3, fig. 56; pl. 10, fig. 33a-b; pl. 12a, fig. 13. One microsphere was observed, and it differed from the meg- alospheres only in the greater number of chambers in the coiled portion of the test. The species is highly variable in shape, with specimens ranging from elongate with parallel margins, to broad, stubby ones in which the margins diverge. The number of ribs range from ten to twenty and in strength from faint striations to prominent ribs. This species is similar to Planularia beterana (Gumbel), but is distinguished by the presence of ribs and the shape of the chambers which are much lower in relation to the breadth than in the latter species. 304 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY Terquem described this species from the Lias. The one speci- men available from the Terquem Lias collection in the Muséum d’Histoire Naturelle is much larger and more robust than the present ones, but otherwise they compare well, and I consider them identical. Terquem included many striate forms in Cris- tellaria semunvoluta and Cristellaria subinvoluta and possibly many of these belong here as well. PLANULARIA INCONSTANS (Schwager) Cristellaria inconstans Schwager, 1867, p. 658, pl. 34, fig. 15. Cristellaria dialatata Wisniowski, 1890, p. 211, pl. 9, fig. 10. Cristellaria dorsoarcuata Wisniowski, 1890, p. 211, pl. 9, fig. 11. Cristellaria (Astacolus) inconstans, Bartenstein and Brand, 1937, p. 171; pl. 10, fig. 34a-e. This species resembles Planularia protracta (Borneman), but can be distinguished by the broader, lower chambers and the absence of a coil. Specimens are rare and were recovered only from the Wat- tonensis Beds. Material deposited at M.C.Z. No. 3333. PLANULARIA PROTRACTA (Borneman) Plate 3, figures 12-13 Cristellaria protracta Borneman, 1854, p. 39, pl. 4, fig. 27a-b. Cristellaria lista Borneman, 1854, p. 39, pl. 4, fig. 28. Cristellaria protracta, Macfadyen, 1941, p. 32, pl. 2, fig. 24. This species is similar to Planularia beierana (Gumbel), but is distinguishable by the more oval, less compressed test and by the much greater curvature of the peripheral margin at the base of the test. PLANULARIA sp. A Test compressed, shghtly twisted; thin walled, transparent ; chambers numerous, the first three forming incomplete coil, the remaining four arranged in a linear series, increasing regularly and rapidly in height as added, moderately in breadth; sutures depressed, particularly in later chambers; aperture terminal, radiate. CIFELLI: BATHONIAN FORAMINIFERA OF ENGLAND 305 The specimens are similar in outline to Planularia pauperata Jones and Parker, but the test is twisted, and the last chambers tend to be bulbous. The material consists of three specimens from three separate localities in the Upper Fuller’s Earth Clay. Material deposited at M.C.Z. No. 3335. PLANULARIA sp. B Plate 3, figure 18 Several poorly preserved specimens are included here which consist of linear tests with low, broad chambers, arched in the early portion, but not actually coiled. The sutures are depressed and slightly curved. The shapes vary from broad, short forms, to long, slim ones. The most distinctive feature is the presence of an elevated border on both margins of the test, against which the sutures abut. This feature is characteristic of Planularva arguta Reuss, a Cretaceous form. The present specimens are similar to fragments identified as this species and described by Blake (1876, p. 464, pl. 19, fig. 10), from the Lias of Yorkshire. The Bathonian material, however, is poorly preserved, and the raised margins, which are not present on all of the specimens, may be due to squashing in the central portions of the tests. The forms resemble Vaginulina legumen (Linné), but are smaller and flatter ; in side view they tend to be more triangular. The specimens are rare, and oceur only in the Cotswolds, Oxford and Northampton areas. Genus MARGINULINA d’Orbigny, 1826 MARGINULINA DESLONGCHAMPSI Terquem Plate 3, figure 21 Marginulina deslongchampsi Terquem, 1864a, p. 188, pl. 8, fig. 8a-b. Marginulina oolithica, Franke (non Terquem), 1936, p. 74, pl. 7, fig. 17a-b. Marginulina oolithica, Bartenstein and Brand, 1937, p. 160, pl. 2a, fig. 11; Diy hy wigs 2 te: pls 3, fie. sosepls 6} digs dines ples, fee 2Oa-bipolemllO fig. 24. Specimens from the Enghsh Bathonian are somewhat less curved in the initial portion of the test than are shown in the figures of Terquem, but are still distinctly marginuline, and otherwise agree well with the figures. 306 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY The shape of the chambers is variable and in some specimens the chambers are bulbous and twisted slightly from the longi- tudinal axis. The degree of curvature is also variable, and some tests are almost dentaline. This species is rare, and occurs only in the Lower Fuller’s Earth Clay and Wattonensis Beds on the Dorset Coast. MARGINULINA TERQUEMI d’Orbigny Plate 3, figures 19-20 Marginulina terquemi, Terquem (non d’Orbigny), 1858, p. 609, pl. 3, fig. la-e. Marginulina glabra, Haeusler (non d’Orbigny), 1890, p. 106, pl. 14, figs. 35-40, 42-43. Marginulina glabra, Bartenstein and Brand, 1937, p. 160, pl. 10, fig. 25a-b; plo 1b; fie? 1s= pil 13, fig. W6a-b. Specimens included here vary considerably in the height of the coil, and the shape of the cross-section. The chambers are less bulbous and the tests are slimmer than in Terquem’s figures, but are otherwise identical. This species is very similar to Marginulina glabra d’Orbigny, which is a Recent form, and perhaps identical. Genus DENTALINA d’Orbigny, 1826 DENTALINA BICORNIS Terquem Plate 4, figure 7 Dentalina bicornis Terquem, 1870b (pars), p. 370, pl. 29, figs. 13-16 (non uifeay DEP) e Dentalina bicornis, Bartenstein and Brand, 1937, p. 138, pl. 10, fig. 17a-e; pl. 15a, fig. 8. Dentalina bicornis, Seibold and Seibold, 1953, p. 59, pl. 4, fig. 15; pl. 5, soieas Af This species is rare, and was observed only in the Kemble Beds at Oxfordshire. DENTALINA COMMUNIS d’Orbigny Plate 4, figure 11 Dentalina communis d’Orbigny, 1826, p. 254, no. 35. Dentalina pseudocommunis Franke, 1936, p. 30, pl. 2, fig. 20a-b. CIFELLI: BATHONIAN FORAMINIFERA OF ENGLAND 307 Dentalina ventricosa Franke, 1936, p. 33, pl. 3, fig. la-b. Dentalina communis, Bartenstein and Brand, 1937, p. 136, pl. la, fig. 6; pl. 1b, figs. 14-15; pl. 2a, fig. 5; pl. 2b, figs. 9-10; pl. 3, fig. 8; pl. 4, fig. 23a-b; pk 5, fig. 13; pl, 6, fig. 10: Dentalina ventricosa, Bartenstein and Brand, 1937, p. 140, pl. 4, fig. 20; ple .o, ties iD: Dentalina communis, Macfadyen, 1941, p. 39, pl. 2, fig. 34. Dentalina pseudocommunis, Barnard, 1950b, p. 19, pl. 3, figs. 6, 7. This species was originally deseribed from Recent sediments of the Mediterranean, and there is much disagreement about the systematic position of the Jurassic forms. Macfadyen studied Recent forms from the Mediterranean and concluded that those from the Jurassic are identical. Barnard found the Jurassic forms to be highly variable, and referred his specimens to D. pseudocommunis Franke, as he felt it unlikely that such a variable species could have such a long range in time. Brotzen (1953, p. 35) does not believe that the Jurassic forms are closely related to the modern ones, suggesting that they may not even belong to the genus Dentalina. I agree with Macfadyen, and can see no morphological distinction between the Recent and the Jurassic forms. It is possible that D. communis is a morpho- logieal type of polyphyletie origin occurring in Mesozoic and Cenozoic strata, but the lineages have yet to be demonstrated. Material deposited at M.C.Z. No. 3340B. DENTALINA aff. D. communis d’Orbigny Plate 4, figures 14-15 Test small, arched, oval in cross-section, with four to six chambers, higher than wide, increasing in size as added, early ones irregular in shape, last ones much larger than preceding and bulbous; sutures depressed, only slightly oblique; prolocu- lus small, spherical; aperture central, radiate. These specimens are similar to Dentalina communis d’Orbigny but differ in their smaller size, less oblique sutures and more central position of the aperture. In addition, the early chambers are characterized by irregular growth, and the shape and size of these are very variable. The relationships of this form are uncertain, and the resem- blanees with D. communis are probably superficial. The speci- mens recorded here appear to belong to a distinct species, but 308 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY they are rare, occurring less than three to a sample, so that they cannot be adequately described. The distribution is of some interest. Although specimens are rare they occur at a number of localities in the Upper Fuller’s Karth Clay, Bradford Clay and Forest Marble on the Dorset Coast and the Bath area. They have not been recovered from lower horizons. DENTALINA CONFERTA Schwager Plate 4, figure 3 Dentalina conferta Schwager, 1865, p. 106, pl. 3, fig. 9. Dentalina bullata Schwager, 1865, p. 107, pl. 3, fig. 23. Dentalina brevis, Issler (non d’Orbigny), 1908, p. 63, pl. 3, figs. 150-154. Dentalina linearis, Issler (non Roemer), 1908, p. 64, pl. 3, fig. 155. Dentalina suboligostegia Franke, 1936, p. 25, pl. 2, fig. 1. Dentalina numismalis Franke, 1936, p. 25, pl. 2, fig. 4. Dentalina bullata, Seibold and Seibold, 1956, p. 128, pl. 7, fig. 8; text fig. 6a-b. Dentalina conferta, Seibold and Seibold, 1956, p. 129, pl. 7, fig. 14; text fig. 5y. This species is rare, and occurs only in the Wattonensis Beds at the Dorset Coast. DENTALINA INTORTA Terquem Plate 4, figures 16-19 Dentalina intorta Terquem, 1870b, p. 364, pl. 27, figs. 26-34. Dentalina ventricosa, Bartenstein and Brand (non Franke), 1937, p. 140, pl, 4 oie. 20: spliwo,eaic: ollp. Dentalina lateralis, Bartenstein and Brand (non Terquem), 1937, p. 135, pl. 5, fig. 16a-b. This species is highly variable in shape and size of the test. Most of the specimens are short and broad, but these grade into individuals that are long and sim. The proloculus varies in shape from globular, equidimensional types to long, narrow ones. In all specimens observed the proloculus is large, and no micro- spheres were recognized. The later chambers tend to become bulbous and overlap the preceding ones on the convex side. In some specimens the axis of the last chambers tends to rotate from CIFELLI: BATHONIAN FORAMINIFERA OF ENGLAND 309 the axis of the preceding ones, sometimes as much as ninety degrees. This results in a plan of growth that is similar to species included with the Polymorphinidae. These specimens are identical with those figured by Terquem and the specimens in the Terquem collection in Paris. One of the specimens in the collection shows a shift on the axis of the last chamber from the preceding ones, such as has been described above. This species is common in the English Bathonian. In north- west Germany, identical forms described by Bartenstein and Brand are recorded from the Lias. DENTALINA MUCRONATA Neugeboren Plate 4, figures 1-2; text-figure 4 Dentalina mucronata Neugeboren, 1856, in Ellis and Messina, Catalogue of Foraminifera. Dentalina cornuformis Terquem, 1870b, p. 367, pl. 28, fig. 16. Dentalina turgida, Wisniowski (non Schwager), 1890, p. 201, pl. 1, fig. 48. Dentalina mucronata, Bartenstein and Brand, 1937, p. 138, pl. 10, fig. 16; pl. 11b, fig. 5a-b; pl. 12b, fig. 4a-b. The shape of the test is highly variable. Most individuals are moderately broad, but some are long and slender while others are stout. Some of the broad forms are sli¢htly arched at the base and have low, broad proloculi. Such specimens are morpho- logically indistinguishable from Vaginulina legumen (Linné), and ean be identified only by their association with transitional and more typical forms in a single assemblage. The variation and resemblance of ‘‘end forms’’ to Vaginulina legumen is illustrated in Figure 4. A similar form is Dentalina communis d’Orbigny, but in this latter species the margins are smooth and the chambers are not inflated. This species was originally described from the Tertiary of the Vienna Basin. The Bathonian forms compare well with Neugeboren’s figures, but specimens from the Tertiary were not examined. Specimens of Dentalina cornuformis from the Ter- quem collection in Paris are identical with those recorded here. This species is fairly common in the English Bathonian, particularly in the Upper Fuller’s Earth Clay of the Dorset Coast. 310 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY Material deposited at M.C.Z. Nos. 3344C, 3344D. DENTALINA NODIGERA Terquem and Berthelin Plate 4, figure 8 Dentalina nodigera Terquem and Berthelin, 1875 (pars), p. 25, pl. 1, fig. dla only. Dentalina nodigera, Franke, 1936, p. 26, pl. 2, fig. 7a-b. Specimens are rare, and consist only of fragments. The tests are thin, transparent and composed of pyriform chambers which are inflated on one side and compressed on the other. The sutures are depressed and the last chambers have rather long necks with bulbous accumulations of calcite on the ends. The apertures are somewhat indistinct, but appear to be rounded. A 6 c D E F G H Figure 4. Variation of Dentalina mucronata (Neugeboren) and Vaginu- lina legumen (Linné), showing similarity of end forms in the two species. DENTALINA OOLITHICA Terquem Plate 4, figure 10 Dentalina oolithica Terquem, 1870b, (pars), p. 366, pl. 28, figs. 5-6 only. Dentalina plebeia Terquem 1870b (pars), p. 369, pl. 29, figs. 3-6 only. CIFELLI: BATHONIAN FORAMINIFERA OF ENGLAND Su lal Terquem included a variety of forms in this species. The present specimens agree well with those in the Terquem collec- tion and with the illustrations cited above. The remaining figures are possibly illustrations of variants of other species, but the drawings appear schematic and unlike anything I have encoun- tered in the English Bathonian. This species occurs throughout the English Bathonian, and is particularly common in the Upper Fuller’s Earth Clay. DENTALINA PROPINQUA Terquem Plate 4, figures 4-6 non Dentalina jurensis Gumbel, 1862, p. 220, pl. 3, figs. l4a-e. Dentalina jurensis Terquem, 1870b, p. 362, pl. 27, figs. 6-16. Dentalina propinqua Terquem, 1870b, p. 365, pl. 28, figs. 1-2. Dentalina sinemuriensis, Bartenstein and Brand (non Terquem), 1937, p 1395 ply das fie. Ss spl. 2b, fies 12 ple 46 figs) plsvo, fig. 29s pl. 6: fig. 12; pl. 8, fig. lla-e. Dentalina arbuscula, Bartenstein and Brand (non Terquem), 1937, p. 139, pl ta, fies 7. pls 2by figseal2 Fs plaas, fice eh epee tio ALON pl eaos fig 18: non Dentalina jurensis, Seibold and Seibold, 1955, p. 112, pl. 138, fig. 9; text fig. 2n. This species is intermediate in structure between Dentalina and Nodosaria. Most of the tests are arcuate, but some are straight and the sutures are transverse. There is much variation in the sutures, which are normally depressed, resulting in con- striction of the chambers along the margins. Such forms, how- ever, grade into those having perfectly flush sutures and smooth margins. The size of the aperture is also variable. Where there is ttle constriction of the final chamber it is large, but where the chamber is much constricted the aperture is narrow and typically dentaline. The aperture is only faintly radiate, and in some specimens may be rounded. The name Dentalina jurensis by which Terquem designated this species was preoccupied by D. jurensis of Gumbel, as pointed out by Seibold and Seibold (1955, p. 113). However, D. pro- pinqua appears to be identical with it, so that that name is used here to designate Terquem’s species. This species occurs throughout most of the Bathonian, but is most common in the Upper Fuller’s Earth Clay. 312 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY DENTALINA SUBPLANA Terquem Plate 4, figures 12-13 Dentalina subplana Terquem, 1870b (pars), p. 367, pl. 28, figs. 19, 20, 22 only. Dentalina punguinculus, Bruckman (non Reuss), 1904, p. 28, pl. 3, figs. 29-31, This species is similar to Dentalina terquemi G’Orbigny, and is probably closely related. It differs from the latter in the smaller size of the test, the more oblique sutures, and the lack of bulbous chambers in the last part of the test. D. terquenu appears to be restricted to the Lias in both England and north- west Germany. Forms comparable to D. subplana have not been recorded from northwest Germany. Another similar form is D. vetustissima d’Orbigny as described by Bartenstein and Brand (1937,-p. 137, pl. 4, fie. 25: "pl. 5,, fig: 12;).. "Ehis latter: species is more apiculate, and the chambers are less drum-like than in the present specimens. D. vetustissima was recorded from the Lias only, by Bartenstein and Brand. The forms recorded here compare well with the figures cited, and with the specimens in the Terquem collection. This species is common only in the Wattonensis Beds of the Dorset Coast, but also occurs in the Lower Fuller’s Earth Clay of the Dorset Coast, the Fuller’s Earth Rock, and the Bradford Clay of the Bath area. DENTALINA TORTA Terquem Plate 4, figure 9 Dentalina torta Terquem, 1858, p. 599, pl. 2, fig. 6a-b. Dentalina torta, Franke, 1936, p. 27, pl. 2, fig. 9. Dentalina torta, Bartenstein and Brand, 1937, pl. 3, fig. 9; pl. 4, fig. 24; Dione. 14 ple Os how ie Dentalina varians, Bartenstein and Brand (non Terquem), 1937 (pars), p. 135, pl. 10, fig. 14; pl. 12b, fig. 3. Dentalina ensis, Bartenstein and Brand (non Wisniowski), 1937, pl. 8, fig. 9a-b. This species is similar to Dentalina intorta Terquem with which it may possibly intergrade. However, the tests of the specimens included here are larger and not as compressed as CIFELLI: BATHONIAN FORAMINIFERA OF ENGLAND 313 those of the latter species, but the sutures are more depressed, resulting in more bulbous chambers. The English Bathonian forms differ somewhat from Terquem’s figure, which illustrates a much more tapered test with less bulbous chambers and a proloculus which is much smaller than any observed in my material. The differences are minor, how- ever, and these specimens are considered identical. yenus NODOSARIA Lamarck, 1812 NoposaRIA CLAVULA (Terquem) Plate 4, figure 20 Dentalina clavula Terquem, 1870b, p. 366, pl. 28, fig. 4. These specimens compare well with Terquem’s figure, except that they do not show the depressed sutures, indicated in Ter- quem’s figure. This species is very rare, and was discovered only from the Forest Marble on the Dorset Coast. NODOSARIA HORTENSIS Terquem Plate 4, figures 22-24 Nodosaria hortensis Terquem 1866b, p. 476, pl. 19, fig. 13. Nodosaria fontinensis Terquem 1870b, p. 3538, pl. 26, figs. 1-5. Nodosaria variablis Terquem 1870b, p. 353, pl. 26, figs. 6-12. Nodosaria mutablis Terquem and Berthelin 1875, p. 20, pl. 1, fig. 19a-b. Nodosaria hortensis, Macfadyen, 1941, p. 438, pl. 3, fig. 41a-b. Nodosaria hortensis, Barnard, 1950b, p. 19, pl. 3, fig. 3. An exhaustive synonymy of this species has been provided by Macfadyen (1941, p. 43). In their remarks on Nodosaria fon- finensis (a synonym) Bartenstein and Brand (1937, p. 148) noted that the Dogger specimens tend to have finer, less sharp costae than those from the Lias. A comparison of the present material with that of the Lias of Byfield described by Barnard reveals similar differences. NODOSARIA INGENS (Terquem) Plate 4, figure 25 Dentalina ingens Terquem 1870b, p. 361, pl. 27, fig. 5. Nodosaria guttifera, Bartenstein and Brand (non d’Orbigny), 1937, p. 144, pits; figs 47. 314 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY The material consists of a few fragments composed of two or three bulbous chambers connected by long necks. The apertures are radiate. A sunilar species is Dentalina guttifera d’Orbigny, which is a Tertiary form, originally described from the Vienna Basin. In the present species, however, the chambers are more inflated and tapered less towards the apertural neck. Noposarta aff. N. ISSLERI Franke Plate 4, figures 36-37 Test small, shehtly tapered, composed of four to six chambers increasing in height and breadth as added; sutures depressed, transverse; growth somewhat irregular in early stages, resulting sometimes in slightly arched test; last chamber considerably larger than preceding and flaring; surface ornamented with ten fine ribs extending continuously from base of proloculus to end of last chamber; proloculus small, rounded; aperture ter- minal, rounded, situated on a short neck. The English Bathonian forms are similar to Nodosaria issleri Franke, and differ only in the greater persistence of the ribs which extend up to the end of the apertural neck. Paratypes of Nodosaria issleri were examined at the Senckenberg Museum, Frankfurt a.M., and the present specimens are undoubtedly closely related to this species. Another similar and probably related form is that figured by Issler (1908, p. 52, figs. 73-78) as Nodosaria raphanus (Linné) which lacks the apertural neck, but is otherwise indistinguishable. NODOSARIA LIASSICA Barnard Plate 4, figure 33 Nodosaria obscura, Bartenstein and Brand (non Reuss), 1937, p. 147, pl. 8, figs V8: Nodosaria obscura var. liassica Barnard, 1950b, p. 18, text fig. 10. A few specimens are identical with those described by Barnard from the Upper Lias of Byfield. The Jurassic forms are treated here as a separate species, since Reuss’s figure of NV. obscura is not clear and the relationship with the Cretaceous form is uncertain. CIFELLI: BATHONIAN FORAMINIFERA OF ENGLAND 315 This species is rare, and was observed only in the Lower Fuller’s Earth Clay of the Dorset Coast. NoposARIA OPALINI Bartenstein Plate 4, figures 26-31 Nodosaria opalini Bartenstein and Brand, 1937, p. 147, pl. 8, fig. 13a-b; pl. 10, fig. 18a-b. This species exhibits much variation in the amount of sutural depression and in the strength of the costae. In many specimens the sutures are flush, and the chambers show no constriction. In others, the sutures in the later portion of the test are depressed, and the last chambers are constricted. Some specimens have con- stricted chambers throughout the length of the test. Costae vary from strong, well defined ribs, to faint, barely perceptible stria- tions. They are, however, always present and are numerous. This species is widely distributed throughout the English Bathonian, and is particularly common in the Upper Fuller’s Earth Clay. There are no previous records of it in England. In Germany it has been recorded from the Dogger Alpha (Lower Bajocian) by Bartenstein and Brand. NopOSARIA PECTINATA (Terquem) Plate 4, figures 34-35 Dentalina pectinata Terquem, 1870b, p. 360, pl. 26, figs. 26-30. The material consists of fragments composed of three or four chambers. Although the tests are distinctly arched, the trans- verse sutures and centrally located apertures suggest that the specimens are better placed in Nodosaria than Dentalina. The specimens have been compared with those in the Terquem collection and are identical. NODOSARIA PLICATILIS Wisniowski Plate 4, figure 21 Nodosaria plicatilis Wisniowski, 1890, p. 194, pl. 1, fig. 16. Nodosaria plicatilis, Paalzow, 1922, p. 16, pl. 2, fig. 4. Nodosaria plicatilis, Bartenstein and Brand, 1937, p. 149, pl. 15a, fig. 14. 316 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY This species resembles N. hortensis in its ornamentation and in its general shape. The two species often occur together, but N. plicatilis is distinguished by being more oval in cross-section, with fewer costae. Noposaria aff. N. prima d’Orbigny Plate 4, figure 32 Test straight or slightly arched, tapered; chambers numerous, generally seven or eight composing the test, increasing in size as added, but occasional chamber smaller than previous, longer than broad, only shghtly inflated; proloculus large, spherical ; surface covered with about fourteen prominent costae extending continuously from tip of proloculus to tip of aperture; aperture terminal, radiate, generally centrally located, but sometimes shifted towards margin; sutures transverse or very slightly oblique, depressed. The shift of the aperture from the center to the margin in some specimens, as well as the occasional curvature of the test is reminiscent of Dentalina, and this form is somewhat intermediate in structure. These specimens are similar to the types of NV. prima as re- figured by Macfadyen, but these illustrations do not show the curvature of the test or the shift of the aperture towards the margin. This species resembles NV. hortensis, but has a radiate aperture in addition to more numerous and persistent costae. Noposaria sp. A Plate 4, figure 38 The material consists of a single, tiny specimen which is very slender and is composed of three chambers. The sutures are flush, so that the outline of the test is smooth. The chambers are almost twice as high as broad, with the center one being the longest. There is no visible ornamentation and the aperture is central and radiate. The specimen resembles NV. pyriformis Terquem figured by Macfadyen (1941, p. 46, pl. 3, fig. 44), but the chambers are not as pyriform. It is also similar to N. pyrula d’Orbigny as figured CIFELLI: BATHONIAN FORAMINIFERA OF ENGLAND old by Haeusler (1890, p. 28, pl. 5, figs. 25-26), but is a much slimmer test and has fewer chambers. The single specimen occurs in the Sharp’s Hill Beds, Oxfordshire. NoposaRia sp. B Plate 4, figure 39 The material consists of several fragments composed of four or five chambers arranged in a straight series, with the first chambers broader than long, and the later ones longer than broad. The surface is covered with numerous, fine costae which are barely perceptible. The sutures are transverse, with the earlier chambers impressed on the later ones. The aperture is central and rounded. These specimens are similar to V. pectinata (Terquem), but are much smaller, and have much fainter costae. The impressed sutures have not been observed on N. pectinata. Genus TRISTIX Macfadyen, 1941 TRISTIX SUPRAJURASSICUM (Paalzow) Plate 5, figure 12 Rhabdogonium suprajurassicum Paalzow, 1932, p. 125, pl. 9, fig. 7. Triplasia somaliensis Macfadyen, 1935, p. 11, pl. 1, fig. 7a-b. Trifarina bradyi, Bartenstein and Brand (non Cushman), 1937, p. 185, pl. 15b, fig. la-b. Trifarina oolithica, Bartenstein and Brand (non Terquem), 1937, p. 186, pl. 11b, fig. 26; pl. 15b, fig. 2a-c. Tristia triangularis Barnard, 19538, p. 190, text-fig. A5. The aperture is finely but distinctly radiate. The proloculus is large and spherical and no microspheres have been recognized. In most specimens there are distinct crenulated keels on the three borders which extend into the prolocular area. In one test the borders are rounded and there are no keels, suggesting synonymy with Tristix liasina (Terquem and Berthelin). How- ever, specimens are rare and relationships between the two species cannot be demonstrated from the present material. 318 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY Genus PSEUDONODOSARIA Boomgart, 1949 PSEUDONODOSARIA VULGATA (Borneman) Plate 5, figure 3 Glandulina vulgata Borneman, 1854, p. 31, pl. 2, figs. la-b, 2. Glandulina tenuis Borneman, 1854, p. 31, pl. 2, fig. 2a-b. Pseudoglandulina vulgata, Bartenstein and Brand, 1937, p. 150, pl. 4, fig. Ale pl figs M6. pl 8 ties LOR pledba, tie: 15: Specimens are rare, but appear to fall well within the range of variation of this highly variable species. The chambers are more bulbous and constricted at the sutures than is shown on Borneman’s figures. The specimens most closely resemble the figures of Bartenstein and Brand. PSEUDONODOSARIA HYBRIDA (Terquem and Berthelin) Plate 5, figures 7-11 Glandulina hybrida Terquem and Berthelin, 1875, p. 22, pl. 1, fig. 26. Pseudoglandulina irregularis, Bartenstein and Brand (non Franke), 1937, (pars), p. 151, pli. 8, fig. 21 only. Test long, slender, straight or slightly arcuate, apiculate, rounded or suboval in cross-section ; chambers numerous, varying in number from four to ten, generally greater in height than breadth, but highly irregular in early portion with some cham- bers very low, greatest increase in height after second chamber ; sutures depressed or flush, varying considerably in single speci- mens; proloculus rounded, tiny; aperture terminal, large, rounded or slightly radiate. The variational pattern of this species is similar to that of Pseudonodosaria vulgata (Borneman) as described by Barnard (1950b, p. 24) from the Lias of Byfield, and most of the chambers are highly irregular in size and shape. The two species are similar, but P. hybrida, which is the more common one in the Enelish Bathonian, is distinguished by the more slender test and the blunter, less bullet-like shape of the last chamber. The shape of the last chamber is remarkably constant, even though the earlier ones are highly variable. This species is common in the Cotswold area of the English Bathonian. Material deposited at M.C.Z. No. 3862D. CIFELLI: BATHONIAN FORAMINIFERA OF ENGLAND 319 PSEUDONODOSARIA OVIFORMIS (‘Terquem ) Plate 5, figures 1-2 Glandulina oviformis Terquem, 1864a, p. 168, pl. 7, fig. 4a-b. Nodosaria humilis, Jones and Parker (non Roemer ), 1860, pl. 19, fig. 6. Nodosaria glans, Jones and Parker (non d’Orbigny), 1860, pl. 19, fig. 7. Pseudoglandulina oviformis, Bartenstein and Brand, 1937, p. 149, pl. 4, fig. 40. Pseudoglandulina oviformis, Macfadyen, 1941, p. 47, pl. 3, fig. 47. Pseudoglandulina oviformis, Barnard, 1950b, p. 24, pl. 1, fig. 6. The specimens agree well with Terquem’s figures, except that the outline of the test is perfectly smooth, and there is no con- striction of the chambers at the sutures. In addition, the apertures are distinctly radiate, which is not indicated in the figure. The minute size, smooth surface and slightly fusiform shape distinguish the species. This species is characteristic of the Lower Jurassic, and has been recorded from numerous locali- ties. In the English Bathonian it was observed only in the Lower Fuller’s Earth Clay on the Dorset Coast, where the material consists of three specimens. PSEUDONODOSARIA PUPOIDES (Borneman) Plate 5, figures 4-6 Orthocerina pupoides Borneman, 1854, p. 35, pl. 3, fig. 16a-b. Pseudoglandulina pupoides, Bartenstein and Brand, 1937, p. 150, pl. 5, fig. 30; pl. 8, fig. 20. This species resembles P. oviformis, but is a larger, heavier form and is not as fusiform. Occasional specimens, however, are small and have thin, transparent walls. This species occurs throughout most of the English Bathonian, but is common only in the Upper Fuller’s Earth. Most of the specimens are poorly preserved, and more adequate material may reveal the present specimens to be variants of some other species of Pseudonodosaria. Genus VAGINULINA d’Orbigny, 1826 VAGINULINA CLATHRATA (Terquem ) Plate 5, figures 20-22 Marginulina longuemari var. clathrata Terquem, 1864a, p. 192, pl. 8, figs. 16, 19a-b. 320 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY Frondicularia fenestrata Terquem, 1886, p. 43, pl. 4, fig. 35. Vaginulina proxima, Franke (non Terquem), 1936, p. 85, pl. 8, fig. 34. Vaginulina proxima, Bartenstein and Brand, 1937, p. 164 (pars), pl. 6, fig. 23 only. Vaginulina clathrata, Barnard, 1950b, p. 29, pl. 1, figs. 8-10. Material of this species from the Lias of Byfield has recently been described in great detail by Barnard (1950b). It is highly variable, particularly in the ornamentation and curvature of the test. Barnard illustrated three forms, in one of which the test is absolutely straight and in the other two the early portions are strongly curved. He interpreted the differences as being indica- tive of dimorphism, and designated the straight type as the megalosphere and the other two as the microspheres. However, there are no appreciable size differences among the proloculi of the three forms. If anything, the proloculi of the two curved types are slightly larger than that of the straight one. Further- more, examination of Barnard’s material reveals that all three types are linked by transitional forms. Consequently, there is no clear indication of dimorphism, although the forms figured by Barnard nicely illustrate the more important variational aspects of the species. The specimens from the Bathonian are identical with the forms described by Barnard and show the same variational patterns, although they are considerably less abundant than in the Lias. They are also identical with one of the forms of V. proxima (Ter- quem) deseribed by Bartenstein and Brand, but the variability of those specimens is not known. Payard (1947) dealt with this and similar species described by Terquem, and having access to the Terquem collection erected five new species and two varieties. All of them appear to be individual variants of a single species. This species (sensu stricto) occurs in the basal beds of the Lower Fuller’s Earth Clay, and has not been observed in younger strata. VAGINULINA CLATHRATA EYPENSA 0. subsp. Plate 5, figures 23-26 Vaginulina proxima, Bartenstein and Brand (mon Terquem), 1937, p. 164 (pars), pl. 8, fig. 27a-b; pl. 11), fig. 13; pl. 14b, fig. 6a-b. CIFELLI: BATHONIAN FORAMINIFERA OF ENGLAND az Test large, stout, broad in relation to height, straight or slightly curved; chambers numerous, about five to seven com- posing the test, increasing rapidly in breadth as added, only slightly so in height; sutures straight or shghtly curved, de- pressed ; surface covered by numerous strong costae, extending over greater portion of test; proloculus in megalosphere large, lobular; aperture at peripheral margin, radiate. Only one microsphere was observed, and it differed from the megalospheres only in the smaller size of the proloculus; there is no difference in the curvature of the early portion of the test. There is much modification in the development of the ribs, and although they are always strong, they are not always per- sistent. Some specimens are very broad, and on these the ribs tend to be less well developed away from the central portion of the test. This subspecies differs from the forms of V. clathrata from the Lias, described by Barnard (see above), in being broader and in having a more irregular development of the ribs. There is much overlap between the two forms and they differ only in their patterns of variation; the subspecies lacks the slim, elon- gate types. This subspecies is common in the Fuller’s Earth Rock and the Wattonensis Beds, and was not observed in older rocks; con- sequently it stratigraphically succeeds V. clathrata (sensu stricto). The illustrations of Bartenstein and Brand suggest that V. proxima consists of two comparable forms which also occur in chronologic succession ; the long straight form referable to V. clathrata (sensu stricto) occurs in the Lias Zeta while the broader forms referable to V. clathrata eypensa are first shown to occur in the Dogger Alpha. Material deposited at M.C.Z. No. 3366D. VAGINULINA CONTRACTA (Terquem) Plate 5, igure 17 Marginulina contracta Terquem, 1868, p. 125, pl. 8, figs. 13-24. Marginulina contracta, Terquem, 1886, p. 25, pl. 2, figs. 44-54. Vaginulina contracta, Bartenstein and Brand, 1937, p. 162, pl. 12b, fig. 8; pl. 15a, fig. 23a-b; pl. 15e, fig. lla-b. Soe BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY This species is similar to Vaginulina legumen (Linné), but is flatter, smaller, and generally has fewer chambers. Occasional specimens have last chambers overlapping on the oral margin and are chevron shaped. This species is common throughout the Bathonian. VAGINULINA HARPA Roemer Plate 5, figures 18-19 Vaginulina harpa Roemer, 1841, p. 96, (pars), pl. 15, fig. 12 only. Vaginulina harpa, Haeusler, 1890, p. 108, pl. 15, figs. 22, 28, 27. Vaginulina harpa, Macfadyen, 1935, p. 12, pl. 1, fig. 8a-b. Vaginulina harpa, Bartenstein and Brand, 1937, p. 163, pl. 14b, fig. 7; pl. 14¢, fig. 10; pl. 15a, fig. 24a-b; pl. 15¢, fig. 12a-d. The present specimens agree well with the figures cited, and are identical with the specimens illustrated by Bartenstein and Brand. The species is rare, occurring only in the Lower Fuller’s Earth Clay and the Wattonensis Beds on the Dorset Coast. VAGINULINA ef. V. HECHTI Bartenstein Plate 5, figure 14 A single, broken specimen recovered from the Wattonensis Beds on the Dorset Coast consists of the first three chambers of the test. There are three oblique ribs on the proloculus. The pro- loculus is larger than that described by Bartenstein and Brand (1937, p. 165, pl. 8, fig. 21a-b), but otherwise compares favorably. VAGINULINA LEGUMEN (Linné) Plate 5, figures 15-16 Vaginulina legunen Blake, 1876, p. 464, pl. 19, fig. 11. Vaginulina legumen Haeusler, 1890, p. 107, pl. 14, fig. 49. Vaginulina legumen Paalzow, 1917, p. 237, pl. 45, figs. 10-11. Vaginulina legumen Bartenstein and Brand, 1937, p. 162, pl. 15a, fig. 22. The present specimens exhibit much variation, particularly in the breadth of the test and in the shape of the proloculus. In some variants the test is slim, and there is no curvature in the © 9 CIFELLI: BATHONIAN FORAMINIFERA OF ENGLAND 323 early portion, so that they are structurally identical with Denta- lina. These variants are morphologically indistinguishable from Dentalina mucronata (Neugeboren), and can be separated from it only by their association with transient and more typical forms. The similarity of the ‘‘end forms’’ of the two species is illustrated in Figure 4A-J. There are many records of this species from the Jurassic, although the type is a Recent form from the Mediterranean. The figure of the type in the Ellis and Messina Catalogue of Foraminifera reveals a large, heavy form which is distinetly keeled and has lmbate sutures. Some variant forms in the Jurassic are comparable, but most of them are smaller with depressed sutures and lack keels. The relationships are uncertain and the Jurassic forms should probably be referred to a separate species. Material deposited at M.C.Z. No. 3370B. VAGINULINA MACILENTA (Terquem) Plate 5, figure 13 Marginulina macilenta Terquem, 1869, p. 12, pl. 7, figs. 1-18. Vaginulina macilenta, Bartenstein and Brand, 1937, p. 163, pl. 11b, fig. 12a-b. The present specimens are identical with those in the Barten- stein and Brand collection at the Senckenberg museum, Frank- furt a.M. These authors record this species from the Dogger Alpha through Delta in northwest Germany. In the present material it was observed only in the Lower Fuller’s Earth Clay, where it is very rare. VAGINULINA sp. A. Test small, curved, oval in cross-section, only slightly com- pressed, rounded on periphery; chambers varying in number from four to six, increasing in breadth as added, only shghtly in height, last one very large compared to previous; sutures straight, thin, flush; proloculus round; aperture radiate. These specimens, which occur sporadically throughout the 3athonian, are probably variants or immature forms of some species such as Vaginulina legumen or V. contracta, with which they often occur. Material deposited at M.C.Z. No. 3372. 324 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY Genus CITHARINA d’Orbigny, 1839 CITHARINA COLLIEZI (Terquem) Plate 6, figures 1-2 Marginulina colliezi Terquem, 1866a, p. 430, pl. 17, figs. 10a-e. Marginulina flabelloides Terquem, 1868, p. 102, pl. 6, figs. 1-30. Vaginulina strigillata, Jones and Parker (non Reuss), 1860, p. 457, pl. 20, figs. 30-35. Vaginulina ef. V. flabelloides, Deeke, 1884, p. 29, pl. 1, fig. 16. Vaginulina flabelloides, Franke, 1936, p. 87, pl. 8, figs. 43, 44a-b. Vaginulina flabelloides, Bartenstein and Brand, 1937, p. 164, pl. 6, fig. 27; pl. 8, fig. 30a-c; pl. 19, fig. 27; pl. 12b, fig. 9; pl. 14b, fig. 8 only. Citharina colliezi, Barnard, 1950b, p. 14, pl. 3, fig. 1. This species is represented by few individuals and is a dis- tinctive form in the English Bathonian. However, in other regions where it occurs in abundance it is highly variable and is not easily distinguished from other species, notably Citharina harpula and C. heteropleura. This is particularly true at Fontoy (Moselle), where Terquem originally described these and other comparable species. The specimens in the Terquem collection in- clude many that are transitional between very unlike types. Highly variable are the strength and number of ribs, and the outhne of the test. The ribs range from fine striations typical of the species to heavy costae characteristic of the Vaginulina proxima and V. inconstans group. The shape of the tests varies from fanlike, as in Citharina heteropleura, to slim, narrow types as is found in Vaginulina clathrata. It is not known which if any of these specimens are the types, nor are the exact horizons at which the specimens occur known, so that the relationships of this complex group can not be ascertained until the type locality has been carefully recollected and topotypes have been examined. The present specimens have been compared with those de- seribed by Barnard from the Lias, and are identical. The species is not common in the Bathonian, and perhaps for this reason exhibits little variation. It occurs in the Lower Fuller’s Earth Clay and Wattonensis Beds of the Dorset Coast and the Bath areas, but was not observed at younger horizons. CITHARINA HETEROPLEURA (Terquem) Plate 6, figures 4-5 Marginulina heteropleura Terquem, 1868, p. 116, pl. 7, figs. 19-25. Marginulina pinguis Terquem, 1868, p. 119, pl. 7, figs. 28-29. CIFELLI: BATHONIAN FORAMINIFERA OF ENGLAND 325 Marginulina distensa Terquem, 1868, p. 120, pl. 7, fig. 30. Marginulina flabelloides, Bartenstein and Brand (non Terquem), 1937, p. 164 (pars), pl. 14e, fig. 11; pl. 15a, fig. 25a-b. This species is similar to Citharina colliezi but differs from it in the broader, more fan-shaped test. The sutures are de- pressed, resulting in chambers that are slightly bulbous. The ribbing is very fine, but is persistent and crosses the sutures. There is no morphologic overlap of the two species in the present material but the morphologic affinities appear to be close and perhaps they represent two stratigraphic subspecies. C. hetero- pleura was recovered only from the Upper Fuller’s Earth Clay, while (@. colliezi was not observed above the Fuller’s Earth Rock. In northwest Germany Bartenstein and Brand have included both of these species in Vaginulina flabelloides. From examination of their figures, which are arranged stratigraph- ically, forms comparable to Citharina colliezi occur in the Lias, and the others comparable to C. heteroplewra oceur in the Dog- ger. CITHARINA sp. Plate 6, figure 3 Test large, thick, highly compressed, subtriangular in side view; chambers numbering eight, increasing rapidly in breadth as added, only slightly in height; sutures flush, mostly obscured by thirteen fine costae covering the surface, extending from pro- loculus to end of last chamber; proloculus small, subelliptical ; aperture radiate, on a short neck formed by junction of margins. The material consists of a single specimen recovered from the Wattonensis Beds on the Dorset Coast. The costae and plan of growth are similar to Vaginulina harpa, but the angular outline and thickness are distinctive. Genus LINGULINA d’Orbigny, 1826 LINGULINA DENTALINIFORMIS Terquem Plate 6, figures 11-12 Lingulina dentaliniformis Terquem, 1870b, p. 339, pl. 25, figs. 1-3. Lingulina dentaliniformis, Terquem, 1886, p. 12, figs. 27-29. Lingulina dentaliniformis, Bartenstein and Brand, 1937, p. 152, pl. 10, fig. 20. 326 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY The test is smooth, but occasional specimens reveal faint striations when viewed under high power. This species is very similar to Frondicularia dentaliniformis and can be distinguished from it only by the sutures, which are transverse or slightly arched. The convexity of the sutures is variable, however, and some specimens are difficult to identify. This species is rare in the English Bathonian. LINGULINA DOLIUM (Terquem) Plate 6, figures 9-10 Frondicularia dolium Terquem, 1870b, p. 338, pl. 23, figs. 9-12. non Lingulina dolium Terquem, 1870b, p. 340, pl. 25, figs. 4-5. Test smooth, compressed, initial end conical, remainder rec- tangular; chambers varying in number from four to ten, in- creasing rapidly in height after second or third chamber, then increasing gradually as added, with no increase in breadth; sutures slightly depressed, generally tranverse, but often with sinuous pattern; proloculus minute, globular; aperture elongate, slit-hke. Terquem’s figure shows a projected initial end, with the proloculus and sueceeding chamber combined, so that the minute proloculus is not apparent. This is probably an inaccuracy in the drawing, as the specimen in the Terquem collection clearly has a tiny proloculus, identical with those in the present speci- mens. The sutures of the specimens in the collection are decidedly more sinuous than shown in Terquem’s figure. A similar form is Lingulina ovalis Schwager (non Terquem and Berthelin) described from the Lower Oxfordian of Germany. This species exhibits the same sinuous sutural pattern, but the proloculus is much larger, and the chambers are more constricted at the sutures. LINGULINA LAEVISSIMA (Terquem) Plate 6, figures 6-8 Frondicularia laevissima Terquem, 1866b, p. 481, pl. 19, fig. 19. Lingulina laevissima, Franke, 1936, p. 62, pl. 6, fig. 11. Lingulina taenoides Franke, 1936, p. 62, pl. 6, fig. 12. CIFELLI: BATHONIAN FORAMINIFERA OF ENGLAND 32 The original description of this species by Terquem is as follows (translated from the French): ‘‘Test elongate, com- pressed, straight, smooth, obtuse at both ends, periphery rounded, composed of eight transverse chambers, the first ones plane, quadrangular, the last two a little extended, arched in the middle, aperture oval.’’ In these specimens the last chambers are not inflated, and there is only the slightest trace of arched sutures. In all eases the apertures are indistinet, and it can not be determined if they are radiate or rounded. They agreee best with Lingulina tae- noides Franke, which I believe is a synonym of L. laevissima. There is much divergence of opinion concerning this species, and a variety of forms have been included under this name. Macfadyen (1941, p. 50, pl. 3, fig. 5la-b), for example, has ignored Franke’s L. laevwissima, and has, instead, included Franke’s L. lanceolata as a synonym. This is a very slim form with highly arched sutures. Barnard (1956, p. 272) regards Franke’s L. laevissima and L. lanceolata as one species. In my opinion none of these forms agree as well with the figure of the type as do those figured by Franke. This species occurs commonly in the Bathonian, but individuals are not common. Specimens are generally squashed and poorly preserved. LINGULINA sp. A Plate 6, figure 15 Test elongate, shghtly rounded in eross-seection; chambers numbering eight, first five minute, then increasing rapidly in width, producing spiked appearance to lower portion of test, last chamber elongate, bullet shaped; sutures slightly depressed, indistinet, shg¢htly arched; aperture rounded. The arched sutures resemble Frondicularia, but are not truly “V-shaped. The apiculate initial end, composed of four minute chambers, distinguishes this form from any other encountered, though possibly it may be a variant of one of the smooth Lingu- lina species or even of one of Frondicularia. This form is rare and was observed only in the Cotswolds. 328 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY LINGULINA sp. B Plate 6, figure 14 Test highly perforate, compressed, elongate, oval in cross- section ; chambers numbering three, last one about equal in length to other two combined; aperture central radiate. The highly perforate test and very long third chamber are very distinctive, and the test is unlike that of any other form encountered. The material consists of one specimen, which was recovered from the Wychwood Beds, in the Kirtlington Cement Works, Oxfordshire. LINGULINA sp. C Plate 6, figure 16 Test compressed, smooth, triangular, shghtly twisted; cham- bers numbering six or seven, increasing in size as added, broader than high, but growth somewhat irregular, constricted at the margins; sutures slightly convex, mostly concentric, but with sinuous pattern in last two chambers; proloculus tiny, rounded ; aperture central, indistinct in all specimens, but apparently rounded. This form bears some resemblance to Lingulina dentaliniformis but is much broader, and exhibits a sinuous sutural pattern in the later chambers. The wall is heavier, and the surface appears to be rugose, although this may be due to the state of preservation. Specimens are rare, and were observed only in the Cotswolds. Genus FRONDICULARIA Defrance, 1824 FRONDICULARIA DENTALINIFORMIS Terquem Plate 6, figures 17-19 Frondicularia dentaliniformis Terquem, 1870b, p. 319, pl. 23, figs. 1-8. Frondicularis dentaliniformis, Bartenstein and Brand, 1937, p. 153, pl. 10, fig. 22. All of the specimens are characterized by the constricted chambers, and the markedly depressed, convex sutures. The CIFELLI: BATHONIAN FORAMINIFERA OF ENGLAND 329 aperture is variable, but the radiate form appears to be the exception. The distribution of the species is sporadic, and speci- mens per sample are few, so that the variation could not be observed in single populations, and the description here is based on isolated specimens from various localities and horizons. This species resembles, and undoubtedly is closely related to, Frondicularia nodosaria and Lingulina dentaliniformis. It differs from the former in the lack of clear-cut costae, the greater constriction of the chambers and in the more deeply depressed sutures. It can be distinguished from the latter by the greater convexity of the sutures. It is possible that all three species are transitional, as is suggested by Bartenstein and Brand (1987, p. 109); however, the relationships are not apparent on the basis of the present rather sparsely distributed material. FRONDICULARIA INVOLUTA Terquem Plate 7, figures 1-3 Frondicularia tnvoluta Terquem, 1866a (pars), p. 403, pl. 15, fig. 3a-b only. Frondicularia complanata, Jones and Parker (non Defrance), 1860, p. 457, plat, figs 19° Frondicularia oolithica Terquem, 1870b, p. 315 (pars), pl. 22, figs. 1-6. Frondicularia complanata, Blake, 1876, p. 468, pl. 19, fig. 20. Frondicularia lignaria, Bartenstein and Brand (non Terquem), 1937, p. 152, pls 8; fig: *25a-b; pl. 10s fies 21s pl? Maw ties 7; pl. dilb, figs 8 ‘ple 12a, fienoce pl tab he: Wels dss fies dl pls a homo. Frondicularia involuta, Barnard, 1950b, p. 13, pl. 2, figs. 8-9. In the Bathonian this species exhibits little variation, and the specimens are consistently broad. Barnard noted the same situa- tion in the Lias. The present specimens agree well with those described from the Lias by Barnard, except that the Bathonian ones are more robust and the chambers are slightly higher. FRONDICULARIA INTUMESCENS Borneman Plate 6, figure 13 Frondicularia intumescens Borneman, 1854, p. 36, pl. 3, fig. 19a-e. Frondicularia nitida, Franke (non Terquem), 1936, p. 68, pl. 7, figs. 1-3. Prondicularia sacculus, Franke (non Terquem), 1936, p. 68, pl. 7, fig. 4. Frondicularia intwmescens, Franke, 1936, p. 67, pl. 6, fig. 27. 330 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY Frondicularia nitida, Bartenstein and Brand, 1937, p. 153, pl. 4, fig. 45. Frondicularia intwmescens, Bartenstein and Brand, 1937, p. 155, pl. 2b, figs 19s pl. 4; fies do) plo, fice. Although these specimens differ somewhat from the figure of the type, they compare well with some of the figures cited, and appear to fall within the limits of this highly variable species. The material consists of 3 small specimens, from the Forest Marble, possibly representing immature individuals. Previous records of this species are all from the Lias. FRONDICULARIA LIGNARIA TTerquem Plate 6, figures 20-21 Frondicularia lignaria Terquem, 1866b, p. 480, pl. 19, fig. 14. Frondicularia varians Terquem, 1866b, p. 480, pl. 19, fig. 15a-b. non Frondicularia lignaria, Bartenstein and Brand, 1937, p. 152, many figures. Frondicularia lignaria, Barnard, 1950a, p. 371, text-fig. 3g. This species is distinguished by the tiny proloculus, sharply angular sutures and smooth margin. Barnard’s figure shows a more rounded outline, sutures which are less angular, and ereater overlap of the last chamber which composes about two- thirds of the test. Despite these differences there is good agree- ment, and the Lias form is considered identical. This species is rare, and was observed only in the Lower Fuller’s Earth Clay. FRONDICULARIA NODOSARIA Terquem Plate 7, figures 4-6 Frondicularia nodosaria Terquem, 1870b, p. 319, pl. 22, figs. 25-30. Frondicularia nodosaria, Bartenstein and Brand, 1937, p. 155, pl. 12a, fig. 6; pl. 13, fig. 14; pl. 15a, fig. 21. Lingulina nodosaria, Barnard, 1950b, p. 29, text-fig. 16. The specimens are not as broad as those figured by Terquem, and the sutures are always distinetly arched. This species is similar to Frondicularia oolithica Terquem figured by Barten- stein and Brand (1987, p. 155, many figures), but the latter form consistently lacks costae or any visible striations. CIFELLI: BATHONIAN FORAMINIFERA OF ENGLAND 331 FRONDICULARIA NODOSARIA Terquem var. A n.var. Plate 7, figures 7-5 This variety differs from other forms of this species encoun- tered in the present material in that the sutures are decidedly less convex and are almost transverse. Specimens are rare, and consist mostly of fragments. The observed distribution is con- fined to the Lower Fuller’s Earth Clay. I’RONDICULARIA SPISSA Terquem Plate 6, figures 22-24 Frondicularia spissa Terquem, 1870b, p. 317, pl. 22, fig. 10. Frondicularia spatulata Terquem, 1870b, p. 317, pl. 22, figs. 11-17. Frondicularia spatulata, Franke, 1936, p. 66, pl. 6, fig. 23. Frondicularia spatulata, Bartenstein and Brand, 1937, p. 154, pl. 138, fig. 13a-b; pl. 14¢, fig. 8; pl. 15a, fig. 19a-b; pl. 15e, fig. 9a-b. Frondicularia lingulaeformis, Bartenstein and Brand (non Schwager), 1937, p. 154, pl. 14c¢, fig. 7. This species is similar to Frondicularia nodosaria, but is gen- erally broader than the latter species, and is not as constricted at the sutures. Specimens are identical with those examined in the Terquem collection. Genus SARACENARIA Defrance, 1824 SARACENARIA TRIQUETRA (Gumbel) Cristellaria triquetra Gumbel, 1862, p. 225, pl. 3, fig. 28a-e. Cristellaria (Saracenaria) triquetra Bartenstein and Brand, 1937, p. 170, pl. 14b, fig. 12a-b; pl. 15a, fig. 2la-b; pl. 15e, fig. 16. Saracenella triquetra Barnard, 1953, p. 190, text-fig. B4a-b. The delicate keel is barely visible on some specimens, probably because of preservation. Specimens also show marked differences in the degree of coil, some being almost completely uncoiled as in Saracenella. Material deposited at M.C.Z. No. 3389. Genus LAGENA Walker and Boys LAGENA sTRIATA (d’Orbigny) Oolina striata d’Orbigny, 1839, p. 21, pl. 5, fig. 12. Lagena natrii Blake, 1876, p. 453, pl. 8, fig. 8. Lagena striata, Haeusler, 1890, pl. 8, fig. 12. 332 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY These rare specimens are small and have distinctly radiate apertures. Material deposited at M.C.Z. No. 3390. LAGENA GLOBOSA (Montagu) Material deposited at M.C.Z. No. 3390. Vermiculum globosum Montagu, 1803, in Ellis and Messina, Catalogue of Foraminifera. Lagena globosa, Seibold and Seibold, 1953, p. 67, pl. 4, fig. 17. This species differs from the preceding in being larger, more robust and lacking striations. It is rare in the English Bath- onian. Material deposited at M.C.Z. No. 3391. Family POLYMORPHINIDAE Genus EOGUTTULINA Cushman and Ozawa, 1930 KOGUTTULINA ANGUSTATA (Terquem) Plate 7, figure 12 Polymorphina angustata Terquem, 1864b, p. 296, pl. 12, figs. 33-35. Eoguttulina polygona, Bartenstein and Brand (non Terquem), 1937, p. 179, pl. 13, fig. 19a-e. The test is small, compressed, and composed of four to five chambers arranged biserially for the most part, with a small spiral in the early portion. Cushman and Ozawa (1930, p. 8) state that the chambers in this species are added in planes ninety degrees apart from one another, and the chamber arrangement is tetraserial. This inter- pretation is based on Terquem’s type figures, which, however, show side views only, with no clear indications of the chamber arrangement. This species is similar to Hoguttulina liassica, but is more compressed, has more depressed sutures, and later chambers that are arranged biserially. There is much variation, however, and additional material may show these species to be transi- tional. This species is comparatively rare, although it occurs at a number of localities in various parts of the Bathonian. CIFELLI: BATHONIAN FORAMINIFERA OF ENGLAND 333 KogurruLina LriAssica (Strickland ) Plate 7, figures 9-11 Polymorphina liassica Strickland, 1846, p. 31, text fig. b. Eoguttulina liassica, Cushman and Ozawa, 1930, p. 7, pl. 1, fig. 2a-c. Eoguttulina liassica, Bartenstein and Brand, 1937, p. 178, pl. la, fig. 24a-b; pl. 2a, fig. 23; pl. 2b, fig. 35; pl. 3, fig. 49; pl. 4, fig. 74a-b; pl. 5, fig. 69a-b. Eoguttulina oolithica Bartenstein and Brand, 1937, p. 179, pl. 8, fig. 34a-b; pl. 10, fig. 43a-b; pl. 11a, fig. 18a-c; pl. 11b, fig. 24a-b; pl. 12b, fig. ALSye pl. 15a, fig. 37a-b. Most often the tests consist of three chambers, but occasional specimens have four or five chambers. Each of these is farther removed from the base than the previous one, and the arrange- ment is vaguely reminiscent of a spiral. The test contains an internal tube which is clearly visible in the well preserved speci- mens under transmitted light. This species is common throughout the Bathonian, and is particularly abundant in Northamptonshire and Rutland. Genus PALEOPOLYMORPHINA Cushman and Ozawa, 19380 PALEOPOLYMORPHINA PLEUROSTOMELLOIDES (Franke) Plate 7, figure 13 Polymorphina pleurostomelloides Franke, 1928, p. 121, pl. 11, fig. 11. Paleopolymorphina pleurostomelloides, Cushman and Ozawa, 1930, p. 112, pl. 28, fig. 5a-b. This species was selected by Cushman and Ozawa as the genotype of Paleopolymorphina and their description of the species is as follows: ‘‘Test small, more or less cylindrical, taper- ine towards the initial end; chambers inflated, oval, but little embracing, alternating, arranged in an entirely biserial series, each farther removed from the base; sutures oblique, much de- pressed, distinct, wall smooth; aperture radiate.’’ However, the description of the genus is as follows: ‘‘Test with early chambers spiral, later ones becoming biserial-Genotype-Poly- morphina pleurostomelloides (Franke).’’ It can be seen that the description of the genotype does not correspond to that of the venus, as this species is biserial throughout. 334 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY This species 1s very rare in the Bathonian, but specimens included here compare well with figures of the type from the Cretaceous, except that the chambers of the Bathonian forms are somewhat more elongate, tending to become uniserial in the latest portion, and there are six, instead of seven chambers. Genus GLOBULINA d’Orbigny, 1839 GLOBULINA sp. Plate 7, figure 14 Rare, tiny, globular specimens are best referred to this genus. The sutures are indistinct, so that the growth plan is uncertain. Superficially, however, they closely resemble other forms of Globulina. Family HETEROHELICIDAE Genus PLECTOFRONDICULARIA Liebus, 1903 PLECTOFRONDICULARIA PARADOXA (Berthelin ) Plate 7, figure 16 Frondicularia paradoxa Berthelin, 1879, p. 33, pl. 1, figs. 12-17. Frondicularia paradoza, Issler, 1908, p. 58, pl. 3, figs. 119-121. Flabellina paradoxa, Bartenstein and Brand, 1937, p. 168, pl. 4, fig. 63a-e. Plectofrondicularia paradoxa, Macfadyen, 1941, p. 66, pl. 4, figs. 67-68. The present material consists of two specimens, one of which is a microsphere and the other a megalosphere. The microsphere is the larger of the two, and has minute spines projecting at the margins. Both specimens agree well with those previously described from the Lias, except that in these the biserial portion constitutes a smaller part of the test than appears to be typical of this species. Family BULIMINIDAE Genus BOLIVINA d’Orbigny, 1839 BouIvVINA RHUMBLERI Franke Plate 7, figure 15 Bolivina nitida, Haeusler (non Brady), 1890, p. 79, pl. 12, figs. 38-39. Bolivina rhumbleri Franke, 1936, p. 126, pl. 12, fig. 21. CIFELLI: BATHONIAN FORAMINIFERA OF ENGLAND 335 Bolivina rhumbleri rhumbleri, Bartenstein and Brand, 1937, p. 184, pl. 4, fig. 73a-c; pl. 5, fig. 72a-c. Specimens are very rare. The aperture is slit-like, and the test is smooth with flush sutures. The specimens compare best with Haeusler’s figures, and are slimmer and more fusiform than that illustrated by Franke. Family ROTALIIDAE Genus SPIRILLINA Ehrenberg, 1843 SPIRILLINA INFIMA (Strickland) Orbis infimus Strickland, 1846, p. 30, text fig. A. Spirillina polygyrata, Bartenstein and Brand (non Gumbel), 1937, p. 181, plea, fesd Oe ple 6; fieo7s) plai4e Migs 35h pl. da; tigst2a-csapl. 5c, fig. la-d. Spirillina punctilata, Bartenstein and Brand (non Terquem), 1937, p. 132, pl. 12b, fig. 2a-d; pl. 14¢, fig. 4a-b. Included here are small, planispiral tubular forms which have thin walls and a vitreous luster. When observed under high power pores are visible on some tests. Poorly preserved speci- mens in which the wall has been altered are inseparable from Cornuspira or Ammodiscus. The vitreous appearance of the wall does not offer a clue to the generic position, as Mr. MeNichol of University College, London, has observed that altered specimens of Cornuspira sometimes have a vitreous luster, but can be seen to be imperforate under very high magnifications (personal com- munication, 1955). The problem of separation, therefore, is difficult. A similar form is Spirillina tenuwissima Gumbel. Topotypes of this species, however, are considerably larger and more robust. The forms included here are extremely common throughout the Bathonian. Genus PATELLINA Williamson, 1858 PATELLINA OOLITHICA Terquem Plate 7, figures 21-22 Patellina oolithica Terquem, 18838, p. 382, pl. 45, figs. 3-4. Spirillina oolithica Terquem, 1886, p. 59, pl. 7, figs. 2-4. The present specimens resemble Trocholina nidiformis (Bruck- man), but the last whorl is divided into four chambers, and 336 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY there are no clearly defined pillars on the ventral surface. The Jurassic forms of this genus are clearly more simple than their recent counterparts, particularly the genotype, Patellina corru- gata Williamson. However, the authority of Cushman (1930, p. 14) is followed in retaining the Jurassic forms in this genus. Genus DISCORBIS Lamarck, 1804 DIscOoRBIS DREHERI Bartenstein Plate 7, figures 23-25 Discorbis dreheri Bartenstein in Bartenstein and Brand, 1937, p. 192, pl. 6, fig. 45a-b; pl. 8, fig. 42a-e; pl. 10, fig. 47a-d. A number of specimens collected from various localities com- pare well with the figures cited, but the preservation is rather poor, and the structure is not entirely visible. Most individuals are plano-convex, although some are biconvex, and the ventral margin is slightly indented. Most of the specimens are covered with a layer of calcite which obscures many of the surface features and the aperture. On two specimens, however, a slt-like aperture can be seen which extends along the umbilical margin of the last chamber. The sutures are flush or slightly depressed, and the chambers are slightly bulbous. Ifofker (1953) has referred this species to Rheinholdella on the basis of tooth plate structure. Several sections were made from the present material, but no internal structure was revealed, possibly due to poor preservation. Genus EPISTOMINA Terquem, 1883 EPISTOMINA STELLIGERA (Reuss) Plate 7, figures 17-20 Rotalina stelligera Reuss, 1854, p. 69, pl. 25, fig. 15a-c. Epistomina stelligera, Bruckman, 1904, p. 23, pl. 4, figs. 10-11. Epistomina caracolla, Paalzow (non Roemer), 1932, p. 142, pl. 11, figs. 12-14. Epistomina stelligera, Bartenstein and Brand, 1937, p. 191, pl. lla, fig. 2la-e; pl. 11e, fig. 3a-f; pl. 12a, fig. 27a-b; pl. 12b, fig. 20a-b; pl. 13, fig. 27a-d; pl. 14b, fig. 2la-c; pl. 14e, fig. 18a-b. Many of the specimens are coated with secondary calcite, and much of the structure is not visible. However, on those CIFELLI: BATHONIAN FORAMINIFERA OF ENGLAND 337 tests which are free of this, the peripheral aperture can be seen to be situated close to the ventral margin. No aperture was observed at the base of the apertural face. The keel is very small, delicate, and is distinctly serrate ; however in most specimens the keel is not present, probably because of erosion. A poorly de- veloped reticulate pattern appears on the umbonal region of some specimens. This species resembles Hpistomina mosquensis, but is con- siderably smaller and lacks the heavy seulpturing on the um- bonal region. Kamily ANOMALINIDAE Genus ANOMALINA, d’Orbigny, 1826 ANOMALINA LIASSICcA Issler Plate 7, figure 26 Anomalina liassica Issler, 1908, p. 93, pl. 7, figs. 331-332. Anomalina liassica, Franke, 1936, p. 128, pl. 12, figs. 27a-b. Anomalina liassica, Bartenstein and Brand, 1937, p. 191, pl. 6, fig. 44; pl. 8, fig. 41. The material consists of one specimen which is very small and has six chambers in the last whorl. The test is evolute, and only shghtly trochoid. The aperture is small and slit-hke and is situated at the base of the last chamber, extending shehtly on the dorsal side. The specimen is smaller than those figured by Issler and is less stout, but is otherwise identical. The specimen was recovered from the Upper Fuller’s Earth Clay at the Combe Hay Fuller’s Earth Works, Bath. BIBLIOGRAPHY ADAMS, C. G. 1957. A study of the morphology and variation of some Upper Lias Foraminifera. Micropaleontology, v. 3, pp. 205-226. ARKELL, W. J. 1933. The Jurassic System in Great Britain. Oxford Press, 681 pp. 1947a. The geology of the country around Weymouth, Swanage, Corfe and Lulworth. Mem. Geol. Surv. Great Britain, 386 pp. 1947b. The geology of Oxford. Oxford Press, 267 pp. 338 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 1951. The English Bathonian Ammonites. Part I. Pal. Soc., v. 104, pp. 1-46. 1956. Jurassic geology of the world. Oliver and Boyd (London), 806 pp. 1957. Ammonites from the Fuller’s Earth Rock, Whatley, Somerset. Geol. Mag., v. 94, pp. 322-325. ARKELL, W. J. and D. T. DONOVAN 1952. The Fuller’s Earth of the Cotswolds and its relation to the Great Oolite. Quart. Jour. Geol. Soe. London, v. 107, pp. 227-253. BARNARD, T. 1950a. Foraminifera from the Lower Lias of the Dorset Coast. Quart. Jour. Geol. Soc. London, v. 105, pp. 347-391. 1950b. Foraminifera from the Upper Lias of Byfield, Northampton- shire. Quart. Jour. Geol. Soe. London, v. 106, pp. 1-36. 1952. Foraminifera from the Upper Oxford Clay, Huntingtonshire. Proce. Geol. Assoc., v. 63, pp. 336-350. 1953. Foraminifera from the Upper Oxford Clay (Jurassic) of Redeliff Point, near Weymouth, England. Proc. Geol. Assoc., v. 64, pp. 183-197. 1956. Some Lingulinae from the Lias of England. Micropaleontology, v. 2, pp. 271-282. 1957. Frondicularia from the Lower Lias of England. Micropaleon- tology, v. 3, pp. 171-181. BARTENSTEIN, H. 1948. Taxonomische Abgrenzung der Foraminiferen-Gattungen Pal- mula Lea, Flabellina d’Orbigny und Falsopalmula n.g., gleizeitig eine Revision der Jura-arten von ‘‘Flabellina.’’ Senckenbergi- ana, v. 28, pp. 119-137. and E. BRAND 1937. Mikro-paliiontologische Untersuchungen zur Stratigraphie des nordwest-deutschen Lias und Doggers. Abh. Senckenb. Naturf. Ges., no. 439, pp. 1-224. BERTHELIN, G. 1879. Foraminiféres du Lias Moyen de la Vendée. Rev. Mag. Zool., 3rd ser., v. 7, pp. 24-41. BLAKE, J. F. 1876. Class Rhizopoda. In: Tate and Blake, The Yorkshire Lias. London, pp. 449-473. CIFELLI: BATHONIAN FORAMINIFERA OF ENGLAND 339 BORNEMAN, J. G. 1854. Uber die Liasformation in der Umgegend von G6ttingen und- ihre organischen Einschliisse. Diss. (Berlin), pp. 1-45. Brapy, H. B. 1867. Foraminifera. Jn Moore, C., On the Middle and Upper Lias of southwest England. Proc. Somerset. Archaeol. Nat. Hist. Soce., 1865-66, v. 13, pp. 119-244. BrorzgEn, F. 1953. Problems in the nomenclature of the Foraminifera. Micro- paleontologist, v. 7, pp. 35-38. BRUCKMAN, R. 1904. Die Foraminiferen des litauische-kurischen Jura. Schr. Phys-- Okon. Gesell. Konigsberg, v. 45, pp. 1-36. BucKMAN, 8S. S. 1922. Jurassic chronology. II. Preliminary studies. Certain Jurassic strata near Eypesmouth (Dorset); the Junction Bed of Watton Cliff and associated rocks. Quart. Jour. Geol. Soc. London, v. 78, pp. 378-436. CHaTwIn, C. P. 1948. British Regional Geology. The Hampshire Basin and adjoining areas. 2nd ed., Geol. Surv. and Mus. London, 99 pp. Cox, L. R. 1941. Report of the Easter Field Meeting, 1940, Bath. Proc. Geol. Assoc., v. 52, pp. 16-35. CUSHMAN, J. A. 1918- The Foraminifera of the Atlantic Ocean. Bull. U. S. Nat. 1931. Museum, v. 104, pts. 1-8, 106 pp. 1930. Some notes on the genus Patellina. Contr. Cushman Lab. Foram. Res., v. 6, pp. 11-17. 1937. A monograph of the foraminiferal family Valvulinidae. Cush- man Lab. Foram. Res., Spee. Publ. 8, 210 pp. 1946. A supplement to the monograph of the foraminiferal family Verneuillinidae. Cushman Lab. Foram. Res., Spec. Publ. 7A, 43 pp. 1948. Foraminifera, their classification and economic use. Harvard Press, 4th ed., 605 pp. — and Y. Ozawa 1930. A monograph of the foraminiferal family Polymorphinidae, Recent and fossil. Proc. U. S. Nat. Museum, v. 77, art. 6, 185 pp. 340 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY DEEKE, W. 1884. Die Foraminiferenfauna der Zone des Stephanoceras humphrie- sianum im Unter-Elsass. Abh. Spezialk. Els. Lothring., v. 4, pp. 1-68. 1886. Les Foraminiféres de 1’Oxfordien des environs de Montbeliard (Doubs). Jn Kilian, W., Notes géologiques sur le Jura du Doubs, pt. 4. Mém. Soc. d’Emul. Montbeliard, v. 16, pp. 293-335. 1898. Foraminiferen aus den Dobbertiner Jurensis-Mergeln. Archiy. v. Freund. Naturg. Mecklenburg, Gustrow, pp. 48-57. SDMUNDS, F. H. and K. P. OAKLEY 1947. British Regional Geology. The Central England District, 2nd ed. Geol. Surv. and Mus. London, 80 pp. Euuis, B. F. and A. R. Messina 1940- Catalogue of Foraminifera. Spee. Publ. Amer. Mus. Nat. Hist., 1958. New York. FRANEE, A. 1928. Die Foraminiferen der oberen Kreide Nord- und Mitteldeutsch- lands. Abh. Preus. Geol. Landesanst., n.s., no. Ma spp. 1-207 1936. Die Foraminiferenfaunen des Lias. Abh. Preus. Geol. Lan- desanst., n.s., no. 169, 138 pp. FRENTZEN, K. 1941. Die Foraminiferenfaunen des Lias, Doggers und Malms der Umgegend von Blumberg (Oberes Wutachgebiet). Beitr. Naturk. Forsch. Oberheing. Karlsruhe, pp. 125-402. 1944. Die agglutierenden Foraminiferen der Bermensdorf Schichten (Transversarius-Zone in Schwammfazies) des Gebietes um Blum- berg in Baden. Paliont. Zeitschr., v. 23, pp. 317-342. GALLOWAY, J. J. 1933. A manual of Foraminifera. Bloomington (Indiana), 483 pp. GREEN, G. W. and R. V. MELVILLE 1956. The stratigraphy of the Stowell Park Borehole. Bull. Geol. Surv. Great Britain, no. 11, pp. 1-33. GUMBEL, C. W. 1862. Die Streitberger Schwammlager und ihre Foraminiferen- einschliisse. Jahrheft. vaterl. Naturkd. Wiirttemberg, v. 18, pp. 192-238. HLAEUSLER, J. 1890. Monographie der Foraminiferen Fauna der Schweizerischen Transversarius-Zone. Abh. schweiz. paliiont. Ges., y. 7a pO. 1-134. CIFELLI: BATHONIAN FORAMINIFERA OF ENGLAND 341 THOFKER, J. 1953. The genus Epistomaria Galloway, 1983 and the genus FE pisto- mariodes Uchio, 1952. Paliont. Zeitschr., v. 27, pp. 129-142. ISSLER, A. 1908. Beitriige zur Stratigraphie und Mikrofauna des Lias in Schwa- ben. Palaeontographieca, v. 55, pp. 1-104. JONES, T. R. and W. K. PARKER 1860. On some fossil Foraminifera from Chellaston near Derby. Quart. Jour. Geol. Soc. London, v. 16, pp. 452-458. KELLAWAY, G. A. and F. B. A. WELCH 1948. British Regional Geology. Bristol and Gloucester District. 2nd ed. Geol. Surv. and Mus. London, 99 pp. KELLAWAY, G. A. and V. WILSON 1941. An outline of the geology of Yeovil, Sherborne and Sparkford Vale. Proc. Geol. Assoc., v. 52, pp. 131-174. IXLAHN, H. 1921. Die Foraminiferengeschlechter Rhabdogonium, Frondicularia und Cristellaria der elsassischen und badischen Juraschichten. Freiburg, pp. 1-75. Lorsuicu, A. R. and H. TAPPAN 1952. The foraminiferal genus Triplasia Reuss, 1854. Smithson. Misc. Coll:s ve 17eno..5, Gill pp: MACFADYEN, F. A. 1935. Jurassic Foraminifera. In: The Mesozoie paleontology of British Somaliland. London. Pt. Il. pp. 7-20. 1936. D’Orbigny’s Lias Foraminifera. Jour. Roy. Micros. Soe., ser. 3, v. 56, pp. 147-153. 1941. Foraminifera from the Green Ammonite Beds, Lower Lias, of Dorset. Philos. Trans. Roy. Soe. London, ser. B, Biol. Sci., no. 576, v. 231, pp. 1-73. Mutir-Woop, H. M. 1936. A monograph of the Brachiopoda of the British Great Oolite Series. Pt. I, The Brachiopoda of the Fuller’s Earth. Palaeont. Soc., v. 89, pp. 1-144. D’ORBIGNY, A. 1826. Tableau méthodique de Ja classe des Céphalopodes. Ann. Sci. Nat., ser. 1, v. 7, pp. 96-314. 342 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 1839. Voyage dans 1|’Amérique meridionale. Paris-Strasburg, pp. 1-86. 1846. Die fossilen Foraminiferen des Tertiaeren Beckens von Wien. Paris, pp. 1-312. PAALZOW, R. 1917. Beitriige zur Kenntnis der Foraminiferenfauna der untern Weis- sen Jura in Siiddeutschenland. Abh. Nat. Geol. Niirnberg, v. 19, pp. 203-248. 1922. Die Foraminiferen der Parkinsoni-Mergel von Heidenheim am Hahnemkamm. Abh. Naturk. Ges. Niirnberg, v. 22, pp. 1-35. 1932. Die Foraminiferen aus den Transversarius-Schichten und IJm- pressa-Tonen der Nordostlichen Sehwabischen Alb. Jahresh. vaterl. Naturk. Wiirttemberg, v. 88, pp. 81-142. PAYARD, J. 1947. La faune des Foraminiféres du Lias Superieur du Détroit Poitevin. Univ. Paris, thésis, 254 pp. Reuss, A. HE. 1854. Beitrige zur Charakteristik der Kreideschichten in den Ostalpen im Gosauthale und am Wolfgangsee. Denksehr. K. Akad. Wiss., ve 7, pp. 1-156: 1863a. Die Foraminiferen des norddeutschen Hils und Gault. Sitzber. Math.-nat. Cl, K. Akad. Wiss., v. 46, Abt. 1, pp. 5-100. 1863b. Die Foraminiferen-Familie der Lagenideen. Sitzber. Math. nat. Cl. K. Akad. Wiss., Abt. 1, pp. 308-342. RICHARDSON, L. 1907. The Inferior Oolite and contiguous deposits of the Bath- Doulting District. Quart. Jour. Geol. Soe. London, v. 63, pp. 383-423. 1909. Exeursion to the Frome District, Somerset. Proc. Geol. Assoc., v. 21, pp. 209-228. 1910. On a Fuller’s Earth section at Combe Hay, near Bath. Proce. Geol. Assoc., v. 21, pp. 425-428. 19lla. On the sections of the Forest Marble and Great Oolite on the railway between Cirencester and Chedworth, Gloucestershire. Proc. Geol. Assoc., v. 22, pp. 95-115. 1911b. The Inferior Oolite and contiguous deposits of the Chipping- Norton district, Oxfordshire. Proce. Cottes. Nat. Field Club, v. 17, pp. 195-231. 1929. The country around Moreton in Marsh. Mem. Geol. Surv. England and Wales, 162 pp. 1953. The country around Cirencester. Mem. Geol. Surv. England and Wales, 116 pp. CIFELLI: BATHONIAN FORAMINIFERA OF ENGLAND 343 RICHARDSON, L., W. J. ARKELL, and H. G. DINES 1946. The geology of the country around Whitney. Mem. Geol. Surv. Great Britain, 150 pp. RoEMER, F. A. 1839. Die Versteinerungen des norddeutschen Oolithen-Gebirges. Han- over, 59 pp. 1841. Die Versteinerungen des norddeutschen Kreidegebirges. Han- over, 145 pp. SCHLUMBERGER, C. 1893. Monographie des Miliolidées du Golfe de Marseille. Mém. Soc. Zool, France, v. 6, pp. 199-228. SCHWAGER, C. 1865. Beitrag zur Kenntnis der mikroskopischen Fauna jurassischer Schichten. Jahresh. vaterl. Naturk. Wiirttemberg, v. 21, pp. 82-151. 1867. In: Waagen, W., Uber die Zone des Ammonites sowerbyi. Geogn.-Palion. Beitrige v. Bennecke, v. 1, pp. 654-668. SEIBOLD, E. and I. SEIBOLD 1953. Foraminiferenfauna und Kalkgehalt eines Profils im gebankten Unteren Malm Sehwabens. Abh. Neues Jahrb. Geol. Palaont., v. 98, pp. 28-76. 1955. Revision der Foraminiferen-Bearbeitung C. W. Gumbels (1862) aus den Streitberger Schwamm-Mergeln (Oberfranken, Untern Malm). Abh. Neues Jahrb. Geol. Paliont., v. 101, pp. 91-134. 1956. Revision der Foraminiferen-Bearbeitung C. Sechwagers (1865) aus den Impressaschichten (Unterer Malm) Siiddeutschlands. Abh. Neues Jahrb. Geol. Paliont., v. 103, pp. 91-154. STRICKLAND, H. E. 1846. On two species of microscopie shells found in the Lias. Quart. Jour. Geol. Soe. London, v. 2, pp. 30-31. SYLVESTER-BRADLEY, P. C. 1957. The Forest Marble of Dorset. Proce. Geol. Soe. London, No. 1556, pp. 26-28. SYLVESTER-BRADLEY, P. C. and F. Hopson 1957. The Fuller’s Earth of Whatley, Somerset. Geol. Mag., v. 94, pp. 312-322. TAPPAN, H. 1955. Foraminifera from the Arctic slope of Alaska. Part Il. Jurassic Foraminifera. U. S. Geol. Surv. Prof. Pap., v. 236-B, pp. 21-90. 544 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY TERQUEM, O. 1858. 1862. 1864a. 1864b. 1866a. 1866b. 1868. 1870b. 1883. 1886. Recherches sur les Foraminiféres du Lias du Départements de la Moselle. Mém. Imp. Acad. Metz, v. 39, pp. 563-656. Recherches sur les Foraminiféres de |]’Etage Moyen et de VEtage Inferieur du Lias. Mém. Imp. Acad. Metz, v. 42, pp. 415-466. Troisiéme Mémoire sur les Foraminiféres du Lias des Départ- ments de la Moselle de la Céte d’Or, du Rhone, de la Vienne et du Calvados. Mém. Imp. Acad. Metz, v. 44, pp. 147-228. Quatriéme Mémoire sur les Foraminiféres du Lias, comprenant les Polymorphines des Départements de la Moselle, de la Cote d’Or et de 1’Indre. Mém. Imp. Acad. Metz, v. 44, pp. 229-308. Cinquiéme Mémoire sur les Foraminiféres du Lias des Départe- ments de la Moselle, de la Céte d’Or et de 1’Indre. Mém. Imp. Acad. Metz, pp. 309-454. Sixiéme Mémoire sur les Foraminiféres du Lias des Départe- ments de 1’Indre et de la Moselle. Mém. Imp. Acad. Metz, pp. 455-532. Premier Mémoire sur les Foraminiféres du Systeme Oolithique Etude du Fuller’s-Earth de la Moselle. Bull. Soc. Hist. Nat. Moselle, v. 11, pp. 1-138. . Deuxiéme Mémoire sur les Foraminiféres du Systeme Oolithique zone & Ammonites parkinsoni de la Moselle. Mém. Imp. Acad. Metz, v. 50, pp. 403-456. Troisiéme Mémoire sur les Foraminiféres du Systeme Oolithique comprenant les genres Frondicularia, Flabellina, Nodosaria, ete., de la zone a Ammonites parkinsoni de Fontoy (Moselle). Mém. Imp. Acad. Metz. v. 51, pp. 299-380. Cinquiéme Mémoire sur les Foraminiféres du Systéme Oolithique de la zone Ammonites parkinsoni de Fontoy (Moselle). Paris, pp. 339-406. Les Foraminiféres et les Ostracodes du Fuller’s-Earth des environs de Varsovie. Mém. Soc. Géol. France, ser. 3, v. 4, 112 pp. TERQUEM, O. and G. BERTHELIN 1875. UHLIG, V. 1883. Etude microscopique des marnes du Lias Moyen d’Essey-les- Nancy, zone inférieure de 1’Assise & Ammonites margaritatus. Mém. Soc. Géol. France, ser. 2, v. 10, 126 pp. Uber Foraminiferen aus den tjason’schen Ornatenthon. Jahrb. K.K. geol. Reichsanstalt, v. 33, pp. 735-774. CIFELLI: BATHONIAN FORAMINIFERA OF ENGLAND 345 UsBECK, I. 1952. Zur Kenntnis von Mikrofauna and Stratigraphie im Untern Lias Alpha Sehwabens. Abh. Neues Jahrb. Geol. Palaont., v. 95, pp. 371-476. WISNIOWSKEI, T. 1890. Mikrofauna ilow ornatowyeh okolicy Krakowa. I. Otwornice gornego Kellowayu w Grojeu. Pam. Akad. Umiej. Krakéwie, v. 17, pp. 181-242. Woop, A. and T. BARNARD 1946. Ophthalmidium: a study of nomenclature variation and evolu- tion in the Foraminifera. Quart. Jour. Geol. Soe. London, v. 102, pp. 77-109. Woopwarb, H. B. 1894. The Jurassic rocks of Britain. IV. The lower Ooolitic rocks of England (Yorkshire excepted). Mem. Geol. Surv. United King- dom, 628 pp. y CHECK LIST OF BATHONIAN FORAMINIFERA FROM THE DORSET COAST AREA, DORSETSHIRE LITHOLOGIC UPPER FOREST UNIT FULLER'S EARTH MARBLE [spectes serie] s [23 [4 [s[e {7 [elo [rofielial |37| 39/39 femoris TTT TTT Bao [Ammopacuitien fontinensie | Bolivin |_| c Bouvet: Bed feouvina mummers TTT fevmarine covers LOT [ol | [ole fcimnarine reteropteure TT TT Cithorina heteropleura 1 [cimmerina sp Dentalina mucronata Dentalina propinqua Dentatina ocolithica Dentalina subpiana Discorbia drehert Stet Ho BeGSS0Esa re a rhumbier ‘ q Eoguttulina angustota Eoguttulina trassice Epistomina stellegera T Flabellamina altnoffi et : EE Haplophragmium suprajurrassicum | lo Lageno striate SS j : ot PP Diiiieas tit |_| ae Lenticuling quenstedt: var B a Bea O ® Lenticulina sowerby! oy lo Lenticulina subalata Lenticulina tricarinelia Lenticulina tricarinella vor A Lenticulina turgida f i Lenticulina vorians Lingulina dentaliformis Lingulina faevissima Marginulina daslongschampsi Marginulina terquem! Massiling dorsetonsis Nodosoria clavula Nodosaria hortansis Nodosaria ingens Nodosaria liassica Planularia protracta Planularia sp. A Problematina cf P liassica Pseudonodosaria hybrida (o) ttt Ie Chie) ae HERES Peas ae < ° o > < 3 e ° < z e o z < o = 3 = > ry 3 6 Q . 2 cy CIFELLI: BATHONIAN FORAMINIFERA OF ENGLAND 34 SUPPLEMENTARY LOCALITY DATA Dorset Coast Sample No. shown in Table 1 Field sample No. 1 DoBu4 2 DoBu3 3 DoBud 4 DoBul 5 DoBu2 6 DoEm0 fi DoEm1 8 DoEm2 9 DoEm3 10 DoKm4 11 DoEm5 2 DoEm6 13 DoEm7 14 DoH2 15 DoH1 16 DoH3 Ae/ DoH4 18 DoWl1 19 DoC10 20 DoC9 21 DoC8 22 DoC7 23 DoC6 24 DoC5 25 DoC4 26 DoC2 27 DoCl1 28 DoH7 29 DoH6 30 DoH5 31 DoH8 32 DoWA 33 DoWC 34 DoWB 35 DoWD 36 DoWE 37 DoWG 38 DoWF 39 DoWH 40 DoWI 348 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY Table 2. Foraminifera from the Lower Fuller’s Earth Clay at Midford (Richardson, 1910, p. 427). Sample BMF1 Bolivina rhumbleri Rare Cornuspira liasina Common Eoguttulina liassiea Rare Frondicularia nodosaria var. A Few Haplophragmium suprajurassicum Rare Lenticulina munsteri Few Lenticulina quenstedti Abundant Lenticulina quenstedti var. A Rare Lenticulina tricarinella Rare Lenticulina turgida Rare Nodosaria hortensis Few Nodosaria opalini Rare Planularia beierana Common Pseudonodosaria pupoides Rare Pseudonodosaria vulgata Rare Spirillina infima Few Spirophthalmidium concentricum Common Vaginulina eclathrata Rare Sample BMF2 Dentalina oolithica Rare Eoguttulina liassica Rare Frondicularia nodosaria Rare Lenticulina galeata Few Lenticulina quenstedti Common Lenticulina quenstedti var. A Few Lenticulina tricarinella Common Lenticulina turgida Rare Nodosaria hortensis Rare Nodosaria opalini Rare Patellina oolithica Few Planularia protracta Few Pseudonodosaria vulgata Rare Reophax multilocularis Rare Saracenaria triquetra Rare Spirillina infima Abundant Trochammina haeusleri Rare Vaginulina elathrata Few CIFELLI: BATHONIAN FORAMINIFERA OF ENGLAND 349 Table 3. Foraminifera from the Lower Fuller’s Earth Clay, at the road cut between Maperton and Charleton-Horetorne (Richardson, 1909, p, 213). Sample BaM1 Cornuspira liasina Abundant Dentalina communis Rare Dentalina mucronata Rare Dentalina oolithica Rare Dentalina propinqua Rare Dentalina subplana Few Eoguttulina liassica Few Frondicularia dentaliniformis ? Rare Frondicularia nodosaria Rare Frondicularia spissa Rare Lenticulina munsteri Common Lingulina laevissima Rare Marginulina terquemi Rare Nodosaria opalini Rare Nodosaria aff. N. prima Common Pateilina oolithica Few Planularia beierana Abundant Saracenaria triquetra Few Spirophthalmidium concentricum Common 350 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY Table 4. Foraminifera from the Lower Fuller’s Earth Clay?, at Box. Sample BC012 (about 12 feet below the contact of the Fuller’s Earth Rock ?) Citharina colliezi Rare Dentalina communis Few Dentalina subplana Rare Eoguttulina lassica Few Lenticulina munsteri Common Lenticulina subalata Rare Nodosaria opalini Rare Planularia beierana Abundant Spirillina infima Rare Vaginulina clathrata eypensa Few Vaginulina sp. A Rare Sample BC014 (about 5 feet below contact of Fuller’s Earth Rock?) Dentalina intorta Few Lenticulina munsteri Rare Lenticulina subalata Rare Lenticulina tricarinella Rare Lingulina laevissima Rare Patellina oolithica Rare Planularia beierana Abundant Pseudonodosaria pupoides Rare CIFELLI: BATHONIAN FORAMINIFERA OF ENGLAND 351 Table 5. Foraminifera from the Fuller’s Earth Rock, between Maperton and Charleton-Horetorne (Richardson, 1909, p. 213). 1— (Sample BaM2) In marls of bed f, about 1% ft. below bed e. 2— (Sample BaM3) In clay parting of bed e, about 2 ft. above bed f. 3—(Sample BaM4) In ‘‘Ornithella marl.’’ A, abundant; C, common; F, few; R, rare. Citharina colliezi Dentalina conferta Dentalina intorta Dentalina oolithica Dentalina propinqua Dentalina subplana Eoguttulina lassica Frondicularia dentaliniformis Frondicularia nodosaria Lenticulina munsteri Lenticulina quenstedti Lenticulina quenstedti var. B Lenticulina subalata Lenticulina tricarinella Lenticulina turgida Lingulina laevissima Marginulina terquemi Nodosaria hortensis Nodosaria aff. N. issleri Nodosaria opalini Nodosaria aff. N. prima Patellina oolithica Planularia beierana Planularia protracta Pseudonodosaria pupoides Pseudonodosaria vulgata Spirulina infima Trochammina haeusleri Vaginulina clathrata eypensa Vaginulina contracta Vaginulina sp. BHOnWaa =e) Qn HAA AH oJ t= = oto t= <= 2) ot =o = = oO Dmeawmwran By BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY Table 6. Foraminifera from the interval between the Acuminata Bed and the ‘‘Fuller’s Earth Rock proper,’’ Cross Ways Inn (Cox, 1941, p. 20). Sample BCRI1 Dentalina intorta Rare Dentalina propinqua Rare Flabellamina althoffi Rare Frondicularia nodosaria Rare Haplophragmium suprajurassicum Common Lenticulina munsteri Common Lenticulina quenstedti Few Lenticulina tricarinella Abundant Nodosaria subplana Rare Planularia beierana Abundant Pseudonodosaria pupoides Rare Spirillina infima Few Spirophthalmidium concentrieum Rare Triplasia bartensteini Common Vaginulina clathrata eypensa Common Vaginulina sp. A Rare Table 7. CIFELLI : BATHONIAN FORAMINIFERA OF ENGLAND 353 Foraminifera from the Upper Fuller’s Earth Clay, Combe Hay Fuller’s Earth Works, Bath. (Samples BCH1, BCH3, BCH4) Ammobaculites agglutinans Ammobaculites fontinensis Anomalina liassica Citharina heteropleura Citharina sp. A Cornuspira lasina Dentalina communis Dentalina aff. D. communis Dentalina intorta Dentalina propinqua Koguttulina angustata Koguttulina lassica Frondicularia involuta Frondicularia nodosaria Lenticulina munsteri Lenticulina subalata Lingulina dentaliniformis Lingulina laevissima Marginulina terquemi Nodosaria hortensis Nodosaria aff. N. issleri Nodosaria opalini Patellina oolithica Planularia beierana Pseudonodosaria hybrida Spirillina infima Spirophthalmidium concentricum Trochammina haeusleri Vaginulina contracta Deen Quasi Cc A R 354 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY Table 8. Foraminifera from the Upper Fuller’s Earth Clay, Vernham Wood. 1—(BE'W5) From middle of bed e 2—(BFW4) From clay band in bed d 3—(BFW2) From bottom of bed b 4—(BFW1) From top of bed b i 2 3 4 Dentalina communis R Dentalina intorta C C Dentalina propinqua R Koguttulina angustata R Eoguttulina liassica C C KF Haplophragmum suprajurassicum R F R R Frondicularia involuta R R Frondicularia spissa R Lenticulina munsteri C R Lenticulina subalata F Lenticulina turgida? R Lenticulina varians Cc R A F Lingulina dolium R Nodosaria hortensis R R R Nodosaria opalini R R R Patellina oolithica F Planularia anceps R R Planularia protracta F F R Pseudonodosaria pupoides R Spirillina infima A A A A Tristix suprajurassicum R Triplasia bartensteini F KF Trochammina globigeriniformis R Trochammina haeusleri R C R Vaginulina harpa R Vaginulina legumen F A A F CIFELLI: BATHONIAN FORAMINIFERA OF ENGLAND Co ON Table 9. Foraminifera from the Upper Fuller’s Earth Clay, Henley Hill, Box. On 1—(Sample BC016) about 10 feet below contact of Great Oolite Limestones. 2_(Sample BC015) 2 feet below contact. Cornuspira liasina Dentalina aff. D. communis Dentalina intorta Dentalina oolithica Dentalina propinqua Hoguttulina liassica Frondicularia dentaliniformis Frondicularia involuta Frondicularia nodosaria Lenticulina munsteri Lenticulina subalata Lenticulina tricarinella Lenticulina turgida Lingulina dolium Lingulina laevissima Nodosaria hortensis Nodosaria opalini Planularia beierana Planularia protracta Pseudonodosaria hybrida Pseudonodosaria pupoides Saracenaria triquetra Spirillina infima Spirophthalmidium concentricum Trochammina globigeriniformis Trochammina haeusleri Vaginulina harpa Vaginulina sp. A GLa eye ry PAaQnDReeADDDPeP aA 356 BULLETIN : Table 10. 1—(Sample BBC5) 2— (Sample 3— (Sample 4— (Sample Ammobaculites agglutinans Dentalina communis Dentalina aff. D. communis Dentalina intorta Dentalina oolithica Dentalina propinqua Dentalina subplana EKoguttulina angustata Eoguttulina liassica BBC4) BBC3) BBC1) Frondicularia dentaliniformis Frondicularia nodosaria Frondicularia spissa Lingulina dolium Lingulina laevissima Marginulina terquemi Massilina dorsetensis Nodosaria hortensis Nodosaria opalini Patellina oolithica Planularia beierana Pseudonodosaria hybrida Pseudonodosaria vulgata Spirillina infima Spiroloculina lanceolata Spirophthalmidium concentricum Trochammina globigeriniformis Vaginulina contracta MUSEUM OF N 1 WwW COMPARATIVE ZOOLOGY ft. above fossil bed. ft. above fossil bed. ft. above fossil bed. ft. above fossil bed. i 2 A A R R R FE F F Fr F R R R R R R R R R R R R R C F R R R Ow Dy yy a F Foraminifera from the Bradford Clay, Bradford-on-Avon. Hons ee a on oles) CIFELLI: BATHONIAN FORAMINIFERA OF ENGLAND 300 Table 11. Foraminifera from the Forest Marble, Monk Quarry, Corsham. 1—(Sample BCO1) 38 ft. above Upper Rag Bed. 2—(Sample BCO2) 3 ft. above 1. 3—(Sample BCO3) 1 ft. above 2. 4—(Sample BCO5) 7 ft. above 3. 5—(Sample BCO6) 31% ft. above 4. 6—(Sample BCO7) 3% ft. above 5. 7—(Sample BCO8) 3 ft. above 6. 8—(Sample BCO9) 2 ft. above 7. 1 2 a} 4 5) 6 7 8 Ammobaculites fontinensis R Citharina heteropleura R R Cornuspira liasina iy F Dentalina intorta R t R Dentalina oolithica R Diseorbis dreheri Eoguttulina angustata R F Eoguttulina liassica FE F FE F Frondicularia involuta R R R R Frondicularia nodosaria R Haplophragmium suprajurassicum R Lenticulina munsteri F A A A A Lenticulina varians F R R Nodosaria opalini R R R R Patellina oolithica Planularia anceps ? R Planularia beierana C Pseudonodosaria pupoides F R R R C Reophax multilocularis R R Spirillina infima R R Spiroloculina lanceolata ? R Tristix suprajurassicum R Trochammina globigeriniformis R Trochammina haeusleri R Vaginulina contracta R Vaginulina legumen R C R R A C mm & ry bo ye © 398 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY Table 12. Foraminifera from the Great Oolite Limestones, Cirencester-Chedworth railway cuttings (Richardson, 1911a). 1—(Sample CTCH6) Taynton Stone, bed 36, about 10 ft. from top (Ched- worth). 2—(Sample CTCH7) Taynton Stone, bed 36, about 5 ft. above 1 (Ched- worth). 3— (Sample CTCH3) White Limestone, bed 19a (Stony Furlong). 4— (Sample CTCH2) White Limestone, bed 19 (Stony Furlong). 5—(Sample CTCH11) White Limestone, marl layer, about bed 16 (Ald- grove). 6—(Sample CTCH4) White Limestone, bed 14 or 15 (Stony Furlong). 7—(Sample CTCH8) White Limestone, base of bed 14 (Aldgrove). 8—(Sample CTCH12) White Limestone, bed 14, about 5 ft. above 9 (Ald- grove). 9—(Sample CTCH14) White Limestone, bed 8 (Folly Barn). 10—(Sample CTCH15) White Limestone bed 4 or 5 (Folly Barn). iI 2 3 4 Bs) 6 7 8 9 10 Ammobacultes agglutinans R Re ot Ammobaculites fontinensis R In 1s 18 R Cornuspira lasina 1) 6 13 Dentalina intorta F ER R Dentalina mucronata 1 R R Dentalina propinqua R R Dentalina torta F Discorbis dreheri R Eoguttulina angustata F R Eoguttulina liassica ea aR (6) Be 6G Frondicularia dentaliniformis R Frondicularia involuta R Frondicularia spissa R Haplophragmium suprajurassicum R Tey day AR R Lagena striata R Lenticulina galeata R R R Lenticulina multangulosa RR R R Lenticulina munsteri R F Lenticulina subalata R Lenticulina varians R R R Han CIFELLI: BATHONIAN Lingulina dolium Lingulina sp. A Lingulina sp. C Marginulina terquemi Nodosaria hortensis Nodosaria aff. issleri Nodosaria opa‘ini Patellina oolithica Planularia anceps Planularia beierana Planularia breoni Planularia protracta Planularia sp. A Planularia sp. B Pseudonodosaria hybrida Spirillina infima Spiroloculina lanceolata Triplasia bartensteini Tristix suprajurassicum Trochammina globigeriniformis Vaginulina contracta Vaginulina legumen Vaginulina sp. FORAMINIFERA OF y R F KF R R R F ENGLAND R 18: R R R R R R F CaP Raa R R R R R F R Ke F R R R 309 R 360 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY Table 13. Foraminifera from the Great Oolite Limestones, Hampen railway cuttings (Richardson, 1929, pp. 104-105). 1—(Sample CTH18) Stonesfield Slate Beds, about 22 ft. below top (prob- ably bed 7b). 2 (Sample CTH17) Stonesfield Slate Beds, 4 ft. above 1. 3—(Sample CTH16) Stonesfield Slate Beds, 3 ft. above 2 (probably bed 6). 4—(Sample CTH14) Stonesfield Slate Beds, top of Stonesfield Slate Beds (Rhynchonella Bed). 5—(Sample CTH13) Taynton Stone, about 20 ft. below top. 6—(Sample CTH12) Taynton Stone, 5 ft. above 5. 7—(Sample CTH2) Hampen Marly Beds, about 10 ft. below top. 8—(Sample CTH1) Hampen Marly Beds, 2 ft. above 2. ] 2 3 4 5) 6 7 S Ammobaculites agglutinans R RK R Cornuspira liasina C KF R F Dentalina intorta R R R R Dentalina mucronata R R R EKoguttulina angustata R R Joguttulina liassica C K A (G R F R Flabellamina althoffi R F Frondicularia dentaliniformis R C R Frondicularia involuta 1 F Haplophragmium suprajurassicum R F R 1 R Lenticulina multangulosa R Lenticulina munsteri R R Lenticulina quenstedti iy Lenticulina quenstedti var. B iy Lenticulina subalata iy R Lenticulina turgida R Lenticulina varians R A A Lingulina dohum R F Lingulina laevissima FE Nodosaria opalini R Planularia anceps R Planularia beierana R Planularia protracta R Planularia sp. A R Pseudonodosaria hybrida R F Pseudonodosaria pupoides R R R Spirillina infima F F R R R C R Spiroloculina lanceolata R Triplasia bartensteini R F Trochammina haeusleri R Trochammina sp. R R Vaginulina contracta F Vaginulina legumen R F R F C CIFELLI: BATHONIAN FORAMINIFERA OF ENGLAND 361 Table 14. Foraminifera from Great Oolite Limestones, quarry at Salperton. 1—(Sample CTH9) White Limestone, about 2 ft. above base. 2— (Sample CTH8) Hampen Marly Beds, about 2 ft. below top. 3—(Sample CTH7) Hampen Marly Beds, about 4 ft. below top. 1 2 3 Ammobaculites agglutinans R Ammobaculites fontinensis R R Cornuspira liasina F K Dentalina intorta R Dentalina mucronata F F Dentalina torta C K Discorbis dreheri R Eoguttulina liassica R C Flabelamina althoffii F Frondicularia dentaliniformis R Globulina sp. R Lenticulina multangulosa F Lenticulina munsteri R R Lenticulina quenstedti var. B ? R Lenticulina subalata C Lingulina dolium R Lingulina laevissima R Lingulina sp. A R Lingulina sp. C R Nodosaria opalini R C Nodosaria sp. B R Planularia beierana R Planularia eugenii R Pseudonodosaria hybrida R R Spirillina infima R Triplasia bartensteini R R Trochammina globigeriniformis R Trochammina haeusleri 1g R Vaginulina clathrata eypensa ? R Vaginulina contracta R F F Vaginulina legumen R R 362 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY Table 15. Foraminifera from the Great Oolite, Nirtlington Cement Works, Kirtlington. 1—(Sample OXK4) White Limestone, 15 ft. below top. 2— (Sample OXK3) White Limestone, 1 ft. below top. 3—(Sample OXK9) Kemble Beds, 1 ft. below top. 4—-(Sample OXK8) Wychwood Beds, 4 ft. below top. 5—(Sample OXK7) Wychwood Beds, 4 ft. below top. 1 2 } 4 Cornuspira liasina, R R R iy Dentalina ef. D. bieornis R Dentalina communis R Dentalina intorta R F K Dentalina mucronata F R Dentalina oolithica R © Dentalina propinqua R R Eoguttulina angustata R Eoguttulina liassica iy R Cc Frondicularia nodosaria F Frondicularia spissa F C Haplophragmium suprajurassicum R R Lagena striata Lenticulina munsteri F Lenticulina subalata K Lingulina laevissima R Lingulina sp. B R Nodosaria hortensis C R Nodosaria opalini R EF R Nodosaria pectinata R Nodosaria pleatilis R Patellina oolithica R Pseudonodosaria hybrida R Pscudonodosaria pupoides R Spirillina infima C F R C Spirophthalmidium concentricum C R Vaginulina contracta R (0. Vaginulina legumen C R R Vaginulina sp. A R “~~ F R R =e} R R CIFELLI: BATHONIAN FORAMINIFERA OF ENGLAND 363 Table 16. Foraminifera from Sharp’s Hill Beds, Sharp’s Hill Quarry. 1—(Sample OXS5) Lower Sharp’s Hill Beds, about 5 ft. below top. 2— (Sample OXS2) Top of Lower Sharp’s Hill Beds. 3—(Sample OXS3) Upper Sharp’s Hill Beds, about 4—(Sample OXS4) Upper Sharp’s Hill Beds, 1% ft. from top. Cornuspira Hasina Dentalina communis Dentalina conferta Dentalina ef. D. communis Dentalina intorta Dentalina oolithica EKoguttulina angustata Joguttulina Hassica Epistomina stelligera Frondicularia dentaliniformis Frondicularia involuta Haplophragmium suprajurassicum Lagena striata Lenticulina munsteri Lingulina sp. A Trochammina haeusleri Nodosaria hortensis Nodosaria opalini Nodosaria sp. A -aleopolymorphina pleurostomelloides Patellina oolithica Planularia sp. B Pseudonodosaria hybrida Spirillina infima Spirophthalmidium concentricum Vaginulina contracta Vaginulina legumen 1 P — -~ ww 3 ft. from top. » ER 364 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY Table 17. Foraminifera from the Upper Estuarine Beds, Clipsham New Quarry, Rutland. 1—(Sample RuC6) Upper Estuarine Beds about 16 ft. below top, in ‘* Astarte fimbriata’’ Beds. 2—(Sample RuC3) Upper Estuarine Beds, about 8ft. above 1. 3—(Sample RuC2) Upper Estuarine Beds, about 31% ft. below top. 4—(Sample RuCl) Upper Estuarine Beds, immediately below Great Oolite Limestone. 1 2 3 4 Cornuspira liasina F F Soguttulina angustata R EF Koguttulina lassica F A A A Frondicularia spissa R Haplophragmium suprajurassicum R Lingulina laevissima R Nodosaria opalini F Pseudonodosaria hybrida g Spirillina infima A C Thurammina tunberosa R C CIFELLI: BATHONIAN FORAMINIFERA OF Table 18. Foraminifera from the Upper Estuarine Beds, Twywell Ironstone Pit. 1—(Sample NHT3) Upper Estuarine 2— (Sample NHT2) Upper Estuarine 3—(Sample NHTO) Upper Estuarine 4—(Sample NHT7) Upper Estuarine Ammobaculites fontinensis Dentalina intorta Dentalina oolithica EKoguttulina hassica Globulina sp. Haplophragmium suprajurassicum Lagena striata Lenticulina subalata Lenticulina varians Nodosaria opalini Planularia sp. B Pseudonodosaria hybrida Spirllina infima Spiroloculina lanceolata ? Vaginulina contracta Vaginulina legumen 3eds, about 9 ft. below top. Beds, + ft. below top. Beds, 2 ft. below top. Beds, immediately below top. 1 A R ENGLAND » =o mo FY « ) ) R = “ 365 # R — ~~ ew Index of generic and specifie names aequale, 285 cornuformis, 309, 310 agglutinans, 283, 284 Cornuspira, 288, 335 althoffi, 283, 286 corrugata, 336 Ammobaculites, 283, 284 crepidula, 299 Ammodiscus, 288, 335 Cristellaria, 290-292, 294-299, 302- anceps, 298 304, 331 angustata, 332 cultrata, 292 Anomalina, 337 Darbyella, 295 arbuseula, 311 ee : Dentalina, 280, 281, 306-313, 315, 316 dentaliniformis, 325, 326, 328 deslongchampsi, 299, 302, 305 arguta, 305 arietis, 303 Astacolus, 291, 292, 295, 297, 303, 304 dialatata, 304 bartensteini, 284, 285 Discorbis, 336 beierana, 281, 299-303 distensa, 325 bicornis, 306 dolium, 298, 326 Bolivina, 334, 335 dorsetensis, 266, 281, 286 bradyi, 317 dorsoarecuata, 304 breoni, 308 dreheri, 336 brevis, 308 nsis, 312 bullata, 308 ensis, Koguttulina, 332, 333 : are peer oa OY r 997 canalieulata, 282 Epistomina, 281, 336, 337 eugenii, 280, 281, 303 eypensa, 266, 281, 320, 321 canariense, 290 casis, 294 Centenarina, 286 Citharina, 281, 324, 328 clathrata, 266, 280, 319-321, 324 elavula, 281, 313 colliezi, 281, 324 communis, 306, 307, 310 Falsopalmula, 299, 302 feifeli, 295 fenestrata, 320 tilosa, 299 Flabelammina, 283, 286 : fe. 281. 307 Flabellina, 334 communis, aff., 2 307 2 c ASi< 3 ay flabelloides, 324, 32 av « complanata, 329 ess : ace Se fontinense, 284 compressa, 23/ 7 é a, eRe oe ! s aoe fontinensis, 284, 313 coneentricum, 288, 289 5 : SG ergs franconica, 284 conferta, 30! ; 2 3 é Aare oon : 6 ' Frondicularia, 280, 281, 320, 326, coniea, 2S8$ ASS e a ; Rags 327-331, 334 contracta, 321, 323 contraria, 287 galeata, 280, 290 coprohthiforme, 283 georgiae, 254 cordiformis, 299, 302 glabra, 306 CIFELLI : Glandulina, 318, 319 glans, 319 globigeriniformis, 290 globosa, 332 globosum, 332 Globulina, 334 guttifera, 313, 314 289 Haplophragmium, 283, 284, haeusleri, 280, 285 harpa, 281, 322, harpula, 324 hechti, 322 helios, 294 heteropleura, ene. 325 90-7 281, 324, 325 hortensis, 313, 316 humilis, 319 hybrida, 318 oO inconstans, 304, 52 infima, 335 infimus, 335 inflata, 291, 297 ingens, 313 inquisita, 297 intorta, 281, 308, 309, 312 intumescens, 281, 829, 330 involuta, 329 irregularis, 318 314 ails issleri, issleri, jJurassica, 284, 299 jurensis, 311 laevissima, 326, 327 Solaooe lanceolata, 287, 327 lateralis, 308 legumen, 805, 310, 280, liasina, 288, 317 lassiea, 980. 285, 314, 330 Lagena, 9909 oun 290-298 329 IN) eat 281, OUST IG) 399 Iaw, lignaria, 280, linearis, 308 BATHONIAN FORAMIN VOR, DOO, OG VIFERA OF ENGLAND lingulaeformis, 331 Lingulina, 325-330 lista, 304 macilenta, 280, 32: Marginulina, 299, 305, 306, 324, 325 Massilina, 266, 281, 286 matutina, 292 mauritli, 284, 285 mosquensis, 337 mucronata, 309, 323 multangulosa, 291 282 291, 292, 294 multilocularis, munsteri, 281, mutablis, 3813 natril, 331 nidiformis, 335 nitida, 329, 33 nodigera, 310 nodosaria, 280, 281, 330, 331 280, 281, 311, 313-317 numisalis, 808 Nodosaria, obliqua, 299, 802 obseura, 314 Oolina, 331 oolithiea, 305, 310, opalini, 280, 315 288 317, 329, 330 orbieula, Orbis, 335 Orthocerina, 319 ovalis, 326 oviformis, 319 ovula, 287 Paleopolymorphina, 333 papillata, 282 paradoxa, 334 Patellina, 335, 336 pauperata, 300, 3802, 305 pectinata, 281, 315, 317 pinquis, 32 Planularia, 280, 281, 295, 298-305 . t f ved 1 39, 321), 368 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY Plectofrondicularia, 334 pleurostomelloides, 333 plieatilis, 3815, 316 polonica, 292 polygona, 032 polygyrata, 335 polymorpha, 295, 296 Polymorphina, 332, 333 prima, 316 prima, aff., 281, 316 Problematina, 285 propinqua, 281, 311 protracta, 304 proxima, 3820, 321 pseudocommunis, 306, 307 Pseudoglandulina, 818, 319 Pseudonodosaria, 318, 319 punctilata, 335 punguineulus, 312 pupoides, 319 pyriformis, 316 pyrula, 316 quadricostata, 3803 quenstedti, 280, 291, 292, 293 Quinqueloculina, 287 radiata, 308 raphanus, 314 recte, 297 Reophax, 2 Rhabdogonium, 317 Rheinholdella, 336 rhumbleri, 334, 335 Robulina, 291 Rotalina, 336 rotundata, 287 s0 sacculus, 32¢ Saracenaria, 291, 295, 331 Saracenella, 331 scorpius, 282 semiinvoluta, 299, 302, 304 sinemurensis, 311 somaliensis, 317 sowerbyi, 295 spatulata, 331 Spirillina, 288, 335 Spirolina, 283 Spiroloculina, 287, 288 Spirophthalmidium, 288 spissa, 281, 331 stelligera, 281, 336 striata, 303, 33 strigillata, 324 subalata, 281, 294 subangulata, 290 subinvoluta, 304 suboligostegia, 308 subplana, 281, 312 suprajurassicum, 317 taenoides, 326, 327 tenuis, 318 tenuissimum, 289 terquemi, 306, 312 Tetraplasia, 286 Textularia, 284 Thurammina, 282 Thuramminopsis, 282 torta, 312 triangularis, 289, 317 tricarinella, 280, 281, 295 Trifarina, 317 Triplasia, 284, 285, 286, 317 triquetra, 331 Tristix, 317 Trochammina, 280, 282, 289, 290 Trocholina, 335 tuberosa, 282 turgida, 297, 309 Vaginulina, 266, 280, 281, 305, 310, 319-325 Valvulina, 289 variabilis, 285 vaniablis, 313 varians, 294, 297, 312, 330 ventricosa, 307, 308 Vermiculum, 332 Verneuilina, 284 vetustissima, 312 vulgata, 318 PLATES bo Ol 9-10 25-26 97.98 -i-— 29 PAT Except where specified, all figures are & 60 Ammobaculites agglutinans (d’Orbigny): Bradford Clay, Bath area (BBC5). MCZ No. 3304. Ammobaculites fontinensis (Terquem ). 2—Hampen Marly Beds, Cotswolds (CTH7). MCZ No. 33054, 3, 4—Upper Fuller’s Earth Clay, Dorset, (DoC4). MCZ No. 3305B. 5—Upper Estuarine Beds, Northamptonshire (NTHO). MCZ No. 3305C. Reophax multilocularis Haeusler. 6—Upper Fuller’s Earth Clay, Dorset (DoC10). MCZ No. 3302A. 7—Elongata Beds, Dorset (DoH1). MCZ No. 3302B. ?Problematina cf. P. lTiassica (Jones). Elongata Beds, Dorset (DoH1). MCZ No. 3308. Haplophragmium suprajurassicum Schwager. 9—Wattonensis Beds, Dorset (DoEM1). MCZ No, 3309A. 10-——Cross Ways Inn section, near Bath (BCR1). MCZ No. 3309B. Triplasia bartensteini Loeblich and Tappan; Cross Ways Inn see- tion, near Bath (BCR1). MCZ No. 3310. Spirophthalmidium concentricum (Terquem and Berthelin); Upper Fuller’s Earth Clay, Dorset (DoC5). MCZ No. 3314. Massilina dorsetensis n. sp.; Forest Marble, Dorset (DoWA). 15, 16—Paratypes. MCZ No. 33114. 17—Holotype. MCZ No. 3311B. Spiroloculina lanceolata (Terquem and Berthelin); Upper Fuller’s Earth Clay, Dorset (DoW1). MCZ No. 3312. Trochammina sp.; Hampen Marly Beds, Cotswolds (CTH1). MCZ INO} ollie Trochammina haeusleri (Galloway); Upper Fuller’s Earth Clay, Box (BCol6); * 100. MCZ No. 3315. Trochammina globigeriniformis (Parker and Jones). °23—Wattonensis Beds, Dorset (DoEm2). MCZ No. 3316A. 24—Upper Fuller’s Earth Clay, Bath area (BC016). MCZ No. 3316B. Flabellamina althoffi Bartenstein. 25—Cross Ways Inn section, Bath area (BCR1). MCZ No. 33034. 26—Upper Fuller’s Earth Clay, Dorset (DoC4). MCZ No. 3303B. Cornuspira liasina Terquem; Forest Marble, Dorset (DoWF). MCZ INO cola: Thurammina tuberosa Haeusler; Upper Estuarine Beds, Rutland, (RuC3). MCZ No. 3301. PLATE | 5-17 23 PLATE 2 Exeept where specified, all figures are & 60 Lenticulina subalata (Reuss); Upper Fuller’s Earth Clay, Dorset (DoC5). MCZ No. 3324. Lenticulina munsteri (Roemer); Upper Fuller’s Earth Clay, Dorset (DoC5). MCZ No. 3320. Lenticulina quenstedti (Gumbel); JWattonensis Beds, Dorset (DoEm1). MCZ No. 3321. Lenticulina quenstedti var. B n.var.; Wattonensis Beds, Dorset (DoEm1). MCZ No. 3323. Lenticulina quenstedti var. A n.var. 9—Lower Fuller’s Earth Clay, Dorset (DoBu4). MCZ No. 3322A. 10—Lower Fuller’s Earth Clay, Bath area (BMF1). MCZ No. 3322B. Lenticulina varians (Borneman). 11—Hampen Marly Beds, Cotswolds (CTH2). MCZ No. 3328A. 12, 13—Forest Marble, Dorset (DoWA). MCZ No. 3328B. Lenticulina multangulosa (Schwager); Hampen Marly Beds, Cots- wolds (CTH7). MCZ No. 3319. Lenticulina galeata (Terquem); Lower Fuller’s Earth Clay, Dorset (DoBud). MCZ No. 3318. Lenticulina turgida (Schwager); Wattonensis 3eds, Dorset (DoEm4). MCZ No. 3329. Lenticulina tricarinella (Reuss); Fuller’s Earth Rock, Bath area (BaM4). MCZ No. 3326. Lenticulina tricarinella var. A n.var.; Upper Fuller’s Earth Clay, Dorset (DoC5). MCZ No. 3327. PLATE 2 1-10 16-17 Rahs Except where specified, all figures are & 60 Planularia beierana (Gumbel). 1, 4, 6, 7, 10—IWattonensis Beds, Dorset (DoEm4). MCZ No, 3330A. 2—Upper Fuller’s Earth Clay, Dorset (DoC2). MCZ No. 3330B. 3—Wattonensis Beds, Dorset (DoEm1). MCZ No. 3330C. o—Wattonensis Beds, Dorset (DoEm0). MCZ No. 3330D, 8, 9—Lower Fuller’s Earth Clay?, Bath area (BCO14). MCZ No. 33308. Planularia breoni (Terquem); Hampen Marly Beds, Cotswolds (CTE7). MCZ No: 333i. Planularia protracta (Borneman). 12—Upper Fuller’s Earth Clay, Bath area (BFW4). MCZ No. 3334B. 13—Wattonensis Beds, Dorset (DoEm3). MCZ No. 3334A. Planularia anceps (Terquem); Wattonensis Beds, Dorset (DoEm4). MCZ No. 3329. 14— Megalosphere. 15—Microsphere. Planularia eugenii (Terquem). 16—Wattonensis Beds, Dorset (DoEm4). MCZ No. 3332A. 17—Lower Fuller’s Earth Clay, Dorset (DoBu2). MCZ No. 3332B. Planularia sp. B; White Limestone, Cotswolds (CTCH12). MCZ No. 3336. Marginulina terquemi d’Orbigny; Upper Fuller’s Earth Clay, Dorset (DoC5). MCZ No. 3338. Marginulina deslongchampsi Terquem; Lower Fuller’s Earth Clay, Dorset (DoBu4). MCZ No. 3337. PLATE 3 oO Ol PLATE 4 Except where specified, all figures are & 60 Dentalina mucronata Neugeboren. 1—Forest Marble, Dorset (DoWD). MCZ No. 33444. 2—Upper Fuller’s Earth Clay, Dorset (DoC5). MCZ No. 3344B. Dentalina conferta Schwager; Wattonensis Beds, Dorset (DoEm1) ; < 100. MCZ No. 3342. Dentalina propinqua Terquem; Upper Fuller’s Earth Clay, Dorset (DoW1); Figure 6& 100. MCZ Nos. 3347A, 3347B, 3347C, 3347D. Dentalina bicornis Terquem; Wychwood Beds, Oxfordshire (OXK9). MCZ No. 3339. Dentalina nodigera Terquem and Berthelin; JVattonensis Beds, Dor- set (DoEm6). MCZ No. 3345. Dentalina torta Terquem; Hampen Marly Beds, Cotswolds (CTH7). MCZ No. 3349. Dentalina oolithica Terquem; Upper Fuller’s Earth Clay, Dorset (DoC1); & 100. MCZ No. 3346. Dentalina communis d’Orbigny; Wychwood Beds, Oxfordshire (OXK9); X 25. MCZ No. 3340A. Dentalina subplana Terquem; Wattonensis Beds, Dorset (DoEm4). MCZ Nos. 3348A, 3348B, 3348C. Dentalina aff. D. communis d’Orbigny; Forest Marble, Dorset (DoWG). MCZ Nos. 3341A, 3341B. Dentalina intorta Terquem. 16, 17—Forest Marble, Dorset (DoWB). MCZ 3343A, 33435. 18—Upper Fuller’s Earth Clay, Dorset (DoC1). MCZ No. 3348C. 19—Forest Marble, Dorset (DoWB); & 100. MCZ No. 33438D. Nodosaria clavula Terquem; Forest Marble, Dorset (DoWB). MCZ No. 3350. Nodosaria plicatilis Wisniowski; Wattonensis Beds, Dorset (DoEm0). MCZ No. 3356. Nodosaria hortensis Terquem. 22—Upper Fuller’s Earth Clay, Dorset (DoC5). MCZ No. 33514. 23—Lower Fuller’s Earth Clay, Dorset (DoBu2). MCZ No. 3351B. 24—Lower Fuller’s Earth Clay, Dorset (DoBu3). MCZ No. 3351C. Nodosaria ingens (Terquem); Wattonensis Beds, Dorset (DoEm6) ; x 100. MCZ No. 33524. Nodosaria opalini Bartenstein; Upper Fuller’s Earth Clay, Dorset (DoC2). MCZ No. 3354. Nodosaria aff. N. prima d’Orbigny; Lower Fuller’s Earth Clay, Bath area (BaM1). MCZ No. 3357. Nodosaria liassica (Barnard); Lower Fuller’s Earth Clay, Dorset (DoBud).- MCZ No. 3353. Nodosaria pectinata (Terquem). 34—Upper Fuller’s Earth Clay, Dorset (DoC8). MCZ No. 3355A. 35—Upper Fuller’s Earth Clay, Dorset (DoC5). MCZ No. 3355B. Nodosaria aff. N. issleri Franke; Upper Fuller’s Earth Clay, Bath area (BCH3). MCZ No. 3352B. Nodosaria sp. A; Sharp’s Hill Beds, Oxfordshire (OXS5). MCZ No. 3358. Nodosaria sp. B; Hampen Marly Beds, Cotswolds (CTH7). MCZ No. 3359. | PLATE 20-22 23-26 PLATE 5 Except where specified, all figures are & 60 Pseudonodosaria oviformis (Terquem); Lower Fuller’s Earth Clay, Dorset (DoBul). MCZ No. 3363. Pseudonodosaria vulgata (Borneman); Upper Fuller’s Earth Clay, Dorset (DoC2). MCZ No. 3361. Pseudonodosaria pupoides (Borneman). 4, 5—Forest Marble, Bath area (BCO9). MCZ 3364A. 6—Forest Marble, Bath area (BCO1). MCZ No. 33645. Pseudonodosaria hybrida (Terquem and Berthelin). 7—Upper Fuller’s Earth Clay, Dorset (DoW1). MCZ No, 33624. 8—Upper Fuller’s Earth Clay, Dorset (DoC4). MCZ No. 3362B. 9, 11—Hampen Marly Beds, Cotswolds (CTH7). MCZ No, 3362C. Tristix suprajurassicum (Paalzow); Wattonensis Beds, Dorset (DoEm1). MCZ No. 3360. Vaginulina macilenta (Terquem); Lower Fuller’s Earth Clay, Dorset (DoBud). MCZ No. 3871. Vaginulina ef. V. hechti Bartenstein; MWattonensis Beds, Dorset (DoEm4). MCZ No. 3369. Vaginulina legumen (Linné); Forest Marble, Bath area (BCO8). MCZ No. 3370A. Vaginulina contracta (Terquem) ; Forest Marble, Bath area (BCOS). MCZ No. 3367. Vaginulina harpa Roemer. 18—Wattonensis Beds, Dorset (DoEm0). MCZ No. 3368A. 19—Upper Fuller’s Earth Clay, Bath area (BCH1). MCZ No, 3368B. Vaginulina clathrata (Terquem) ; Lower Fuller’s Earth Clay, Dor- set (DoBu5). MCZ No. 3365A. Vaginulina clathrata eypensa n. subsp. 93, 24, Paratypes Lower Fuller’s Earth Clay?, Bath area (BCO012). MCZ No. 3566A. 25, Paratype Wattonensis Beds, Dorset (DoEm1). MCZ No. 3366B. 26, Holotype Cross Ways Inn section, Bath area (BCR1). MCZ No. 3366C. PLATE 5 1-2 eo va on 9-10 20-21 9994 PLATE 6 Except where specified, all figures are & 60 Citharina colliezi (Terquem). 1—-Wattonensis Beds, Dorset (DoEm0); & 35. MCZ No. 3373A. 2—Wattonensis Beds, Dorset (DoEm1). MCZ No. 3373B. Citharina sp.; Wattonensis Beds, Dorset (DoEm0); & 35. MCZ No. 3375C. Citharina heteropleura (‘Terquem). 4—Upper Fuller’s Earth Clay, Dorset (DoCS); & 100. MCZ No. 3374A. 5—Upper Fuller’s Earth Clay, Dorset (DoC2). MCZ No. 3374B. Lingulina laevissima (Terquem). 6—Upper Fuller’s Earth Clay, Dorset (DoW1). MCZ No. 3378A. 7—Hampen Marly Beds, Cotswolds (CTH7); X& 100. MCZ No. 3378B. 8—Upper Fuller’s Earth Clay, Dorset (DoW1); *& 100. MCZ No. 3378C. Lingulina dolium (Terquem); Upper Fuller’s Earth Clay, Bath area (BC015); Figure 9 & 100. MCZ No. 3377. Lingulina dentaliniformis Terquem; Upper Fuller’s Earth Clay, Dorset, (DoCS). MCZ No. 3376. Frondicularia intumescens Borneman; Forest Marble, Dorset (DoWB); X 100. MCZ No. 3384. Lingulina sp. B; Wychwood Beds, Oxfordshire (OXK9). MCZ No. 3380. Lingulina sp. A; Hampen Marly Beds, Cotswolds (CTH7); & 100. MCZ No. 3379. Lingulina sp. C; Hampen Marly Beds, Cotswolds (CTH7); & 100. MCZ No. 3381. Frondicularia dentaliniformis Terquem. 17, 19—Hampen Marly Beds, Cotswolds (CTH7). MCZ No. 33824. 18—Upper Fuller’s Earth Clay, Dorset (DoCS). MCZ No. 3382B. Frondicularia lignaria Terquem; Lower Fuller’s Earth Clay, Dorset (DoBu4). MCZ No. 3385, Frondicularia spissa Terquem. 22, 24—-Forest Marble, Dorset (DoWG); * 100, MCZ No. 33884. 23—Kemble Beds, Oxfordshire (OXK1). MCZ No. 3388B. PLATE 6 1- 9 26 3 -11 -20 PLATE 7 Except where specified, all figures are « 60 Frondicularia involuta Terquem. 1—Forest Marble, Bath area (BCO8). MCZ No. 33834. 2—Forest Marble, Bath area (BCO9). MCZ No. 3383B. 3—Hampen Marly Beds, Cotswolds (CTH2). MCZ No. 33883C. Frondicularia nodosaria Terquem. 4, 6—Upper Fuller’s Earth Clay, Dorset (DoC7). MCZ No. 3386A, 5—Wattonensis Beds, Dorset (DoEm3). MCZ No. 3386B. Frondicularia nodosaria Terquem var. A n.var.; Lower Fuller’s Earth Clay, Dorset (DoBu4). MCZ No. 3387. Eoguttulina liassica (Strickland). 9, 11—Upper Fuller’s Earth Clay, Dorset (DoC5); x 100. MCZ No. 33934. 10—Hampen Marly Beds (CTH7); X 100. MCZ No. 3393B. Eoguttulina angustata (Terquem); Upper Estuarine Beds, Rutland (RuC3); & 100. MCZ No. 3392. Paleopolymorphina pleurostomelloides (Franke); Sharp’s Hill Beds, Oxfordshire, (OXS2); & 100. MCZ No. 3394. Globulina sp.; Hampen Marly Beds, Cotswolds (CTH7); & 100. MCZ No. 3395. Bolivina rhumbleri Franke; Wattonensis Beds, Dorset (DoEm4) ; x 100: MCZ No. 3397. Plectofrondicularia paradoxa (Berthelin); Lower Fuller’s Earth Clay, Bath area (BMF1); x 100. MCZ No. 3396. Epistomina stelligera (Reuss); Upper Fuller’s Earth Clay, Dorset (DoC6); & 100. MCZ No. 3400. Patellina oolithica Terquem; Forest Marble, Dorset (DoWE). MCZ No. 3398B. 21—-Ventral view; * 100. 22—Dorsal view; & 100. Discorbis dreheri Bartenstein. 23—White Limestone, Cotswolds (CTCH4); dorsal view; & 100. MCZ No. 33994. 24—White Limestone, Cotswolds (CTCH4); ventral view; & 100. MCZ No. 3399A. 25—Wattonensis Beds, Dorset (DoEm7); dorsal view; * 100. MCZ No. 3399B. Anomalina liassica Issler; Upper Fuller’s Earth Clay, Bath area (BCH4); X 100. MCZ No. 3401. PLATE 7 , Mie aha Bulletin of the Museum of Comparative Zoology JNM ASEAN ARH WEIN A ID) (OPO) IEy 1) (Ea Vor. 121, No. 8 SILICIFIED MIDDLE ORDOVICIAN TRILOBITES: Remopleurididae, Trinucleidae, Raphiophoridae, Endymioniidae By H. B. Wuirrineton WITH 36 PLATES CAMBRIibuEH, MASS., U.S.A. PRINTED FOR THE MUSEUM NoveEMBER, 1959 PUBLICATIONS ISSUED BY OR IN CONNECTION WITH THE MUSEUM OF COMPARATIVE ZOOLOGY AT HARVARD COLLEGE BULLETIN (octavo) 1863 — The current volume is Vol. 121. BREVIORA (octavo) 1952 — No. 115 is current. Memorrs (quarto) 1864-1938 — Publication was terminated with Mole aa: JOHNSONIA (quarto) 1941— A publication of the Department of Mollusks. Vol. 3, no.39 is current. OccASIONAL PAPERS OF THE DEPARTMENT oF MoLLusks (octavo) 1945 — Vol. 2, no. 23 is current. PROCEEDINGS OF THE New ENGLAND ZooLogicaAL CLuB (octavo) 1899-1948 — Published in connection with the Museum. Publication terminated with Vol. 24. The continuing publications are issued at irregular intervals in num- bers which may be purchased separately. Prices and lists may be obtained on application to the Director of the Museum of Comparative Zoology, Cambridge 38, Massachusetts. Of the Peters ‘‘Check List of Birds of the World,’’ volumes 1-3 are out of print; volumes 4 and 6 may be obtained from the Harvard Uni- versity Press; volumes 5 and 7 are sold by the Museum, and future volumes will be published under Museum auspices. Bulletin of the Museum of Comparative Zoology ANAM Dat IN 1s WIN IRD) COO) IG AAEM a Vox. 121, No: 8 SILICIFIED MIDDLE ORDOVICIAN TRILOBITES: Remopleurididae, Trinucleidae, Raphiophoridae, Endymioniidae By H. B. WuirtineTon WiTH 36 PLATES CAMBRIDGE, MASS., U.S.A. PRINTED FOR THE MUSEU ™M NOVEMBER, 1959 Trinucleidae, Raphiophoridae, Endynuionidae By H. B. WHITTINGTON CONTENTS Introduction and acknowledgements .............-..0.0.02808.00205 MerMinOlO@y: cis cysts s cies, Sasts ogtieesopoked nae cy NEA PERO B ahr, Bor Sct ewees cs Stratieraphicall occurrence and! localitiesia4es-). 2444 nec ier oes eee nists. of trilobite: faunas). .a0..) sane MP rs ees oa A aun alleatimitl esis see ae sere se ptpsyl RES eas WY. Systematie descriptions and discussions ...... Family Remopleurididae Hawle and Corda, 1847 Be: Remopleurides Portlock, 1843 .... Cee) Seach ty eee Ane Morphology of the holaspid .. Wythe a ee Discussion of development ......... es Eee Ue Tea Jc . caelatus n. Sp. ... 0... Seater IR oes ee Ne . plaesiourus n. sp. ee er Ce Aer REE eae ne . caphyroides n. sp. ...... Te eS Teen SHASDERULUS NeeS Wey, seers actuate cae hy eure aaa oe ROU ee eeu CLUNUUS NASD A fie .6= PING) fies! 2.03) chet). Such an area is present at a similar developmental stage of Menoparia (Ross, 1951, pl. 81, fig. 14), and is reduced in large eranidia by expansion of the glabella (Ross, 1953, pl. 62, fig. 4), but never disappears. In Remopleurides, on the other hand, the glabella rapidly expands between the eye lobes and the crescentic area narrows and then disappears, so that the axial and palpebral furrows approach and merge into a single furrow (PI. 4, figs. 1, 7, las PlsGitios: 2) dean Sill? Plas eties Mla 20 24-428), Other changes that take place in this size series are the general 400 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY increase in convexity of glabella, both longitudinally and trans- versely, and the reduction in width (tr.) of the anterior area of the fixed cheek. Transitory pygidia. In Remopleurides caelatus the next trans- itory pygidia known after those of degree 0 are believed to belong to degrees 5, 6, and 7, and have a long median axial spine (Pl. 4, figs. 5, 6, 138, 16-18, 22). In the development of Menoparia, Ross (1953, pl. 62, fig. 20) found a similar type of pygidium to be next in the series. In R. caelatus no intermediates are known between the degree 0 and degree 5 transitory pygidia. [t is possible that the absence of such specimens implies that they may not have existed; that is, that there was a simultaneous release of four segments into the thorax so that the degree 0 transitory pygidium was succeeded by that of degree 5. In the material at hand the median axial spine appears suddenly and is of extraordinary length. It must develop, perhaps abruptly between moults, from the region represented by the strongly convex terminal axial portion of the degree 0 transitory pygi- dium (Pl. 3, figs. 10-13, 15). Transitory pygidia of degrees 8 to 10 (PI. 4, figs. 23-25, belonging to the developmental stages after the release of the segment bearing the median axial spine into the thorax), progressively approach the appearance of the true pygidium and do not have the strongly convex terminal axial portion of the degree 0 transitory pygidium. Discussion. The development of Remopleurides is best known trom R. caclatus, the less complete developmental series of other species showing, however, that they are similar. The remarkable parallels that have been drawn between the development of the Lower Ordovician species of Menoparia and Middle Ordovician species of Remopleurides, serve to illustrate the principle that related animals display similar early developmental stages. A comparison between the illustrations given here, and those of Ross, shows that the protaspides and to a lesser extent meraspid degree 0 have a family resemblance to each other. Subfamily and generic characters are visible at immediately succeeding degrees — for example, the shape of the glabella, width (tr.) of the anterior area of the fixed cheek, length (sag. and exs.) of the preglabellar area, form of the pleural regions in Meno- paria. When comparison is made between the developmental WHITTINGTON : ORDOVICIAN TRILOBITES 401 series of the new species of Remopleurides, it is evident that specifie characters can be picked out even in the protaspis. The developmental series of R. caelatus and R. plaesiourus, and of R. asperulus and R. eximius are similar to each other, as are the holaspides. Both developmental series and holaspis of R. caphy- roides are distinetly different. REMOPLEURIDES CAELATUS D.sp. Plates 1-3; Plate 4, figures 1-25; Text-figures 4, 5 Holotype. USNM 137675 (PI. 1, figs. 1, 2), locality 2. Other Material. Paratypes USNM 137676a-e; all figured specimens in U.S. National Museum. Geological Horizon and Localities. Edinburg formation, localities 2-4, 6, 7, Iya Description. Cephalon of outline, excluding librigenal spines, sub-oval, maximum width greater than leneth (sag.). Glabella of length shghtly greater than maximum width, which is across a point about three-fifths the length. Occipital ring widest (sag. ) at mid-line, decreasing progressively in width outward, small median tubercle adjacent to occipital furrow. Latter deep medially, rising vertically on anterior side to median glabellar area; distally where it runs along margin of palpebral rim, the occipital furrow is shallow. Median glabellar area gently con- vex, descending abruptly (especially postero-laterally ) to palpe- bral furrow. Three pairs of lateral glabellar furrows are visible because the exoskeleton is thinner, being excavated on the inner side (Pl. 1, figs. 5, 6). First and second furrows similar in leneth, curvature of first stronger, directed inward and_ back- ward ; distal part of first furrow situated in line with maximum width of median glabellar area, furrow becomes wider inward; second furrow distally of similar width to first, but tapering inward; third furrow short, sub-oval in outline, situated in lne (exs.) with inner part of first furrow, the three furrows spaced about equidistantly. Glabellar tongue of same width at base as base of median glabellar area, gently convex, widening forward to anterior extremity, across which it is moderately convex; bounded laterally by axial furrow, which is continuous with preglabellar furrow. Latter furrow becomes progressively shal- lower and closer to anterior margin of tongue as it runs inwar.l, 402 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY and dies out before reaching mid-line (PI. 1, fig. 2). In anterior view outline made by anterior margin of preglabellar area is obtusely angulate (Pl. 1, fig. 2). Eye surface gently convex transversely, moderately convex longitudinally, more strongly curved in posterior than anterior half; external surface with many tiny, low facets arranged in diagonal lines (PI. 2, fig. 22). Palpebral rim (PI. 1, figs. 1, 2) widest adjacent to posterolateral part of median glabellar area, narrowing progressively forward to a minimum width where it joins anterior area of free check, latter merges with preglabellar area; palpebral rim narrows abruptly as it descends to occipital furrow. Anterior branch of suture runs inward in a curve to mid-line; posterior branch runs beside antero-lateral extremity of occipital ring, then out across cheek just inside posterior margin for a short distance before turning to cut the margin immediately outside the fuleral articulating socket (Pl. 1, fig. 5). Cheek outside eye lobe narrow, outward-sloping, separated from eye surface by convex external rim of eye lobe; broad base of librigenal spine in line (tr.) with posterior part of median elabellar area, spine tapers back to sharp point. Inside base of librigenal spine cheek depressed beside external rim of eye lobe, but outer part convex and extended posteriorly in a rounded projection which lies immediately outside the articulating socket (ed bg Ica geae? By Doublure of cephalon includes that beneath occipital rmg and that which extends around free cheek, which is narrow postero- laterally, but widens progressively to the mid-line, where it is crossed by the median suture. At the inner edge, for a short distance on each side of the median suture, there is an upward angulation in the doublure (PI. 1, fig. 10; Pl. 2, fig. 23). Beneath the eye lobe the innermost part of the doublure is thinner and is curled over, the curled portion being widest anteriorly. The edge of the doublure proximal to the curled part is thick and broad (Pl. 2, fig. 23), this edge being that along which the hypostomal suture runs. In the external surface of the doublure, and lying in the same line (exs.) as the outer end of the hypo- stomal suture, is a pit (Pl. 1, figs. 10, 11). On the inner surface of the doublure this pit appears as a cone, inclined so that the apex lies close to the angulation in the beginning part of the anterior sutural branch. The apex of the cone is broken in € WHITTINGTON : ORDOVICIAN TRILOBITES 403 many specimens but appears to have a minute opening at the tip when undamaged (PI. 2, figs. 23, 24). It seems that the tip rested in a pit in the inner surface of the cranidium, at the junction of the axial and preglabellar furrows, which appear as low ridges on the inner surface (PI. 2, fig. 25). Hypostome gently convex, maximum width, across anterior wings, greater than length. Middle body not separated from anterior border by border furrow, but laterally outlined by nar- row furrow and posteriorly by broader, shallower furrow. Most of middle body oceupied by oval areas, most convex posteriorly, and oriented with long axis diagonal; the exoskeleton over these areas is thinner, so that they are more clearly outlined on the inner surface. The subtriangular strip separating the oval areas is not convex. Median boss short, blunt, backwardly directed. Anterior wing a short, rounded projection of noticeably thinner exoskeleton, lacking a wing process. Inner edge of lateral border a sharp ridge, this ridge continued across postero-lateral border and into base of short, blunt spine; posterior border wider than lateral. Exoskeleton at inturned, upwardly-directed edge of lateral and posterior borders thin, the edges scalloped (PI. 1, figs. 12-15) ; in front of the posterolateral projection this exoskeleton erossed by a flexure and part of the inner edge is twisted and projects to form the posterior wing; the edge of the portion along the posterior border displays a broad, deep median notch, outside the edge of which is a small, semicircular notch. The broad, flat anterior edge of the hypostome at the suture is inclined forward medially, twisting to incline back laterally, and thus fits close against the correspondingly inclined surface of the suture on the free cheek (PI. 1, figs. 11, 12). Dorsal external surface of exoskeleton, except in furrows, bearing fine raised lines in Bertillon pattern which varies little between individuals. Minute tubercles are also scattered over the surface, and become coarse and readily visible on the gla- bella adjacent to the palpebral and occipital furrows, and along the posterior edge of the occipital ring. On the doublure and genal spine (upper and lower surfaces) the Bertillon lines run parallel to the margins and heavier terrace lines occur with them. External surface of hypostome bearing lines that run sub-parallel to the margin on the borders, but run longitudinally on the middle body. These lines are stronger than the Bertillon 404 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY lines of the glabella, and on the borders especially are of equal strength to the terrace lines. Because of the thinness of the exoskeleton over the oval areas the lines are prominent over these areas and help to outline them. Number of thoracic segments unknown, but eleven is typical for the genus. Axis moderately convex, tapering back, width more than half, and posteriorly three-quarters, of that of thorax. Axial ring sloping forward medially, gently inflated distally, lateral margin semicircular in outline and descending vertically to axial furrow. Narrow articulating furrow and long (sag.) articulating half-ring. Median axial spine (PI. 2, figs. 11, 12, 18) on fourth segment in front of pygidium (? eighth in thorax) directed backward just above thorax, sharp point extending to, or beyond, tip of pygidium, ventral side flattened, dorsal convex. Inner part of pleura narrow (tr.) in anterior segment, diminish- ing progressively in width posteriorly. Outer part of pleura gently sloping in anterior segments, slope increasing posteriorly ; outline subrectangular in anterior segments, with rounded antero-lateral corner and small point on postero-lateral corner ; outline changes progressively posteriorly as outer part of pleura is more backwardly curved and drawn out into a longer point. Fuleral articulating socket and process, and adjacent edges of inner part of pleura, strongly raised ; apex of triangular flattened areas of inner part of pleura between them leads into a deep depression immediately in front of, and extending outside of, fuleral socket: this depression appears to constitute the pleural furrow, which dies out rapidly distally. Axial socket and process small. Doublure extends inward to fulcrum, except along pos- terior edge where a narrow tongue continues inward along outer edge of fuleral articulating socket; doublure is bent upward at inner edge, and inner posterior portion is crossed transversely by a sharp flexure, the vertical portion of the flexure facing posteriorly. Because of this flexure an inner, posterior part of the doublure forms a depression which accommodates the antero- lateral part of the succeeding segment during enrollment. The amount of angular movement possible between the segments is thus limited, and in those following the axial spine it 1s slight. Pygidium subrectangular in outline, length greater than width. Convex axis semicircular in outline, width three-quarters that of pygidium at anterior margin, but extending little more WHITTINGTON : ORDOVICIAN TRILOBITES 405 than one-quarter of length, divided into two rings. First ring shortest and slightly depressed (sag.) in mid-line, articulating furrow running in curve coneave forward; first ring furrow more strongly curved and outline concave posteriorly except distally, where it runs transversely; lobate outer portion descends ver- tically to axial furrow. Second ring bilobate, with a_ broad, deep median depression. Articulating furrow deepest distally, articulating half-ring long (sag.). Pleural regions gently con- vex medially, descending vertically to lateral margins, promi- nent articulating boss at antero-lateral corner, outside and below which is facet; pleural and interpleural furrows absent, pos- terior margin drawn out into two pairs of pleural spines, the inner pair the larger. Broad doublure, inner part of which is curved dorsally, so that margin hes immediately ventral to sharp flexure at margin of axis. There is some individual variation displayed by the pygidium —in the degree of inflation of the axis, and particularly in the outline, degree of inflation, and relative length of the inner pair of pleural spines. In the original of Plate 2, figure 14, the inner pleural spines are inflated, sepa- rated at the base by a deep cleft, and much longer than the outer pair; in the original of Plate 2, figure 20, the inner spines are relatively shorter and not inflated; in the original of Plate 2, figure 15, though inflated, the inner spines are relatively short and blunt. External surface of posterior margin and inflated distal part of axial rings of thorax and pygidium tuberculate, less conspic- uous tubercles may be scattered over pleural regions. Except in furrows and on articulating half ring Bertillon lines run transversely on axial rings and diagonally on pleurae of thorax, subparallel to margin on pleural region. On median axial spine terrace lines run longitudinally, and stronger ones occur at intervals. Similarly, strong terrace lines run longitudinally on thoracic doublure and subparallel to margins of pygidial doublure. Development Protaspis. Two complete and 15 broken specimens of a pro- taspis from locality 6, and 1 and 6 broken specimens from local- ities 3 and 4 respectively, have been obtained. This protaspis is 406 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY exceedingly like that of the closely related younger species R. plaesiourus. Protaspis (Pl. 3, figs. 1-5; Text-fig. 4) subspherical in shape, excluding spines maximum length 0.62 mm., maximum width 0.81 mm., maximum height 0.85 mm. Cephalon and protopy- gidium subequal in size. Occipital ring of width (tr.) about one-third total width, defined posteriorly by shallow ring furrow, anteriorly by extremely faint occipital furrow, and set off from pleural region by its convexity. Axial furrow in front of occipital ring is indicated by change in slope between gently convex gla- bella and cheek; axial furrows diverge slightly forwards and run in a straight course to margin. Glabella extends to anterior Figure 4. Remopleurides caclatus n.sp. A, B, C, protaspis, lacking free cheeks and hypostome, anterior, posterior, and left lateral views, X 26 (compare Pl. 2, figs. 2, 4). Abbreviations: a, anterior cephalic spine; ab, anterior branch of suture; fa, fixigenal spine; or, occipital ring; p, postero- lateral spine on protopygidium; pb, posterior branch of suture; pl, palpebral lobe; t, tuberele. edge of shield, which is bent down. Pair of tiny tubercles on anterior slope. Fixed cheek gently convex and outward-sloping, palpebral lobe not raised or defined other than by long, curved sutural margin; anterior branch of suture runs from palpebral lobe forward and outward, then curves inward to margin; pos- terior branch runs in ‘‘S8’’ curve and is directed more strongly outward. At anterolateral corner of fixed cheek there is a thick spine directed forward and slightly outward; at posterolateral corner there is a spine of similar size, the fixigenal, directed backward and more strongly outward. There is only the faintest trace of a transverse furrow, directed outward and slightly back, separating fixed cheek from pleural region of protopygidium, and WHITTINGTON : ORDOVICIAN TRILOBITES 407 running out immediately behind the fixigenal spine to reach the margin where it is cut by the extremity of the posterior sutural branch. Free cheek, cephalic doublure, and hypostome unknown. Protopygidium subrhomboidal in outline, well-defined, gently convex axis less than one-third total width, tapering only at blunt tip at posterior margin. In the complete protaspis three axial rings of about equal length (sag.) to the occipital, and a longer tip, may be distinguished. In some broken specimens (PI. 3, fig. 6) there are five axial rings in front of the tip. Pleural region slopes gently to lateral margin, but posterolaterally and beside tip of axis is bent down vertically, prominent spine at postero- lateral corner is situated at upper edge of this flexure. Doublure narrow and curled under, tiny median notch, greatest width posterolaterally, diminishing in width anteriorly to disappear at anterolateral corner. Along posterior margin of pygidium, be- hind main pleural spines, are 4 pairs of tiny spines, the middle pair close together at base of notch in doublure, the outer pair largest, and directly behind main spines. Degree 0. Reconstructed (Text-fig. 5) from the smallest crani- dia, free cheeks and transitory pygidia (PI. 3, figs. 7-15, 17, 18), all of which are like those of the younger species R. plaesiourus. Cranidia are known from localities 3, 4, and 6, free cheeks from 3 and 4, transitory pygidia from 2-4, 6, 7, 15, and 16. Differs from protaspis most obviously in better definition and subdivision of axial region, in reduction of fixigenal spine to a small nub, and absence of main posterolateral spines on transi- tory pygidium. The longitudinal convexity of the cephalon is such that, if the posterior margin of the occipital ring is oriented in the vertical plane, then the anterior slope of the glabella is vertical. The longitudinal convexity of the transitory pygidium and inclination of the articulating half ring is such that the position relative to the cephalon gives a subspherical shape to the exoskeleton. In several specimens a pair of small tubercles may be seen on the glabella, situated in line with the bases of the anterior cranidial spines and near the ill-defined glabellar margin. Palpebral lobe slightly raised above adjacent part of fixed cheek, notably posteriorly, where the angle the margin makes with the posterior sutural branch is more acute and more deeply indented than in the protaspis. Some specimens suggest 408 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY the presence of a faint, narrow palpebral rim. Free cheeks of smallest size (Pl. 3, figs. 17, 18) fit approximately, and are sepa- rated by median suture. The anterior branch of the suture has the characteristic curve so that the dorsal surface of free cheek narrows and disappears at mid-line. Outside eye lobe free cheek of uniform width and lateral outline consequently a convex curve; librigenal spine arises from border at a point opposite two-thirds the length (exs.) of the eye lobe, is broad at the base Figure 5. Remopleurides caelatus n.sp. Outline reconstruction of degree 0 individual, right free cheek omitted. A, exterior, B, oblique exterior view, X 26. Abbreviations: a, anterior cephalic spine; fz, greatly reduced fixigenal spine; lb, librigenal spine; t, tubercle. Based on originals of Plate 3, figures 9, 12. and tapering. Projection from inner part of posterior border of free cheek not developed. Eye surface narrow (tr.), facets relatively large, about 8 in a diagonal row. Doublure relatively broad in front of librigenal spine, narrow behind, inner margin excavated in broad curve along hypostomal suture, pit in doub- lure adjacent to extremity of this suture, pit appearing on inner surface as low, blunt cone. Hypostome unknown. Transitory pygidium subcireular in outline, strongly convex (sag. and tr.), most strongly so posteriorly. Width of axis about one-quarter maximum width of transitory pygidium, gently convex in anterior part, which is divided by furrows into a WHITTINGTON : ORDOVICIAN TRILOBITES 409 prominent articulating half-ring and five axial rings. Rings decrease in length and width posteriorly, the fifth markedly shorter than those in front of it; posterior part of axis strongly convex, undivided, tip steeply sloping. Four pairs of short spines along postero-lateral margin of pleural regions, doublure narrow, curled. Comparison of this transitory pygidium with the protopygi- dium of the protaspis shows the strong resemblance in outline and form between the two. Chief changes are the more convex axis, better definition of rings, and lack of main pleural spines in degree 0. Further development of cephalon. Cranidia of length (sag.) 0.85 (PI. 4, figs. 1, 2) to 0.97 mm. (PI. 3, fig. 16) from localities 2, 6, 15 seem to represent the next stage of growth. Occipital ring is wider (tr.) than the glabella between the posterior part of the palpebral lobes, moderately convex ; large low median tubercle close to anterior margin, row of 12 blunt spines evenly spaced along posterior edge. Occipital furrow with vertical anterior slope so that glabella adjacent to occipital ring stands higher than ring; glabella outlined by its convexity and by shallow axial furrows, expands forwards to maximum width (equal to that of occipital ring) opposite anterior part of palpebral lobe, is pinched-in opposite anterior end of palpebral lobe, then ex- pands forward again to equal greatest width at margin of tongue. Latter is bounded laterally by narrow band of fixed cheek and deep axial furrow, there is no anterior cranidial spine; anterior edge of tongue with evenly curved outline and narrow preglabel- lar field laterally, which narrows almost to disappearance as mid-line is approached. Palpebral lobe with broad, gently con- vex rim which is widest posterolaterally and narrows forward ; between rim and axial furrow is a small, erescentie flat area situ- ated in the inner, anterior portion of the lobe. Free cheek like that of degree 0. By the time the cranidium reaches a length (sag.) of about 1.24 mm. (PI. 4, figs. 7, 8) the glabella has expanded between the palpebral lobes so that the earlier flat part of these lobes is eliminated, and the lateral margin of the glabella is separated from the palpebral rim by a palpebral furrow. This furrow is 410 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY broadest at its mid-point, and in line with this point is the maxi- mum glabellar width, which is greater than that of the tongue or the occipital ring. Beyond this size there is no major change in morphology of the cranidium. As size increases, convexity of the glabella increases, both longitudinally and transversely, and the axial-palpebral furrow becomes narrow and deep, with a vertical slope down from the glabella. Three pairs of muscle scars have been observed on glabella of cranidia of length (sag.) 2.5 mm. and upwards. Small tubercles occur on the external surface of the tongue and on the glabella adjacent to the palpe- bral furrow in cranidia of this same size, the raised lines appear- ing at a larger size. Free cheek that fits a cranidium of length (sag.) about 1.2 mm. (PI. 4, figs. 11, 12) essentially like the largest, with narrow region outside eye lobe, doublure with pit, strong terrace lines and inner edge curled. Hypostomes of the smallest size known (PI. 4, figs. 3, 4) fit a cranidium of length (sag.) 2.5 mm., and differ from the largest in that the spine on the posterolateral corner is longer. Thorax and pygidium. Transitory pygidia from localities 3 and 4 are tentatively placed here. Tiny specimens of this type have not been recovered from localities 2, 6, or 15. Earliest in the presumed series are those having a length (sag., excluding spines) of 0.37 to 0.44 mm. (PI. 4, figs. 5, 6, 13, 16-18, 22), and bearing a prominent upwardly-curving median axial spine of length almost 2 mm. The axis is convex, three-fifths the total width, not sharply bounded from the less steeply-sloping pleural regions. Three pairs of thornlike pleural spines are about equally spaced along the lateral and posterolateral margins, and between the last pair, on the posterior margin, there may be a tiny fourth pair (Pl. 4, fig. 5). The doublure is narrow and curled under, with a median posterior notch. The axis is crossed by broad, shallow ring furrows, the rings gently convex, and in front of the swollen base of the median spine there may be two rings (Pl. 4, figs. 5, 6, 13), one ring (PI. 4, figs. 16-18), or none (PI. 4, fig. 22). In all there is a shallow articulating furrow and an articulating half-ring. In those with the median spine at the anterior end of the axis, axial rings cannot be seen behind the base of the spine. In the holaspis (PI. 2, fig. 12) the median axial spine is on the 4th segment in front of the pygidium, 1.e. WHITTINGTON : ORDOVICIAN TRILOBITES 411 the 8th thoracie segment if the total number is eleven. On this assuiuption the transitory pygidia with 2, 1 and no rings in front of the axial spine would be, respectively, those of degrees 5, 6, and 7. Specimens belonging to any one of these degrees vary in size within the limits mentioned. Those of degree 5 seem always to exhibit the fourth pair of pleural spines, but those of degrees 6 and 7 may, or may not, possess them. These transi- tory pygidia are similar to those described by Ross (1951, pp. 082-3, pl. 83, fig. 19; 1953, p. 634, pl. 62, fig. 20) as belonging to an older remopleuridid species. The second group of transitory pygidia (PI. 4, figs. 23-25) vary from 0.8 to 0.4 mm. in length (sag., excluding spines and articulating half-ring). The convex axis tapers backward and extends close to the posterior border. Up to four rings may be outlined by shallow ring furrows, the articulating furrow and anterior ring furrow being the deepest, those following being progressively shallower and the rings shorter (sag. and exs.). The third ring may show a faint median longitudinal depression, like those on the rings of the holaspid pygidium, and the fourth ring may be bilobate. Pleural regions narrow, less steeply slop- ing, lateral and posterolateral borders bearing triangular spines. These spines are differently developed on different specimens (probably corresponding to different degrees) — three pairs sub- equal in size and a tiny fourth posterior pair (PI. 4, fig. 24), three subequal pairs (PI. 4, fig. 25), or three pairs with the third small (Pl. 4, fig. 23). The doublure is curled under and bears the prominent terrace lines. Since these transitory pygidia bear no trace of the median axial spine, it seems hkely that they belong to degrees 8-10, i.e. those succeeding the release of the spine bearing segment into the thorax. If they belonged to early degrees (1 to 4) they might be expected to show some sign of the median axial spine, unless this spine appears abruptly be- tween moults. Further, the tip of the axis of the degree 0 transi- tory pygidium is strongly convex, quite unlike that of this type. The smallest pygidia that are probably those of the holaspid (Pl. 4, fig. 21) are about 0.49 mm. in length (sag.), and are similar to larger specimens. 412 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY REMOPLEURIDES PLAESIOURUS N..sp. Plate 4, figures 26-30; Plate 5; Plate 6, figures 1-15. Holotype. USNM 137677 (Pl. 5, figs. 3-5), locality 9. Other Material. Paratypes USNM 137678a-e; all figured specimens in U.S. National Museum. Geological Horizon and Localities. Lower Martinsburg shale, localities 9, 10, 12. Description. This species is exceedingly like R. caelatus n.sp., differing from it in minor details of outlines, convexities, and proportions of parts of the exoskeleton. Comparisons between the photographs show that in R. plaesiourus: a) glabellar tongue (compare PI. 5, figs. 3, 4 with Pl. 1, figs. 3, 8) appears relatively narrower, is more inflated so that the slope down to the axial furrow is vertical, and the anterior area of fixed cheek is slightly wider. The preglabellar area dies out well before reaching the mid-line; in exterior view the outline of the anterior margin of the preglabellar area is almost straight, rather than obtusely angulate ; b) outline of palpebral rim in dorsal view (compare PI. 5, fig. 5, with Pl. 1, fig. 5) is less curved in anterior half, more curved in posterior half, and the rim itself is wider (exs.) imme- diately in front of the occipital ring ; c) the different shape of the tongue and palpebral rim means that shg¢ht differences in shape are also present in the free cheek (compare PI. 5, figs. 6, 7, with Pl. 1, figs. 4, 16) — in the outline of the inner margin of the eye surface and the course of the an- terior branch of the suture. In addition, the lbrigenal spine is differently placed, and there is a deeper and wider notch between the base of the spine and the posterolateral part of the cheek ; d) the hypostome (compare PI. 4, figs. 26, 29, 30 with Pl. 1, figs. 9-15) has a different outline, the anterior margin less strongly curved, the posterolateral part of the border slopes more steeply, the median boss is higher and sharper, the oval areas less inflated posteriorly ; e) the pygidium (compare PI. 5, figs. 8, 9, 12-15 with Pl. 2, fies. 14-17, 19-21) shows the same kind of individual variation as that seen in R. caelatus in the relative length and amount of inflation of the inner pleural spine. Yet the inner pair of spines is relatively wider at the base and longer; the lateral margins WHITTINGTON : ORDOVICIAN TRILOBITES 413 of the pygidium are parallel-sided rather than backward converg- ing and a low but distinct post-axial ridge is present, extending about half-way to the base of the notch between the inner pleural spines ; f) pattern of Bertillon lines on eranidium (compare Pl. 5, fig. | with Pl. 1, fig. 1) is different, and there are more granules in the mid-region of the median glabellar area. Median region of pygidium is also more granulate (compare PI. 5, fig. 15 with Ele ariotol Ot Development. The development of Remopleurides plaesiourus is so like that of R. caelatus that much of the description given of the latter would apply to both species. The material of R. plaesiourus is less abundant and less complete in some respects. The protaspis (PI. 5, figs. 18-20, 26) is broken at the anterolateral corners, so that the anterior cephalic spines are not seen. The protopygidium shows five axial rings, the fifth faint, in front of the convex tip of the axis, as do some broken specimens of R. caclatus (Pl. 3, fig. 6). In other respects the protaspides of the two species are alike. The degree 0 ecranidium of R. plaesiourus has shmmer anterior eephalie spines and the outline of the palpebral lobe is typical of this species in being more strongly curved posteriorly (com- pare Pl. 5, fig. 16, with Pl. 3, fig. 9). The transitory pygidium of degree 0 (PI. 5, figs. 21-23) is like that of R. caelatus, and the few specimens show a small range in size. A eranidium of length (sag.) 0.75 mm. (Pl. 6, fig. 1) shows a stage not found in the material of R. caelatus n.sp. It is very little larger than the degree 0 eranidium, the palpebral lobe has a faint rim, and the anterolateral margin continues the line of the anterior palpebral margin forward to meet the transverse an- terior margin at an obtuse angle. Anterior cephalic spines are lacking, and the symmetrical form (three additional specimens are known from loeality 10) suggests that the shape is not the result of breakage but of the inward shift of the anterior branch of the suture following the loss of the spine. The next largest cranidium, of length (sag.) 0.79 mm. (PI. 6, fic. 3) has the tongue with steeply sloping sides, and the anterior area of the free cheek. The clabella is narrowest in front of the occipital furrow and expands forward at an even rate to the 414 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY maximum width at the anterior end of the tongue. The palpebral lobe has a well-developed rim, broadest posteriorly, and enclosed between the are of the palpebral furrow and the straight axial furrow is a horizontal area. This area is larger than that in cranidia of the same size of R. caelatus (P1. 4, figs. 1, 2) because in this latter specimen the glabella expands toward the mid- length of the eye lobes. The free cheek fitting the small erani- dium of R. plaesiourus (PI. 6, fig. 4) has the curve of the inner margin of the visual surface, and deep notch between base of spine and lateral margin, characteristic of this species (com- pare with that of R. caelatus; Pl. 3, fig. 16). In the next two larger cranidia (PI. 6, figs. 2, 5, 7, the original of fig. 5 of length (sag.) 1.27 mm.), the glabella bulges out between the eye lobes, and in larger cranidia (PI. 6, figs. 8-12, 15) occupies the space between the palpebral furrows. From this size onwards there is little further change. Transitory pygidia between degree 0 and probable degrees 8-10 are not known in R. plaesiourus, but those of degrees 8-10? (Pl. 5, figs. 24, 25) bear three pairs of marginal spines and are like those of R. caelatus. REMOPLEURIDES CAPHYROIDES Ni. sp. Plates 7-10; Plate 11, figures 1-4; Text-figure 6 Holotype. USNM 137681 (PI. 7, figs. 1-3, 8,10; Pl. 8, figs. 8, 10), locality 3. Other Material. Paratypes USNM 137682a-e; all figured specimens in U. S. National Museum. Geological Horizon and Localities. Edinburg formation, localities 2-6, 15. Fairly common at localities 2-4, rare at the others, especially at 6, where Rk. caelatus is common. Description. All parts of the exoskeleton of this species may be differentiated from those of Remopleurides caelatus by the following characters: (1) Glabella (compare PI. 7, figs. 1-3 with Pl. 1, figs. 1-3, 5, 8) is more convex (sag. and tr.), especially anteriorly, and the relatively wider and more inflated tongue overhangs the anterior cephalic margin; palpebral rim is narrower in anterior part, and anterior area of fixed cheek is narrower. Outline of anterior edge of cranidium a curve, concave (rather than convex) ven- trally. The two posterior glabellar furrows are faintly visible WHITTINGTON : ORDOVICIAN TRILOBITES 415 as curved, smooth strips, and the short anterior furrow is rarely visible (compare PI. 7, figs. 3, 8 with Pl. 1, figs. 5, 6); all the furrows are less conspicuous than in R. caelatus. Posterior area of fixed cheek short (exs.) and narrow (tr.), the anterior margin a gentle ‘‘S’’ curve. (11) Free cheek (compare PI. 7, figs. 1, 3-7, 9, 10 with Pl. 1, figs. 4, 10-12, 16) with anterior part of eye surface overhanging margin, external rim of eye lobe less conspicuous; doublure ad- jacent to median suture wider (sag. and exs.) and with more curved anterior margin; posterior margin outside suture curves gently forward to base of librigenal spine —there is no pos- teriorly directed projection outside the articulating socket, and the base of the librigenal spine is broader and the spine is dif- ferently directed. As in R. caelatus there is a pit in the doublure, appearing on the inner surface as a long, slim cone, the tip of which rests in a pit at the junction of the axial and preglabellar furrows (compare PI. 8, figs. 8-10 with Pl. 2, figs. 28-25). (111) Hypostome (compare Pl. 7, figs. 4, 5, 7, 9 with Pl. 1, figs. 9-15) strikingly different, being transversely rectangular in outline with wider borders, prominent shoulder and _ postero- lateral projection. Middle body separated from anterior border by shallow furrow, the oval areas with the long axis more widely divergent and enclosing a larger triangular area; on the anterior margin in the mid-line is a low rounded median boss. Anterior wing long and slim, no wing process, curving forward and up- ward so that distally it lies above the curled edge of the free cheek doublure. Posterior wing a large blunt projection. (iv) Thoracie segments (compare PI. 9, figs. 1-5, 7, 8, 12 with Pl. 2, figs. 1-13, 18, 19) have distal parts of axial rings only gently convex, less prominent axial articulating processes and sockets, longer (exs.) and wider (tr.) pleurae, flattened and tapering more gradually. The median spine on the eighth (?) thoracic segment is relatively slimmer and shorter. (v) Pygidium (compare PI. 9, figs. 6, 9-11, with Pl. 2, figs. 11-18, 18-21) lacks ring furrow defining median part of first axial ring, and pleural regions flattened, broadest at mid-length, bearing two pairs of border spines of which the outer are the longer. 416 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY (vi) External surface of glabella granulate in mid-part of median area (PI. 7, fig. 3), with a row of larger granules along the posterior margin of the occipital ring in all but largest indi- viduals (PI. 8, figs. 1, 3, 5). Bertillon lines present on anterior and anterolateral parts of glabella (PI. 7, figs. 1, 2) but not as conspicuous as in R. caelatus. On free cheek and hypostome, ter- race lines run sub-parallel to margins. On axial rings of thorax, lines run transversely and there may be a row of granules along the posterior margin. On dorsal surface of pleural spines of thorax and pygidium each terrace line runs in a characteristic inverted ‘‘V’’ course (PI. 9, figs. 3, 5, 6, 9, 12), and on the doub- lure the lines run sub-parallel to the lateral margins (PI. 9, figs. 2, 4, 8, 11): The eranidium, free cheek and hypostome have been associated by fitting together isolated specimens (Pl. 7; Pl. 8, fig. 7). The form of the occipital ring leads to association of the thoracic segments, and the pleurae of the latter may be matched in form and lines on external surface with pleural regions of the pygi- dium (PI. 9, figs. 1-12). Development The supposed developmental series of Remopleurides caphy- roides is illustrated in Plate 10 and Plate 11, figures 1-8. There is no complete size series of cranidia below a length of 1.7 mm. from any one locality. In this range, therefore, the series is tentative, being built up from specimens from different localities. The development of the cranidium will be described in deseend- ing order of size and the arguments given for the series here advocated. Cranidia of length (sag.) 2.5 mm. (PI. 8, figs. 5, 6) (or larger) down to 1.73 mm. (Pl. 10, figs. 1-4) show a gradual transition —in the smallest example the glabella is relatively narrower, in longitudinal profile seen to be more convex and strongly arched at the base of the tongue, the anterior outline a rounded, inverted ‘V’ rather than a broad curve; the palpebral lobe is relatively wider (tr. and exs.), especially in the posterior part; external surface more coarsely granulate. A free cheek that approximately fits this cranidium is much like larger ones; smaller specimens are not known. WHITTINGTON : ORDOVICIAN TRILOBITES AIT The next smallest cranidium comparable to the original of Plate 10, figures 1-4, is the original of Plate 10, figures 5-7, 0.92 mm. in length (sag.). The anterior part of the glabella has the same inverted ‘V’ outline in dorsal view and is more strongly raised, coming up to a blunt point in the mid-line at the base of the glabellar tongue; between the median part of the palpebral lobes the glabella is much narrower, the sides diverging forwards; a short basal glabellar furrow opposite the posterior part of the palpebral lobe, a longer median furrow opposite the middle part of the palpebral lobe — comparable positions to those of the mt Figure 6. Remopleurides caphyroides n.sp. Outline reconstructions of degree 0 individual, lacking free cheeks and hypostome. A, right lateral, B, oblique views, X 30. Abbreviations: a, anterior cephalic spine; mt, median tubercle of glabella; or, occipital ring; t, tubercle. Based on origi- nals of Plate 10, figures 8-14, 16, 17. furrows in larger glabellae. The palpebral lobe is subsemicircu- lar in outine, horizontal, and with a faint rim. It seems reasonable to suggest that the preceding stage of the eranidium (believed to be degree 0 by analogy with R. caelatus ) is that shown in Plate 10, figures 8-11, 13 (ef. Text-fig. 6) with the striking large median glabellar tubercle situated opposite the anterior end of the palpebral lobes. The occipital ring is relatively narrow, the glabella straight-sided and expanding for- ward ; two pairs of smooth areas are the lateral glabellar furrows, 418 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY situated in the same relative position as in the next larger crani- dium; about mid-way down the anterior slope is a pair of tiny tubercles. Anterior cephalic spines are long and curved, and there is a slight swelling at the tip of the posterior area of the fixed cheek suggesting a fixigenal spine; palpebral lobe gently convex and outward sloping, narrow rim. External surface granulate, prominent median occipital tubercle. The protaspis preceding this degree 0 cranidium can hardly be other than that shown in Plate 10, figures 15, 18-20, and Plate 11, figures 1-4, the cranidium differing in its greater longi- tudinal convexity, and in exhibiting slim fixigenal spines. This protaspis differs from that of Remopleurides caelatus (PI. 3, figs. 1-5) in the more elongate outline of the cranidium and protopy- gidium, the more convex glabella, which is not only more convex anteriorly but has the large median tubercle; narrow palpebral rim, anterior branches of the suture directed slightly inwards. The three pairs of spines— anterior cephalic, fixigenal, and protopygidial — are slimmer than those of R. caelatus. Protopy- eidium with five axial rings in front of terminal portion and four pairs of tiny spines on the posterior margin, as in R. caela- tus. Division between cephalon and protopygidium more clearly shown by shallow furrow that curves outward and backward from beside the posterior margin of the occipital ring and passes just behind the fixigenal spine. The degree 0 cranidium of R. caphyroides (Pl. 10, figs. 8-11, 13) is also distinguished from that of R. caelatus (Pl. 3, figs. 7, 9) by the outline, median glabellar tubercle, narrow palpebral rim, and slimmer anterior cephalic spines. The degree 0 transi- tory pygidium here referred to R. caphyroides (Pl. 10, figs. 12, 14, 16, 17) is extremely like that of R. caelatus (Pl. 3, figs. 8, 10-15), differing in being larger and relatively longer, the out- line being suboval rather than subcircular. These two transitory pygidia occur together at localities 2, 3, and 4, but can be sepa- rated by size (though each type shows some range in size) and outline, especially at locality 2 where both species are common. At locality 6, where R. caphyroides is extremely rare, the transi- tory pygidia are all of R. caelatus type. It may be added that the degree 0 transitory pygidium of R. plaesiourus (PI. 5, figs. 21-23) is of the same size and outline as that of the closely related R. caclatus. WHITTINGTON : ORDOVICIAN TRILOBITES 419 Comparison between the protaspides of R. caphyroides (PI. 10, figs. 15, 18-20; Pl. 11, figs. 1-3) and R. caelatus (Pl. 3, figs. 1-6) shows that the outlines and size of the protopygidia differ in the same way as do the degree (0 transitory pygidia. The de- eree 0 transitory pygidium of FR. caelatus is smaller than the protopygidium of the protaspis of R. caphyroides, whereas the degree 0 transitory pygidium referred to R. caphyroides is larger. Small hypostomes of R. caphyroides (Pl. 9, figs. 16, 17) differ from the largest in the more square outline, greater convexity of the middle body, the presence of a blunt median spine (rather than boss) and a spine at the posterolateral angle. The smallest known pygidium of R. caphyroides (Pl. 9, figs. 18, 22) is 0.52 mm. in leneth and does not seem to be transitory — it differs from larger ones in that the inner pair of pleural spines is close together and partly fused. The degree 0 transi- tory pygidia of R. caphyroides and R. caelatus are similar, but there seems no reason to believe that the transitory pygidia of R. caelatus with the long median axial spine (PI. 4, figs. 5, 6, 13, 16-18, 22) may belong to R. caphyroides — the latter species does not have such a prominent median axial spine in the holaspid thorax and the shape of the pleural regions is different. The material described above has been found at the following localities: protaspis, localities 2-5; degree 0, localities 2, 3 (transi- tory pygidium only), 4, 6, and 15 (cranidium only); parts of larger exoskeletons are relatively abundant at localities 2 and 3, few at localities 4, 6, and 15, and unknown from 5. From loeali- ties 3 and 4+ come eranidia of length (sag.) 1.8 mm. (PI. 11, figs. 5-8) which in size and other characters are intermediate between the originals of Plate 10, figures 1-4 and 5-7 —~in outline of elabella between palpebral lobes, in form of lateral glabellar furrows, and size of papebral lobe. The anterior part of the elabella, however, has a broadly rounded outline in dorsal aspect, and in lateral aspect lacks the sharp convexity. Further, and also only from localities 3 and 4, comes a degree 0 eranidium (PI. 11, figs. 9-11) like the original of Plate 10, figures 8-11, 13, except that it lacks the large, rounded median tubercle on the anterior part of the glabella. No protaspis of this type is known. Two possible explanations occur for these cranidia: (a) they are part of the developmental series of an otherwise unknown species of 420 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY Remopleurides, or (b) they are part of the R. caphyroides devel- opmental series. Acceptance of the latter alternative would mean that in the early meraspid development of the cranidium of some individuals the median glabellar tubercle was abruptly reduced between protaspis and degree 0 and the glabellar tongue conse- quently rounded in dorsal outline. I know of no comparable case of morphological differences between groups of early meras- pides of a single species. REMOPLEURIDES ASPERULUS N.sp. Plates 12, 13. Holotype. USNM 137683 (Pl. 12, figs. 1, 2, 4, 6, 14), locality 13. Other Material. Paratypes USNM 137684 a-e; all figured specimens in U. S. National Museum. Geological Horizon and Localities. Upper Lincolnshire limestone, localities ley ales. Description. This species is the only remopleuridid in the upper Lincolnshire limestone, and the first to appear in the Middle Ordovician of the Shenandoah valley. The silicification is relatively coarse-grained, and the surfaces of most of the specimens have granules (probably of quartz) adhering to them or growing out of them. A fairly complete growth series is known. Cranidium (PI. 12, figs. 1, 2, 4, 6, 14) wider than long, maxi- mum width at the mid-length, the margin of the palpebral lobe projecting farther out than the tip of the posterior area of the fixed cheek. Free cheek (PI. 12, figs. 3, 5, 11, 20) correspondingly narrow, the convex eye surface overhanging the margin antero- laterally, the postero-lateral portion, outside the external rim of the eye lobe, narrow, triangular, steeply sloping, with a blunt, short librigenal spine. Occipital ring moderately convex, longest (sag.) in the mid-line, deep, narrow occipital furrow. Remainder of glabella evenly and moderately convex (tr. and sag.), out- lined by narrow palpebral and axial furrows; glabellar tongue two-fifths maximum width of cranidium, bent down steeply to overhang slightly anterior margin of cephalon. Glabellar fur- rows visible on inner surface as grooves (Pl. 12, fig. 14), basal furrow long, curved, opposite greatest width of glabella, second furrow shorter, curved, opposite a point about one-third length of palpebral lobe. Palpebral lobe widest posteriorly, narrowing WHITTINGTON : ORDOVICIAN TRILOBITES 421 progressively forward and continuous with extremely narrow anterior area of fixed cheek. Doublure of cephalon widest an- teriorly; beside median suture a narrow posterior portion is flexed up vertically and then bent back horizontally (Pl. 12, fig. 3) —this flexure dying out rapidly away from the median suture; anterolaterally doublure curled up at inner edge, and extending back to where the suture cuts posterior cephalic mar- gin. Pit in doublure (PI. 12, fig. 11) appearing as a long, slim cone on inner surface (PI. 12, fig. 3); tip of cone lies close to anterior pit. Hypostome (PI. 12, figs. 7, 8, 12) subrectangular in outline, wider than long, middle body with prominent median boss and diagonally directed oval areas; borders broad, blunt spine at posterolateral angle, anterior wing short, posterior wing small. Thorax of at least nine segments (PI. 12, figs. 15-18). Axis broad, two-thirds width of segments, pleurae steeply sloping, with short pleural spines except for long spine on fifth segment from posterior. Short, blunt median axial spine on fourth seg- ment from posterior. Prominent axial articulating processes and sockets. Doublure extends in to axial furrow, posterior part crossed by a sharp, curving and transversely directed flexure which borders a depressed (in ventral view) posterior area that receives the anterior part of the succeeding segment during en- rollment (see Pl. 12, fig. 17, anterior segments on left side). Pygidium (PI. 12, figs. 15-18) wider than long, broad, short axis tapers rapidly to rounded termination, first axial ring narrow (sag.) medially and expanding laterally into a lobate area, second axial ring bilobate and lying between outer parts of first ring. Steeply sloping pleural regions having two pairs of short, blunt, posteriorly directed spines, deep notch between inner pair, smaller notch between the pairs. Doublure broad, bent upwards, median posterior projection. External surface bearing anastomosing terrace lines — on cheek (PI. 12, fig. 20) running sub-parallel to margins, on thorax and pygidium (PI. 12, fig. 15) transversely on axis, on pleurae curving distally to run subparallel to long axis. Along posterior margin of axial rings a row of low tubercles, largest laterally. On doublure (PI. 12, figs. 11, 17) terrace lines strong and every- where subparallel to inner margin; on hypostome lines run diagonally on middle body, and along the borders. 422 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY Development Protaspis (Pl. 13, figs. 1-4, 10). Subspherical in shape, con- vex cranidium with protopygidium curled closely beneath it. Owing to the coarse grain of the replacement, details cannot be observed. The axis is not defined. In lateral view scalloped lateral margin of cranidium shows position of palpebral lobe and course of anterior branch of suture. A second incomplete speci- men (not illustrated) shows four pairs of tiny spines along the curled posterior margin of the protopygidium, and a larger pair of spines on the steep flanks in front of the outermost marginal pair. Anterior cephalic spines are either absent or not preserved. What may be the fixigenal spine is visible a short distance inward and backward from the base of the palpebral lobe. Degree 0 (Pl. 138, figs. 5-7). Cranidium believed to be this degree, of length 0.83 mm., gently convex transversely, strongly convex longitudinally so that short (sag.) glabellar tongue de- scends vertically. Maximum width of cranidium at about half length and almost same as length (sag.), palpebral margin pro- jects out farther than posterior area of fixed cheek. Occipital ring is well-defined, convex, width about one-third that of erani- dium, with median tubercle, and in front of it axial furrows extend directly forward a short distance, outlining the gently convex posterior part of the glabella, and then die out. One specimen (not illustrated) suggests that a short anterior cepha- lie spine is present; fixigenal spine absent. External surface eranulate. Transitory pygidium (Pl. 13, figs. 11-15) of length ca. 0.9 mm., maximum width four-fifths of length; at three-quarters the length high and strongly convex so that lateral and posterior slopes are vertical, maximum convexity and height at point of greatest width. Axis about one-third total width, with prominent articulating half ring; six gently convex rings progressively shorter (sag.) posteriorly, and a more convex semi-cylindrical posterior portion situated on the posterior slope. Each ring is separated from the others and the posterior portion by a faint ring furrow, and there is a stronger articulating furrow, but axial furrows as such are not present. Doublure is present around postero-lateral margins, curled under, and along the curled edge is a curving row of 5 spines, the innermost pair WHITTINGTON : ORDOVICIAN TRILOBITES 423 close together on the ventrally-facing surface of the doublure immediately behind the tip of the axis, the other spines evenly spaced along the pleural margins. External surface finely granulate, the granules present on the axial rings but not in the furrows. On the posterior parts of the pleural regions curv- ing granulated bands, separated by narrow smooth bands (PI. 13, fig. 14), run out from the last 3 rings to the pleural margin. Each of these bands ends against the base of a marginal spine, and the second, third and fourth pairs seem thus to belong to the three axial rings in front of the terminal axial portion. Further Development. The next smallest cephalon (Pl. 13, fig. 17), eranidium of length (sag.) 1.0 mm., has the axial fur- row visible as a change in slope between glabella and palpebral lobe, extending forward from the margin of the occipital ring to the edge of the tongue in a curve gently convex outward. alabellar tongue is curved gently downward, width about half maximum width of cranidium. Palpebral lobe is gently convex, rim distinct and widest posteriorly. Free cheek with long, slim librigenal spine. At the next size known (Pl. 13, figs. 20-22, cranidium of length 1.4 mm.) the glabella extends between the palpebral furrows and is gently and evenly convex transversely. Width (tr.) of tongue is more than half maximum width of eranidium. As size of cephalon increases (Pl. 13, figs. 23-31), the major changes that take place are relative increase in maxi- mum width of cranidium so that it becomes greater than length, and this is accompanied by increase in curvature of the palpe- bral rim in dorsal view, and steeper down-bending of glabellar tongue so that the tongue becomes barely visible in dorsal view. The librigenal spine is progressively reduced to a blunt, short point. Granulation on the external surface becomes relatively finer, and can scarcely be seen in the larger specimens, partly owing to the preservation. Glabellar furrows are faint or invi- sible in most specimens, an exception being the original of Plate 13, figures 24-26, in which the furrows are represented by shallow grooves in the external surface. Smallest hypostome known (Pl. 13, figs. 8, 9) differs from largest principally in having a longer posterolateral spine. An example of the transitory pygidium (PI. 13, figs. 16, 18, 19) probably belongs to the later degrees of the meraspid period. 424 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY The first ring of the broad axis is continuous with the posteriorly directed spinose pleura, and the articulating half ring and pro- cess 18 typical. Behind and inside this segment the next ring is demareated, and there are two pairs of pleural spines; between the median pair of spines is the deep notch characteristic of the holaspis. REMOPLEURIDES EXIMIUS N.sp. Plates 14, 15, 16. Holotype. USNM 137685 (Pl. 14; Pl. 15, fig. 1), locality 4. Other Material. Paratypes, USNM 137686a, b; all figured specimens in U. 8. National Museum. Geological Horizon and Locality. Lower Edinburg limestone, localities 3 and 4. One specimen of the degree 0 transitory pygidium attributed to this species comes from locality 2. Description. The holotype specimen is an exceptionally com- plete exoskeleton lacking only the right free cheek and the hypostome. Only a few other exoskeletal parts come from locality 4, but these include the hypostome. At locality 3 this species is less common than Remopleurides caphyroides n.sp. and about as abundant as R. simulus n.sp. R. eximius n.sp. resembles closely R. asperulus n.sp. from the Upper Lincolnshire limestone, and it is largely because of this similarity that a hypostome occurring at localities 3 and 4 has been attributed to it. R. eximius may be distinguished from FR. asperulus by the following characters : — Glabellar tongue (compare PI. 15, fig. 1 with Pl. 12, fig. 4) of width (tr.) about half that of glabella, palpebral rim narrower posteriorly and outline in dorsal aspect evenly curved rather than a markedly stronger curvature in the posterior part. Free cheek extremely similar, but having a different curvature of the eye lobe and a more marked depressed region in the posterior part of the doublure. This depressed region receives the facet of anterior thoracic pleura during enrollment (PI. 14, fig. 2). Hy- postome (compare PI. 15, figs. 2, 4, 10 with Pl. 12, figs. 7, 8, 12) differs from that of R. asperulus in details of outline and in hav- ing a larger shoulder. Thorax (Pl. 14; Pl. 15, figs: 16-8; 11,12): of d1csegments;va longer pleural spine on the seventh and a median axial spine on the eighth. Pygidium (compare PI. 15, figs. 9, 18, 14 with PI. 12, WHITTINGTON : ORDOVICIAN TRILOBITES 495 figs. 15, 16, 18) distinguished by the convexity of the axis and consequent depth of furrows, including the median longitudinal ; posterior notch between inner pair of pleural spines broader. Development of Remopleurides eximius u.sp. ? Protaspis. A distinctive protaspis (Pl. 16, figs. 1-7, 9), quite abundant at locality 3 but also present at locality 4, has been attributed to this species because of its resemblance to that of Remopleurides asperulus nu.sp. This protaspis is far better pre- served than that of R. asperulus, though in neither case is the free cheek or hypostome known. Cranidial portion evenly con- vex (sag. and tr.), the occipital ring (PI. 16, fig. 2) faintly out- lined by both occipital and axial furrows and width of about one-third that of the cranidial portion. Conspicuous pair of tubercles close to mid-line and situated near anterior margin of the glabellar tongue (PI. 16, figs. 4,6). Margin of palpebral lobe with narrow rim, branches of suture distinct. Curving outward and backward from occipital furrow is a faint change in slope, the posterior border furrow (PI. 16, fig. 2). Subparallel to this furrow and running out from the posterior margin of the occi- pital ring is a faint furrow that marks the boundary between the cranidial and protopygidial portions (Pl. 16, fig. 7). Short slim anterior cephalic spine, and the fixigenal spine of similar size; the latter situated close to the cranidial margin and at about half the width of the posterior border. Protopygidium strongly convex (sag. and tr.), especially posteriorly. At the curled posterolateral margin may be seen four pairs of tiny spines, a fifth, outer and larger anterior pair situated slightly farther inside the margin (PI. 16, fig. 6) ; these five pairs about equi-distant from each other. Segmentation of the protopygi- diui is revealed by the transverse bands alternately finely gran- ulate and smooth (PI. 16, figs. 7, 9). These bands run subparallel to the faint furrow which separates cranidium and protopygi- dium and curve downwards and backwards over the steep sides. At least five bands can be distinguished following the occipital ring, but separate bands cannot be made out on the vertical lateral portion. Axis extremely faintly outlined in the first two of these bands following the occipital ring. The doublure is rela- tively broad and curled under, extending laterally as far as the posterior edge of the cranidial portion. 426 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY Degree 0. Both the cranidium (PI. 16, figs. 8, 10-12) and tran- sitory pygidium (PI. 16, figs. 13-17) are relatively abundant at localities 3 and 4 and are like those of R. asperulus (Pl. 13, figs. 5-7, 11-15), notably in the distinctive form of the transitory pygidium. Oecipital ring of cranidium convex, with median tubercle, occipital furrow deep, continued across fixed cheek by deep posterior border furrow. Glabella convex (sag. and tr.), tongue bent down vertically, anterior cephalic spine at antero- lateral corner long and slim. Two pairs of lateral glabellar fur- rows visible (Pl. 16, fig. 10) in posteromedian part of glabella as smooth ovate areas; the anterior furrow directed diagonally inward, the posterior directed diagonally inward and forward. In anteromedian part of glabella is a narrow, sagittal, smooth band which dies out at base of tongue. Pair of tubercles close to mid-lne of tongue (PI. 16, fig. 8). No fixigenal spine on pos- terior border. Transitory pygidium with large articulating half ring. Along posterior margin 5 pairs of short spines arranged in a curving line (Pl. 16, fig. 17). Segmentation in the transi- tory pygidium again indicated by alternate granulate and smooth bands which curve downward and backward over the steeply sloping lateral portions. Six granulate bands may be recognized (Pl. 16, fig. 13), separated by five smoother bands, and on the vertical slope behind the sixth band may be seen (PI. 16, fig. 16) a long narrow convexity representing the posterior part of the axis. Developmental stages intermediate between those of this degree 0 and the supposed holaspis are not known. The degree 0 crani- dium (Pl. 16, figs. 8, 10-12) is quite different in outline and convexity from that (PI. 11, figs. 9-11) here placed in R. caphy- roides ? REMOPLEURIDES SIMULUS N.sp. Plate 17; Plate 19, figures 11, 12. Holotype. USNM 137687 (Pl. 17, figs. 1, 2, 4, 5), locality 3. Other Material. Paratypes USNM 137688a-c; all figured material in U. S. National Museum. Geological Horizon and Locality. Lower part of Edinburg limestone, locality 9 vo. WHITTINGTON : ORDOVICIAN TRILOBITES 497 Description. This species, the fourth to be described from locality 3, does not resemble closely either Remopleurides caela- tus or R. caphyroides, but differs from R. eximius in the follow- ing minor characters (these characters also serve to distinguish this species from R. asperulus, which occurs at a lower horizon) : Glabella with relatively slightly wider tongue (compare PI. 17, fig. 1, with Pl. 15, fig. 1); the glabella and tongue are gently convex (sag. and tr.), the longitudinal convexity less than that in R. eximius (compare PI. 17, fig. 2 with Pl. 14 fig. 3). Outline of palpebral rim gently curved, only slightly curved in posterior half, free cheek with slim lbrigenal spine situated in advance of the genal angle, the base being about opposite the posterior part of the eye lobe. Hypostome (compare Pl. 17, fig. 18 with Pl. 15, fig. 2) with strong curve to anterior margin, postero- lateral corner angulate but not drawn out into a spine. Thorax (compare Pl. 17, fig. 11 with Pl. 15, fig. 7) distinguished by the flattened appearance of the axial rings, the tuberculation on the posterolateral margin being fine; pleurae likewise flattened, bladelike, the posterolateral corner extended into a blunt, back- wardly-directed spine. The fifth thoracic segment from the posterior (probably the seventh segment) does not have elongated pleurae but the succeeding segment (probably the eighth) has a slim median axial spine. Pygidium (compare PI. 17, figs. 11, 19 with Pl. 15, figs. 9, 13) with less inflated axis and consequent shallower furrows. Articulating process outside axial furrow prominent. ‘'wo pairs of pleural spines on posterior margin equally spaced from each other and without a markedly deeper median notch between the inner pair. It may be argued that the exoskeletons included here under R. simulus may be sexual dimorphs of those exoskeletons included under FR. eximius. An admittedly inconclusive argument against this view is that R. simulus does not occur at locality 4, where R. eximius is present. While localities 3 and 4 are not far apart geographically there may be a slight difference in horizon be- tween them, as exemplified by the species peculiar to each locality (see lists above). No early developmental stages of this species are known. Remopleurides rugicostatus Raymond (1925, p. 57, pl. 3, fig. 3; this paper, Pl. 18, figs. 23, 24, 26, 27), from the Ridley limestone 428 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY (Wilderness stage) of Tennessee, is based on a single, incomplete, enrolled exoskeleton that probably has the outer exoskeletal layers missing. The cranidium and anterior thoracic segments are like those of R. simulus, and on the anterior pleural band of both species raised lines run in convex curves forward and out- ward from the pleural furrow. The pleura of the seventh seg- ment does not appear to be elongated in R. rugicostatus ; the axial ring of the eighth segment is broken. Free cheek, hypostome and pygidium are unknown in RF. rugicostatus, and because of this lack of knowledge a new name is given to the Virginia material ; clearly R. rugicostatus and R. simulus are closely related, the former being slightly the younger (Cooper, 1956, Chart I). Genus ROBERGIA Wiman, 1905 RopBerciA MAJoR Raymond, 1920 Plate 18, figures 1-22, 25. Material, Localities and Geological Horizon. Lower part of Edinburg lime- stone (Liberty Hall facies), locality 14, a few cranidia and hypostomes from locality 16. Description. This species was illustrated by Raymond (1925, pp. 60-61, pl. 3, figs. 6-10) and later redescribed by Cooper (1953, pp. 22-23, pl. 8, figs. 7-11). At the same time Cooper (1953, p. 22, pl. 12, figs. 10-14) redeseribed the species Robergia athenia Butts, 1926, and some of the material he illustrated came from the present locality 14. Cooper stated (1953, p. 22) that R. athenia ‘‘differs from R. major chiefly in the shorter, more ex- panded frontal lobe of the glabella, by possessing a broad un- tapered axial lobe on the thorax, and by having the middle pair rather than the outer pair of pygidial spines longer than the other two.’’ I consider the first two of these distinctions of doubtful value, and a re-examination of the type material of R. major shows that the pygidium bears three pairs of pleural spines, the outer pair extremely small and short, the middle pair the longest, the inner pair short and close together near the midline. The pygidium of R. athenia appears to be extremely similar and I therefore regard these two species as one, and they come from approximately the same horizon. WHITTINGTON : ORDOVICIAN TRILOBITES 429 In redescribing the type species of Robergia (Whittington, 1950b, pp. 543-544, pl. 71, figs. 1-8), I pointed out that the hy- postome was not known from the Swedish specimens. Raymond described the hypostome of R. major as bifurcated. The original material includes one specimen marked by Raymond as being the hypostome, but this specimen is a poorly preserved external mould of a pygidium. At locality 14 R. major is the most abun- dant of two remopleuridids that are present, and the hypostome described below is thought to belong to it. Cranidium typical of Robergia, the glabella having 3 pairs of lateral glabellar furrows, the middle pair the longest and the most strongly curved. The basal glabellar lobe (PI. 18, figs. 3, 5), situated adjacent to the inner end of the eye lobe, is sub- circular in outline and has a faint independent convexity. Palpe- bral rim continuous with narrow anterior area of fixed cheek and preglabellar area (PI. 18, figs. 3, 4,7). Free cheek (PI. 18, fig. 2 subtriangular in outline, a narrow convex border, the base of the genal spine situated opposite the midpoint of the eye lobe, the spine long and slim. Eye surface (PI. 18, fig. 18) with many tiny facets arranged in diagonal lines. Doublure narrow, crossed anteriorly by a median suture. Hypostome (Pl. 18, figs. 8-14) subsquare in outline, anterior border flattened and continuous laterally with the large anterior wing. Lateral and posterior borders narrow, convex, separated from the middle body by deep furrow, small shoulder, spine at posterolateral angle. Mid- dle body bearing the characteristic pair of gently convex ovate areas; anteriorly these areas separated by a triangular depressed area, this depression interrupted by the median boss; posteriorly the oval areas separated only by a shallow furrow. Doublure narrow, small posterior wing. Pygidium (PI. 18, figs. 15, 19, 22, 25) subrectangular in out- line, shghtly wider than long. Axis at anterior margin one-third of width, tapering evenly back to reach about four-fifths of the length and continued to the posterior margin by a narrow post- axial ridge. Some five axial rings may be distinguished. Pleural regions horizontal, the outer pair of pleural spines situated on the lateral margin at about three-quarters the length, the middle and largest pair at the posterolateral corner, and the third pair close together near the midline. First two interpleural ridges 430 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY curve back to reach the posterior margin of the corresponding spine; third pair of interpleural ribs faint and close to the post- axial ridge. Doublure broad, inner margin reaching a line joining the anterolateral corner to the posterior tip of the axis. The specimens are not well preserved, being somewhat flattened and distorted. Details of the external surface cannot be dis- cerned except for the fine subparallel lines on the oval areas of the hypostome. Development. Small specimens of only the cranidium are known, the smallest being of length (sag.) 1.2 mm. (PI. 18, fig. 16). Glabella clavate in outline, the occipital ring narrow (sag.), convex, defined by a deep occipital furrow; the shallow axial furrows diverge forward to the maximum width opposite the anterior end of the eye lobes; in front of this point the glabella narrows slightly and is rounded. Three pairs of lateral glabellar furrows appear as short broad depressions, their positions being about as in larger cranidia, the posterior pair reaching the axial furrow, the anterior pair commencing inside the axial furrow and the shortest. Palpebral rim broad, convex, defined by a shallow furrow which runs anteriorly into the axial furrow. An- terior branches of the facial suture diverge forwards, then opposite the anterior end of the glabella curve around and run strongly inwards to meet the midpoint. A semicircular horizontal portion of the fixed cheek is thus enclosed between the palpebral rim and the axial furrow, and a further section of the fixed cheek and preglabellar area is enclosed between the anterior branches of the suture. The next largest cranidium (PI. 18, figs. 17, 20, 21) is of length (sag.) 1.88 mm. The glabella is now relatively broader and ex- pands between the eye lobes so that only a narrow crescentic depression separates the glabella from the palpebral rim. In position and length the lateral glabellar furrows are much as in the largest cranidia and the basal lateral lobes display the subcireular outline and gentle convexity. The anterior part of the glabella, however, does not expand in front of the eye lobes but narrows slightly so that outside the axial furrows there is still a relatively broad portion of the fixed cheek. The preglabel- lar area appears relatively shorter (sag. and exs.). The free cheek that fits this size of cranidium is similar in form to that of WHITTINGTON : ORDOVICIAN TRILOBITES 431 the largest specimens known. The next largest cranidium known (Pl. 18, fig. 4) is approximately 4 mm. in length (sag.). Evi- dently during development to this size the glabella expands between the eye lobes so that the lateral margin is adjacent to the palpebral rim, and also expands in front of the eye lobes so that the anterior area of the fixed cheek becomes narrower. The small pygidium (PI. 18, figs. 15, 19), of length 1.5 mm., (sag.) is from locality 16 and is not distorted. It shows clearly the form of the pleural spines, in particular of the outermost pair. Discussion of Development. Small cranidia of R. major (P1. 18, figs. 16, 17, 20, 21) are like those of Menoparia (Ross, 1951, pl. 81, fig. 14; 1953, pl. 62, fig. 3) in width (tr.) of the anterior area of the fixed cheek and shape of the preglabellar area. Robergia may therefore be more closely related to Menoparia than to Remopleurides. In the development of the cranidium of Robergia the same major features are shown as in the development of Remopleurides — the expansion of the glabella between the palpe- bral rims and the reduction in width of the anterior area of the fixed cheek and the preglabellar area. ROBERGIELLA n. gen. Type species. Robergiella sagittalis n. gen., n. sp. Discussion. The material included here occurs with Robergia major at locality 14, and with Remopleurides caelatus at locality 16, and displays holaspid characters that are intermediate be- tween them. The cranidium has rather the form and outline of Remopleurides, the eye lobe being relatively long and the palpe- bral rim broad; the glabellar tongue expands forward, but ap- parently not as strongly as in typical Robergia. The three pairs of lateral glabellar furrows, however, are like those of Robergia, yet the basal glabellar lobe is not subcireular and slightly inflated. The free cheek is relatively broad with a narrow border and doublure, base of the genal spine being opposite the occipital ring. The thoracic segments lack the large axial articulating processes and sockets typical of Remopleurides and have the inner part of the pleurae crossed by a broad shallow pleural furrow as in typi- cal Robergia. Axis of the pygidium includes only two pleural rings, the second divided by a median longitudinal furrow as in Remopleurides, and there is a post-axial ridge. Pleural regions 432 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY are flattened, bordered by two pairs of long, tapering spines directed backwards, a strong interpleural ridge separating the proximal parts of the pleurae. ROBERGIELLA SAGITTALIS nN. gen., n. sp. Plate 6, figures 16-33. TTolotype. USNM 137679 (PI. 6, figs. 16, 17), locality 14. Other Material, Paratype USNM 137680, all figured material in U. S. National Museum. Geological Horizon and Localities. Lower part of the Edinburg limestone, Liberty Hall facies, localities 14, 14a and 16. Description. Occipital ring longest in the midline, tapering distally to a blunt point at the axial furrow, the distal portion lying immediately behind the posterior part of the eye lobe. Median tubercle hes at the anterior margin. In front of the deep, narrow, occipital furrow, width of the glabella (PI. 6, figs. 16-18) about the same as that of the base of the tongue; imme- diately in front of this basal portion the glabella expands be- tween the palpebral rims. The tongue of length (sag.) less than half that of the eye lobe and curved only gently downward; narrow anterior area of fixed cheek. Three pairs of narrow, deep, lateral glabellar furrows, equally spaced from each other and from the occipital furrow, the anterior pair opposite the anterior part of the eye lobe; the furrows are diagonally directed, the first and second the longest, shghtly curved, the third short. Eye lobe moderately curved, convex palpebral rim defined by deep palpebral furrow; eye surface vertical with many tiny facets (Pl. 6, fig. 38), narrow convex external rim. Anterior branch of facial suture (PI. 6, fig. 26) has a forward and out- wardly directed course immediately in front of eye lobe; pos- terior branch of the suture (PI. 6, fig. 16) directed strongly outwards across the posterior border so that it reaches margin at a point about two-thirds of the width of the cheek (tr.). Free cheek (PI. 6, figs. 16, 17, 27) subtriangular in outline, greatest width along the line of the posterior border furrow, which is directed transversely. Anterolateral and lateral border narrow, convex, defined by a deep border furrow; at the genal angle the lateral border is continuous with the long slim genal spine. Posterior border slightly wider (exs.), small noteh in the WHITTINGTON : ORDOVICIAN TRILOBITES 433 margin immediately inside the base of the librigenal spine. Doub- lure (PI. 6, fig. 23) rolled beneath the border and not extending in beyond the lateral and posterior border furrows. Three incomplete thoracic segments are known, one of which is attached to the holotype (PI. 6, figs. 16, 17, 19, 24). Axial ring convex, inner part of pleura horizontal, transversely directed, and erossed by a deep diagonal pleural furrow; adjacent to the axial furrow the pleural furrow is subdivided by a low swelling, subtriangular in shape, which extends a short distance distally. Outer part of pleura curved, flattened, and pointed, the pleural furrow extending onto this portion and becoming shallower dis- tally. One segment with long median axial spine. Pygidium (PI. 6, figs. 29-32) with convex axis tapering rapidly baekwards, extending to about two-thirds the length (sag.), post- axial ridge becoming narrower posteriorly but reaching the margin. Axis with deep articulating furrow and subdivided into two rines by the shallow axial furrow; posterior of the two rings subdivided into two subecireular lobes by a median longi- tudinal furrow. Pleural regions horizontal, divided by an inter- pleural ridge which runs backward and slightly outward from the ring furrow, and by the postaxial ridge. Shallow pleural furrow curves outward and backward on the anterior segment, and adjacent to the axial furrow is subdivided by the low swell- ing. Each pleura continued by a backwardly directed gently tapering spine. Doublure broad, extending inward to tip of axis. External surface bearing fine, raised, anastomosing lines, these lines running transversely on the occipital and axial rings, in curves convex forward on the outer parts of the pleurae of thorax and pygidium. On the free cheek between the eye lobe and the border furrows the raised lines form a reticulate pattern (PI. 6, fig. 27). A row of small tubercles along the posterior margin of the oecipital and axial rings. On the median axial spine of the thorax and the doublure, raised lines run longitudinally, sub- parallel to the margins. Discussion. This species is not common at any of the localities at which it occurs, and the hypostome is not certainly known. A few small cranidia occur at locality 19 (Pl. 6, figs. 20-22) which may belong to this species. The example figured, of length (sag.) 2.13 mm., shows the three pairs of glabellar furrows (the anterior pair faint), and the palpebral rim is relatively wider than in 434 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY larger individuals. This specimen is referred to below in con- nection with the indeterminable small remopleuridid. REMOPLEURIDID gen. et sp. ind. Plate 19, figures 1-10. Material. Figured specimens, USNM 137689a-f. Geological Horizon and Localities. Lower Edinburg limestone, localities 3 and 4. Description. This cephalon oceurs with Remopleurides caela- tus, R. caphyroides, R. eximius, and R. simulus, and the largest eranidium known (Pl. 19, fig. 9) is only of length (sag.) 1.84 mm. It most closely resembles that of R. caelatus but differs from those of comparable size of this species (and of the other species mentioned). From R. caelatus (Pl. 3, fig. 16; Pl. 4, fig. 14) it is distinguished by the wider (especially posterolaterally ) palpebral rim and the wider (tr.), more rapidly expanding and less steeply sloping glabellar tongue. The anterior area of the fixed cheek and preglabellar area are wider, and the outline of the anterior margin is bluntly pointed rather than rounded. The free cheek (Pl. 19, figs. 6, 10) may be distinguished most readily by the subangulate outline of the anterolateral margin, and by the sharp edge along this margin, formed by the acutely angulate junction between the sloping dorsal surface of the free cheek and the doublure. The posterior margin of the free cheek has a less prominent projection and makes a larger angle with the inner margin of the librigenal spine. The characteristic pit is present in the outer surface of the anterior part of the doub- lure. Successively smaller cranidia [sagittal length respectively 126imms) (Pl, W9shfie.8), 91203) mm) (Pl osihies 5:47) | iditter principally from that described in the presence of the crescentic area of fixed cheek between the axial furrow and the palpebral furrow. Free cheeks associated with these cranidia are similar to those of larger size, and show the pit in the doublure. The smallest cranidium placed here (PI. 19, figs. 1, 2), probably that of degree 0, is of length (sag.) 0.85 mm. It differs from cranidia of degree 0 of R. caelatus (Pl. 3, fig. 9) in that the anterior cephalic spines are longer and slimmer, not curved, and in that short, bluntly pointed fixigenal spines are present. A few exam- ples of eranidia of the same size (Pl. 19, figs. 3, 4), lack the WHITTINGTON : ORDOVICIAN TRILOBITES 435 anterior cephalic spine, have the anterolateral margin smoothly eurved, but are otherwise indistinguishable. Evidently the an- terior cephalic spine is lost abruptly between moults. Hypostome, thoracic segments and pygidium unknown. Discussion. Inspection of the figures will show that the original of Plate 6, figures 20-22, of length (sag.) 2.13 mm., the smallest cranidium referred to Robergiella sagittalis n. gen., n. sp., might be considered to be a larger growth stage of the original of Plate 19, figure 9. The latter is the largest cranidium, length (sag.) 1.84 mm., placed in ‘‘remopleuridid gen. et sp. ind.’’ Thus the series placed here might be the early developmental stages of Robergiella sagittalis. However, these two groups of specimens do not occur together at any one locality, though both come from the lower part of the Edinburg formation. Without more com- plete developmental series of both groups a decision as to whether or not they belong to two separate species cannot be made. Should they prove to be a single species, then the early develop- ment of the cephalon (remopleuridid gen. et sp. ind.) is more like that of Remopleurides than that of Robergia (P1. 18, figs. 16, 205211). Distribution and Relationships of Remopleuridid Species from Virginia Remopleurides appears first in the silicified material in the Upper Lincolnshire limestone, where it is represented by a single species, R. asperulus. In the Lower Edinburg limestone a great variety of remopleuridid species appear suddenly. There seems little doubt but that Remopleurides caclatus (the most abun- dant), R. caphyroides, and R. eximius are distinct species. As noted above, the specimens included under R. simulus may be sexual dimorphs of R. eximius or they may be a further dis- tinct species from Lower Edinburg limestone. R. eximius is so like the species from the Upper Lincolnshire that it is presumed to be its descendant. The antecedents of the other species are not known. In addition, in the Lower Edinburg limestone, Robergia major and Robergiella sagittalis are present. The abundance of types of remopleuridids in the Lower Edinburg limestone may be ascribed partly to their having come from a considerable variety 436 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY of localities, but at all these localities the trilobite fauna is richer and more varied than at higher levels in the Edinburg or in the formations immediately below or above. In the Oranda limestone, remopleuridids are not known, but in the succeeding lower part of the Martinsburg limestone one species, Remopleurides plaesiourus, is quite abundant. This species is exceedingly similar to R. caelatus and is most probably descended from it. Comparison of the Virginia Remopleuridid Species with Those of other Areas The morphological characters displayed by the Virginia spe- cies are seen in slightly different combinations in species from northwestern Europe across into China (Lu, 1957, pl. 153, figs. 14, 15,16). Raymond (1925, p. 57) referred some material from Virginia to a Canadian species, and also erected the species R. rugicostatus on material from the Wilderness stage (Cooper, 1956) of Tennessee. This species is extremely like the slightly older R. simulus n.sp. (see above). The type species of Remopleurides, R. colbvi, from the Middle Ordovician of Eire (Whittington, 1950b, pp. 540-543, pl. 70, figs. 1, 2, 4, 5), has a cephalon (including hypostome) quite like those of the present R. caphyroides. However, the thorax and pygidium of R. colbii recall those of R. eximius (Pl. 14) and R. simulus (Pl. 17). There is a remarkable similarity between one specimen that I referred doubtfully to R. colbu (Whittington 1950b, pl. 69, figs. 5, 6), and the holotype of R. eximius. A second species from the same horizon in Hire is R. dorsospinifer (Whit- tington, 1950b, pl. 69, figs. 7-10), a species that has much in common with R. caelatus (Pls. 1, 2) and R. plaesiourus (PI. 5, figs. 1-15). In 1950 (p. 542) I held the view that R. dorsospinifer might belong to a genus separate from Remopleurides. I would now expand the limits of Remopleurides and include R. dorso- spinifer within this genus. A third species from Hire, R. longi- costatus (Whittington, 1950b, pl. 70, figs. 3, 6), has the long genal spines (PI. 7, fig. 4) and the long bladelike pleurae (PI. 9, figs. 1-5, 7,8) characteristic of R. caphyroides. From Middle Ordovician horizons in Scotland come Remo- pleurides girvanensis Reed, (1903, pp. 39-41, pl. 6, figs. 8-15) and WHITTINGTON : ORDOVICIAN TRILOBITES 437 R. biaculeatus Tripp (1954, pp. 664-666, pl. 2, figs. 1-12). The former of these species is ike R. eximius from Virginia. R. biacu- leatus bears a strong resemblance to R. caelatus, and Tripp’s description includes such details as the pit in the cephalic doub- lure and the median boss and oval areas of the hypostome. Thors- lund (1940, pl. 7) illustrated remopleuridids from the Middle Ordovician of Sweden. The cephalon of R. validus Thorslund is like that of R. caelatus, whereas the pygidium of this species is more like that of R. eximius; the pygidium of R. ef. latws Olin is more like that of R. caelatus. The thorax of R. nanus elongatus (Opik, 1926, pl. 2, fig. 17), from the Middle Ordovician of Es- tonia, is like that of R. stimulus, the pygidium (Opik, 1937, pl. 24, figs. 1, 2) of R. caelatus type. There is evidently a closely related group of species of Remopleurides in the areas mentioned. Quite similar species of Robergia are known in Oklahoma and the Appalachians (Cooper, 1953, pp. 22-24), Scotland and Sweden (Whittington, 1950b, pp. 543-4), in Middle Ordovician rocks, but do not seem to be known in Asia (I regard the illustrations referred to by Kobayashi, 1951, p. 20, as being of Remopleurides). Superfamily TRINUCLEOIDAE Hawle and Corda, 1847 Trinucleidae and Raphiophoridae exhibit characters in com- mon in both developmental and holaspid stages. In holaspides the occipital ring is narrow and lacks the doublure; in front of it the glabella expands forward and has the frontal part the most strongly convex. There are three pairs of subcircular or suboval muscle areas ; the basal area is in many genera stronely impressed in the external surface, these impressions defining a narrow ‘*neck’’ or a swollen ring, the occiput, in front of the occipital furrow; median pair is situated well up on the slope of the glabella, anterior pair small. Deep anterior pit. Cheek convex, posterior border defined by furrow which deepens distally and may form a pit; librigenal spine long; eye lobe bearing a com- pound eye absent, eye tubercle in some trinucleids; suture not passing across eye lobe but marginal or submarginal, free cheeks narrow (tr.) on dorsal surface, continuous ventrally, no median or connective suture. Hypostome subtrapezoidal in outline, con- vex middle body, narrow border defined anterolaterally and con- tinuous around lateral and posterior sides; small wings. Seem- ingly hypostome not joined to cephalic doublure alone a suture, but probably held closely beneath glabella by muscles. 438 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY Five or six thoracie segments, articulating furrow with a deep appendiferal pit, the horizontal pleurae bent down only at the extreme outer part, the doublure merely a curled-under edge, the pleural furrow strong. Articulating half ring lacking on first segment, which is bevelled anterolaterally. Pygidium sub- triangular in outline with a gently convex, tapering axis reach- ing back to the rim. The larger part of the pleural regions is horizontal, the outer part bent down to form a steeply sloping border, the margin of this border typically sinuous, with a median posterior notch; doublure formed of the curled-under edge. Some ten segmental divisions may be revealed in the axis by up to five ring furrows and appendifers; muscle areas (com- monly an inner and outer pair) may indicate further segments posteriorly; one to five pleural furrows. Apparently when enrolled, border of pygidium lay against inner edge of cephalic doublure. The protaspides, described here for the first time, are similar in outline, convexity, and breadth of the gently convex axis. Up to three pairs of border spines may be present; in the cephal- ic portion the suture runs around the outermost edge of the pleural regions and narrow preglabellar area, posteriorly cross- ing the doublure in a sharp curve. In larger protaspides the glabella has the clavate form, the axis of the protopygidium is bulbous and vaguely defined, the pleural regions slope vertically posteriorly and have the median notch. Earliest meraspid cran- idia display the clavate glabella with flat, steeply sloping sides and a median longitudinal keel, the occipital ring ill-defined. Convex cheek without border spines, posterior border best de- fined distally, by the pit. Suture is marginal and laterally has a slightly inwardly concave course. Alae, which are not evident in the protaspis, are relatively large and gently convex in trinu- cleids, Ampyx, Ampy.xina, and Raymondella, but not in Lon- chodomas. Small transitory pygidia have a transverse outline, a bulbous, ill-defined axis that does not taper posteriorly, the pleural regions slightly bent down beside the posterior part of this axis, the outermost parts of the pleural regions bent down to form the border. Typically there is a median posterior notch in the outhne. WHITTINGTON : ORDOVICIAN TRILOBITES 439 Between the protaspis and degree 0, border spines are lost while at degree 0 the frontal glabellar spine of raphiophorids makes its appearance. In immediately succeeding degrees the trinucleid fringe develops rapidly. Thus some familial (e.g. the raphiophorid frontal glabellar spine) and generic (e.g. the glabellar carina of Lonchodomas) characters are evident at de- grees 0 to 1, whereas other generic characters (e.g. the expan- sion of the frontal part of the glabella in Tretaspis and Raymon- della, or the fringe of Vretaspis) develop in immediately succeeding degrees. Family TRINUCLEIDAE Hawle and Corda, 1847 The common characters of holaspides of this family have recently been discussed (Whittington, 1959, p. 0420). The pres- sent material, however, shows the development more completely and, by virtue of the excellent preservation, in greater detail than heretofore. The protaspides of Cryptolithus (Pl. 28, figs. 11-16) and Tretaspis (Pl. 26, figs. 2, 3, 5, 6, 8-13) show a con- siderable likeness in form. Only degree 0 specimens were previously known, and they have no marginal spines on cheek lobe or transitory pygidium. The presence of such spines in the protaspis is thus unexpected, and their loss at the end of the protaspid period abrupt. The development of the glabella is similar in each genus — in the protaspis it has only a slight anter- ior expansion, this expanson increasing and a strongly convex, sharply carinate, form appearing early in the meraspid period in Cryptolithus (Pl. 23, figs. 1, 2, 9), in the late protaspid period in Tretaspis (Pl. 26, figs. 10, 13). Only in holaspides does this carinate form give place to a convexity that is rounded in trans- verse profile, while the anterior expansion becomes fully devel- oped. Alae are not discernible in the protaspis, are largest at degree 0 (Pl. 23, fig. 6; Pl. 27, fig. 1), and are progressively reduced so that little or no trace of them remains in the largest holaspides (Pl. 20, figs. 4, 6; Pl. 24, fig. 1). The border of the cephalon is at first a rolled margin, traversed on the upper surface by the marginal suture and at the genal angle continuous with the genal spine. A concentric row or rows of pits first appear (in late degree 0 and degree 1; Pl. 23, 440 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY figs. 5, 7; Pl. 27, figs. 18, 19) in a deep trough which is immedi- ately adjacent to the cheek lobes and preglabellar area, and there is a corresponding trough in the lower lamella. The first com- plete concentric row or rows has almost the same number of pits as in the holaspis. The fringe widens rapidly as additional con- centric rows are added, and not until three such rows are pres- ent is the girder apparent (PI. 22, figs. 1, 2, 5, 6, 10, 12; Pl. 28, figs. 1-4). The complete number of concentric rows appears fairly early in the meraspid development, and in the later meras- pid and early holaspid periods relatively few pits are added and the fringe assumes its characteristic holaspid form — for example (compare Pl. 22, figs. 1, 2, with Pl. 20, figs. 4, 6, 7; Pl. 28, figs. 1-8 with Pl. 24, figs. 1-5), the lower lamella becomes sharply flexed at the girder as the latter becomes deeper, cheek roll and brim appear in T'retaspis, and concentric ridges and radial ridges and sulci appear in the upper and lower lamellae. The abrupt appearance of a new, complete concentric row of pits in the early meraspid degrees is accompanied by a marked increase in dimensions. Thus the early meraspides of Cryptolithus show a series of size groups corresponding to those with one, two and three complete concentric rows. It is hoped that the abundant material of Cryptolithus will permit a more detailed study of this phenomenon. Harly transitory pygidia (Pl. 23, figs: 3, 8, 10; Pl. 27, figs: 14, 15, 17, 18, 21, 23) have a bulbous axis which is broad and rounded posteriorly, the posterolateral parts of the pleural regions are bent down steeply, and the shallow median posterior notch gives a bilobate outline. During the meraspid period (PI. 20, Hes: 17-19) 21) 23- Pl. 27%, fies: 20, 22°24. 25) thexouthine becomes triangular, the axis gradually assumes the tapering form, the rings are better defined, the pleural regions become horizontal, with the outer parts bent down in the characteristic vertical border; in posterior view the outer margin of this border is sinuous and has a median notch. While these parallels can be drawn between the development of Tretaspis and Cryptolithus, it is nevertheless evident that from the protaspis onwards the two genera can readily be dis- tinguished. In the early meraspid degrees, for example, Tretas- pis already exhibits such features as the relatively wider and WHITTINGTON : ORDOVICIAN TRILOBITES 44] more convex frontal part of the glabella, the more prominent eye tubercles and eye ridges, and the wider lateral part of the fringe with more _ pits. Subfamily CRYPTOLITHINAE 0;, LG: Description. Reconstruction of exoskeleton (Text-fig. 7) shows outline and convexity ; length (sag.) of cephalon almost half that of exoskeleton. Glabella (Pl. 29, figs. 1-4) narrowest across occipital ring; this ring is sharply ridged, with a vertical poster- ior slope and a steep slope down to occipital furrow, and not A66 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY separated by a depression at the axial furrow from the posterior border. Posterior, ventral edge of occipital ring thickened, but doublure absent. Occipital furrow broad and shallow, in dorsal aspect both occipital furrow and ring curved convexly backward. Glabella, defined by shallow axial and preglabellar furrows, ex- pands forward to a maximum width at about one-third the length, anterior part rounded in dorsal outline, projecting beyond Figure 7. Ampya virginiensis Cooper, 1953. Reconstruction, A, dorsal, B, left lateral views. Approximately X 2. cephalic margin, and drawn out into long, curved spine, the tip of which is directed upward and slightly backward; glabellar spine round in section, longitudinal lateral furrow in proximal portion; behind base of spine glabella rounded (not carinate) along midline, sides flattened and steeply sloping. Four pairs WHITTINGTON: ORDOVICIAN TRILOBITES 467 of muscle areas (visible by their colour, the thinness of the exo- skeleton over them, or by being impressed): occipital area elongate-oval, situated in outer part of occipital furrow, adjacent to it is subecireular first lateral sear, the anterior, inner and posterior boundaries of which are impressed into glabella, and the outer, posterior corner connected by a shallow depression to the occipital area; second lateral area larger, subcircular in outline, situated immediately in front of first and half way up slope of elabella, not impressed; third lateral area smaller, subcircular in outline, adjacent to axial furrow and at greatest width of elabella. In axial furrow, adjacent to posterior border furrow, there is a narrow (tr.), short (exs.) depression, probably a muscle area; anterior pit deep, elongate, situated in front of third lateral muscle area. Cheek (PI. 29, figs. 5-8, 11) gently convex, lateral and anterior border narrow, flattened, defined by change in slope rather than a furrow; border flexed down at margin and rolled under to form a doublure which is no wider than the border; anterior border and preglabellar furrows eoincident. Posterior border a sharp ridge, anteriorly sloping to broad shallow posterior border fur- row, latter with deep pit distally; distal part of border bent down; on posterior edge, just inside suture, is a socket to receive anterolateral corner of first thoracic segment. Facial suture euts posterior margin near extremity of border, from here runs outward and slightly forward, curves abruptly back over poster- ior border furrow outside distal pit, follows a gently sinuous eourse across outer part of cheek, and two branches meet in smooth curve along anterior margin of border. No median or connective sutures. Librigenal spine long and gently curved, hase formed by extensions of lateral and posterior cephalic borders; posterior border furrow of cephalon is continuous with shallow longitudinal furrow on upper surface of librigenal spine, similar furrow on lower surface of spine so that, proximally, eross-section is hour-glass shaped; furrows die out distally; an elongate depression on inner side of spine near base. Antero- lateral and anterior parts of cephalic doublure divided by a flexure into an outer, wider, flat part and an inner, gently convex part ; flexure has a slightly sinuous course; inner edge of doublure thick, rounded, traversed by fine raised lines. 468 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY Hypostome (PI. 29, figs. 9, 10, 12-16) shield-shaped, anterior margin gently curved convexly forward, medially no anterior border; distally a gently convex border becomes defined and is continuous with lateral and posterior borders; shallow border furrow. Middle body most convex in median part, gently sloping anterolaterally ; thinner area in anterolateral corner adjacent to border, probably a muscle area; shallow pit in border furrow posterolaterally. In lateral and posterior views (PI. 29, figs. 14, 15) border is seen to be deepest (i.e. extended dorsally but not eurled inward) laterally and posteriorly, and outline shows a lateral and median posterior scallop, with a long, pointed postero- lateral projection (the posterior wing) between the scallops. Thorax (Pl. 30, figs. 1-13) of six segments, first the longest (sag. and exs.), succeeding segments progressively shorter, and, after the second, narrower (tr.); axis gently convex, one-third of total width and tapering slightly. Subcircular distal part of axial ring gently inflated, posterior edge of ring with slight median indentation ; articulating furrow broad and shallow, run- ning in curve concave forward, articulating half-ring and doub- lure of ring short (sag.) ; oval area in outer part of articulating furrow is shghtly more depressed than rest of furrow and may be different in colour, and is probably a muscle attachment area. Small axial articulating processes and sockets, absent only be- tween first thoracic segment and occipital ring. Inner part of pleura horizontal, outer part of pleura short (tr.), bent down steeply ; pleural furrow broad, shallow, directed slightly diagon- ally, deepening and curving forward as it reaches fulerum and ends against facet ; in dorsal aspect line of fulerum is inward and forward on first segment, on succeeding segments progressively more strongly inwardly and backwardly directed, giving charac- teristic outline to thorax; in addition outer part of pleura is longer (tr.) on succeeding segments (compare PI. 30, fig. 6 with Pl. 30, fig. 8). Fuleral articulating process and socket on each segment, including process on first segment which fits into socket in posterior border of cephalon where latter is flexed downward. Doublure of pleura short (tr.), curled under outermost part of pleura; each pleura with small facet on outer part, the tip of the facet being received during enrollment by a depression in the doublure of the preceding segment, this mechanism providing a limit to amount of enrollment. WHITTINGTON : ORDOVICIAN TRILOBITES 469 Pygidium (PI. 30, figs. 14, 16, 17, 20) three times as wide as long, gently convex axis tapering back to reach inner margin of border. Articulating furrow with deeper outer part, short (sag.)} articulating half-ring. Eight or nine axial rings indicated by furrows which are deepest medially and become progressively fainter posteriorly. In outer part of each furrow an oval muscle area, corresponding in position with deeper part of articulating furrows of pygidium and thorax. Inner part of pleural region horizontal, crossed by broad pleural furrow running in curve concave forward; area in front of this furrow bent to slope up- ward and forward, most strongly bent distally adjacent to articulating process and inner part of facet. Border slopes steeply and is broad, edge curled under and in posterior view with a characteristic bow-shaped outline imposed by the greatest width being posterolateral and by the median posterior notch. External surface smooth (or extremely finely granulate) but not ‘‘punctate’’ as stated by Cooper (1953, p. 16), fine granu- lation on glabellar spine (PI. 29, figs. 1, 3) and on proximal part of lbrigenal spine, fine raised lines on outer part of cephalic border, border of hypostome, and outer part of pleural regions of thorax and pygidium, these lines running subparallel to the margins. Discussion. Ampyzx virginiensis is extremely like the type spec- ies A. nasutus (Whittington, 1950b, pp. 554-556, pl. 74, figs. 3-9, text-fig. 6) from the Lower Ordovician of Sweden, differing only in such features as the less forwardly projecting glabella, nar- rower anterior and lateral cephalic border and relatively shorter pygidium. Cooper (19538, pp. 16-17, pl. 7, figs. 1-11, 18) erected this species for material from the present localities 2and 6. The relatively longer (sag.) pygidium, with deeper axial fur- rows, of Ampyx americanus (Cooper, 1953, pl. 5, figs. 3-5, 8,9 [figure 3 is X2, not X5 as stated by Cooper]), from a similar horizon in Tennessee, serves to distinguish it from A. virginiensis. A. camurus (Cooper, 1953, p. 16, pl. 5, figs. 1, 2, 6, 7; this paper, Pl. 30, figs. 15, 18, 19), is distinguished from A. virginiensis by the more convex frontal portion of the glabella and the relatively longer pygidium which has a more triangular outline. Both species occur at about the same horizon in Virginia, but A. camu- rus has not been found among the silicified material. From the 470 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY same locality and horizon that vielded A. camurus came also A. lobatus Cooper, 1953 (p. 16, pl. 6, figs. 3, 4), based on a distinctive pygidium that exhibits the second and third pleural furrows extending across the inner pleural regions. Comparable furrows are not present on pygidia of A. virginicnsis of the same size. I have not seen the type material of Ampyxr? hastatus Ruedemann (1901, pp. 48-54, pl. 3, figs. 1-10, 30), but it appears to differ from A. wirginiensis in the presence of the carina on the glabella, the prismatic section of the frontal glabellar spine, and outline of the pygidium. A. virginiensis has one, rather than two, anterior glabellar muscle sears. Whittard (1955, p. 19, pl. 2, fig. 8) has described the hypostome of Ampysx linleyensis. That of A. virginiensis is similar, but wider (tr.) across the anterior wings, the position of which is revealed by the muscle area. Whittard interprets the projection bounding the anterior margin of the notch in the lateral border as the anterior wing in A. linleyensis, but by analogy with A. virgimensis the anterior wing may be in advance of this position. Development Protaspis. The protaspis here attributed to A. virginiensis oc- eurs at localities 3 and 4, where small developmental stages of the species are quite abundant. This protaspis does not seem to be that of Lonchodomas carinatus since small developmental stages of this species are rare at locality 3, and do not occur at locality 4. Smallest specimen (P]. 30, figs. 26, 30) of length (sag.) 0.47 mm., maximum width 0.49 mm. Greatest width at about half the length, in front of which straight lateral margins converge slightly forward to the obliquely angulate anterolateral margin: anterior margin curved gently forward; behind greatest width shield narrows more rapidly and is rounded. Axis of width about one-third that of entire exoskeleton, subparallel sided and rounded at each end. Pleural regions curved down more steeply posterolaterally, almost vertical behind the axis; at anterolateral angle a long slim spine, a second spine on the posterior margin in line with the axial furrow; between posterior pair of spines mar- gin is notched. Doublure narrow, extending forward to about the maximum width; in front of this point margin of the exo- skeleton is sutural. Free cheeks unknown. WHITTINGTON : ORDOVICIAN TRILOBITES 471 A slightly larger example (PI. 30, figs. 21-24, 27; length 0.53 mm.) is similar in form but has the axis divided by a ring furrow ; glabellar portion more strongly convex at the rounded anterior end, protopygidial portion more convex and slightly wider than the basal part of the glabellar portion. Posterior margin of protopygidium deeply notched between bases of spines, doublure of notch bearing a short, ventrally and inwardly directed spine on each side. Doublure extends forward only a short distance in front of posterior spine before being cut off by suture — appar- ently the suture in this specimen extends farther back than in the smaller one, and therefore anterior margin of pleural region of protopygidium must run outward and backward from base of elabella. Free cheek unknown. The largest example (PI. 30, figs. 25, 28, 29) is incomplete and apparently has the border spines broken off; it is similar in form but the glabella has a definitely clavate outline and is narrower across the basal part than across the expanded, convex, anterior part. Faint transverse furrow extends across pleural regions to divide cephalie from protopygidial portion ; thus anterior margin of protopygidium is transverse and pleural region is longer (exs. ) and wider. Degree 0. Ampyx virginiensis 1s rare at locality 16, yet one specimen (Pl. 31, figs. 1, 2) is a degree 0 raphiophorid, and seemingly belongs to this species. Glabella now outlined by shallow axial furrows, sloping steeply posteriorly to the narrow (sag. and exs.), faintly defined occipital ring, anteriorly pro- jecting forward over the cephalic margin (the outline of which is concave medially) and continued by a thick, forwardly directed glabellar spine; shallow depression in side of glabella immediately in front of occipital ring. Fixed cheek gently convex, sloping steeply anterolaterally, bent down along posterior margin but the posterior border not defined. Fixed cheeks united in front of glabella by narrow (sag.) preglabellar area. Fixed cheeks and preglabellar area bounded by facial suture, which on the left side has the characteristic slight inward curvature, and continues around the posterolateral corner. Free cheeks and hypostome un- known. Transitory pygidium more than twice as wide as long, eurved down (sag. and exs.) so that posterior part slopes verti- cally; axis stands above pleural regions, divided by shallow ring furrows into two gently convex rings and a more convex 472, BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY terminal portion; pleural region flat in a transverse direction, narrow raised rim, narrow border curled under as a narrow doublure. Cephalon. Smallest cranidium (PI. 31, figs. 3-5; length exclud- ing glabellar spine 0.8 mm., maximum width 0.69 mm.; length glabellar spine 0.49 mm.) subsemicireular in outline, ovoid glabel- la with rounded median ridge and steeply sloping sides, deep, round pit immediately in front of distal part of low occipital ring, glabellar spine long, round in section, curved slightly down in lateral view. Ala occupies inner one-third of area of cheek, is gently convex, outlined by shallow alar and axial furrows. Posterior border faintly indicated distally by depression. Facial suture curves around posterolateral corner of fixed cheek and course is faintly indented in lateral margin. Next largest cranidium (Pl. 31, figs. 7-9; length excluding glabellar spine 0.45 mm., maximum width 0.94 mm.) has antero- median part of glabella higher and more convex so that it projects farther forward, the glabellar spine is straight, ala is more distinctly outlined, and posterior border furrow is com- plete, ending distally in a pit. Free cheek shows the lateral and anterior borders and border furrows, is relatively narrow (tr.) compared to larger specimens (apparently because of the lesser inward curvature of the suture laterally), and has a backwardly curving librigenal spine. The doublure is narrow. In cephala up to a length (sag., excluding glabellar spine) of 1.5 mm. (PI. 31, figs. 6, 10-20) the glabella becomes slightly more convex and the frontal spine upeurved. The depression in front of the occipital ring becomes recognizable as the basal muscle area, and in cephala of length 1.42 mm. or more the second and the anterior areas are visible (PI. 31, fig. 15). The ala persists as a convex area in the inner corner of the cheek, connected by a low ridge to the extremity of the occipital ring, but becomes progressively smaller. The facial suture curves more strongly inward across the lateral part and the cheek, and the free cheek becomes wider (tr.); the proximal part of the librigenal spine becomes straight and outwardly and backwardly directed. Gran- ulation, present on the external surfaces of cephala of length (sag.) 0.52 mm. (PI. 31, figs. 6, 10), where it is coarsest on the free cheek, becomes finer. Beyond a length (sag.) of 14 WHITTINGTON : ORDOVICIAN TRILOBITES 473 mm. there is little change in the cephalon, except that the ala disappears; a trace of it in the form of a low ridge connecting the inner corner of the cheek to the extremity of the axial ring is visible in cephala of length 3 mm. and slightly larger. The smallest known hypostomes do not differ greatly from the largest. A size series of transitory and true pygidia, believed to be that of Ampyx virginiensis, is shown in Plate 31, figures 21-32. The corresponding series of Lonchodomas carinatus (Pl. 33, figs. 7, 10, 13, 15, 18, 20, 21, 25) is quite similar, especially at the smaller sizes, and such transitory pygidia of these two species are not readily separated. Pygidia of length (sag.) 0.69 mm. (PI. 31, figs. 31, 32) of A. virginiensis differ from larger examples (Pl. 30, figs. 14, 16, 17, 20) in the presence of the second, third, and faint fourth pleural furrows (each one directed more strongly backward), in showing faint interpleural grooves, and in possessing the rim. Following the series through progressively smaller sizes, the convex axial rings become more clearly defined by ring furrows having a distal pit, the outline becomes more rounded, the border narrower and the rim stronger. In the two smallest specimens (PI. 31, figs. 21-24) a posterior indentation appears, and in the smallest of these only two axial rings and a bulbous terminal portion are visible, and the pleural regions are bent down posteriorly. The characteristic bow-shaped outline of the external margin of the border, seen in posterior view, is present throughout the series. This smallest transitory pygi- dium (PI. 31, figs. 21, 22) is larger than that of the degree 0 specimen (PI. 31, figs. 1, 2), and differs in exhibiting pleural furrows and a more acute angle between the outline of the anter- ior and lateral margins. Genus LONCHODOMAS Angelin, 1854 LONCHODOMAS CARINATUS Cooper, 1953 Plates 32, 33; Text-figure 8 Figured Material. Allin USNM Geological Horizon and Localities. Lower Edinburg limestone, localities 2, 3, 5, 6, and Willow Grove, 3 miles south of Woodstock, Shenandoah Co., Virginia. Description. Cephalon (Pl. 32, figs. 1, 4, 5, 7) excluding spines triangular in outline, about twice as wide as long. Glabella 474 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY subdiamond-shaped in outline; occipital ring a narrow (sag.) band, not separated by axial furrow from posterior border and not raised above the adjacent part of the border, posterior mar- gin gently arcuate backward ; occipital furrow extremely shallow ; in front of this furrow glabella becomes moderately convex, with a median longitudinal carina from which the gently convex sides slope steeply down, maximum width in front of the mid-leneth, anterior part drawn out into a long, tapering, frontal glabellar spine which is directed horizontally with an extremely slight upward curvature, the spine square in cross section, each corner of the square projecting slightly to form a longitudinal ridge, the two ridges on the upper surface running back to join and merge into the carina, the two ridges on the lower surface being continuous with the anterolateral edge of the cephalon; muscle areas on the glabella (darker areas, smooth externally — Pl. 32, fig. 23; Pl. 33, figs. 31, 32; Text-fig. 8a) include a small oval area in the distal part of the occipital furrow, in front of this a sub- triangular area, the posterior part of which extends up the side of the glabella quite close to the carina, confluent with this a third elongate-oval area disposed longitudinally on the glabella about half way up the slope; a fourth poorly defined area is situated on the slope of the glabella just inside the anterior pit. The anterior area was not observed in Lonchodomas rostratus (Whit- tington, 1950b, fig. 7) but may be seen in Lonchodomas sp. of Stormer (1949, p. 178, fig. 14b). There seems no evidence for the subdivision of the basal and median areas shown by Hupé (1953, p. 72, fig. 26, 3). Axial furrows not distinct, limits of glabella ill-defined by change in slope and by the deep elongate anterior pit which is situated opposite the maximum width of the glabella. Cheek subtriangular in outline, sloping anterolaterally, inner part gently convex, outer part sloping less steeply towards the acutely angulate margin (PI. 32, fig. 1) so that in profile there is a gentle concavity; posterior border sloping steeply forward, posterior border furrow commencing a short distance outside the occipital ring and ending just inside the facial suture in a deep pit; at about half its width the posterior border is bent down to slope gently outward; librigenal spine thick, hook-shaped, end- ing in a sharp point, prismatic in cross section, each edge raised to form a strong ridge, the surfaces between the ridges concave. wo t~ ORDOVICIAN TRILOBITES WHITTINGTON ‘aumojsodxy Jo Lpoq apppru Jo oqo] sO1ojsod “d tawojsodxy JO Jopaoqg 1otiaysod ‘qd Soanjpqnop atyeydos JO ULsIvU JOUUT UT YoJou fu FBiNoBUT ‘UW ‘a famoysodady Fo Lpoq oppptuat JO oqo] Lorteyue ‘7p -W0A ‘) ‘[B10} Bl JUS ‘gq ‘Tesiop fy ‘SMOTJONAYSMOVY 4] foainjqnop oipeydead jo yavd Lou poyvAvslo JO aspo ISUOTJVIABIQQCY °“%so X Apopyeurxoiddy “sMota [v.14 "EGET ‘tad00D snyvUiDO SDUOPOYOUOT *8 OINSTT 476 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY The lower, outer ridge of the spine continuous with the acutely angulate anterolateral border of the cheek, the lower, inner ridge dying out as it approaches the genal angle, the upper, outer ridge dying out over the genal angle of the cheek, the upper, inner ridge continuous with the sharp edge of the posterior border. The facial suture crosses the posterior margin at about three- quarters the width of the posterior border, runs at first outward and slightly forward, then in an ‘S’ eurve across the posterior border and the outer part of the cheek, reaching the antero- lateral margin almost directly in line (exs.) with the point at which it crossed the posterior margin; from here the suture runs along the acutely angulate margin of the cephalon toward the midline, the two branches meeting on the under side of the frontal glabellar spine in a double curve with a median sinus (Pl. 82, figs. 2, 3, 5; Text-fig. 8c) ; these curves give a character- istic outline to the anterior margin of the doublure. Doublure broadest anteriorly where it is gently convex ventrally, antero- laterally about as wide as the librigenal spine; outline of the inner margin a smooth semicircular curve, interrupted by shallow median anterior notch (n in Text-fig. 8¢); a short dis- tance inside this margin is a flexure (e in Text-fig. 8c) so situated that the narrow inner part of the doublure projects slightly ventrally below the wider, flat, outer part (compare PI. 32, figs. 2, 3). Hypostome (PL 32ietigs: 6.08% 9) 12) 17 = Textsie W8e) Psub- hexagonal in outline, in Senin aspect flexed along ait ‘midline so that the sides slope steeply from the rounded median part. Anterior lobe of middle body subcireular in outline, reaching anterior margin in the midline, anterolaterally curving down to flattened border; near base of small triangular anterior wing this border passes into the gently convex lateral border; lateral border furrow deep just behind anterior wing, becoming shal- lower posteriorly and running into middle furrow; posterior border a narrow convex band, bearing raised lines, margin gently curved convexly posteriorly; macula a small, slightly raised oval area situated midway along middle furrow; posterior lobe of middle body ecrescentic in outline, flattened. Margin of borders bent down but not curled under to form a doublure; posterior wing a small rounded projection from this bent-down i WHITTINGTON : ORDOVICIAN TRILOBITES 47 margin at the posterolateral angle, margin excavated between the wings. The outline and curvature of the transverse median part of the anterior edge of the hypostome is such that it will not fit exactly against the inner edge of the cephalic doublure (compare figs. 6, 9 with figs. 2, 3, Pl. 32). Further, these edges are not thick and sharply transected as are, for example, the edges of the eheek along the dorsal facial suture. This suggests that the anterior edge of the hypostome did not lie against the inner edge of the cephalic doublure along a hypostomal suture. The hypo- stome may have been attached only by musculature to the dorsal cephalic exoskeleton. If the exoskeleton is reconstructed with the tip of the anterior hypostomal wing lying close to the anterior boss, then the anterior part of the hypostome lay both above and in front of the cephalic doublure, as suggested in Text-figure 8c. External surface of occipital ring and median lobe of glabella (excluding the muscle areas) and posterior part of the cheek bearing closely and irregularly spaced small pits (Pl. 32, fig. 23); similar pitting may be visible on the middle body of the hyvpostome. Anastomosing raised lines run longitudinally on the rounded angles of the frontal glabellar spine, the hbrigenal spine, and the anterolateral edge of the cephalon; similar lines on the outer surface of the inner, ventrally projecting part of the cephalic doublure, and on the lateral and posterior borders of the hypostome. The exoskeleton is thin and differently col- ored over the hypostomal maculae. Number of thoracic segments unknown, but presumed in the reconstruction (Text-fig. 8a, b) to be five as in other species of this genus. A few isolated segments have been obtained (PI. 32, figs. 10, 11, 13-15). Axis about one-third of the width, gently convex, ring flattened and with slight convexity distally; articulating furrow narrow and deep, articulating half ring short (sag.), as is the doublure of the axia! ring. Inner part of pleura horizontal, at fulerum bent down vertically, outer part short, with a narrow doublure rolled closely underneath. Pleural furrow narrow and deep, commencing a short distance out from the axial furrow. Axial and fuleral articulating processes pres- ent, a small facet on the anterolateral area of the outer part of the pleura, and a small depression in the posterolateral part of the doublure. These segments are extremely like those of Ampyw virginiensis (Pl. 30, figs. 6-13) but may readily be separated by 478 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY the flatness of the axial ring, narrowness of the articulating and pleural furrows, ete. These segments also match, and fit with, the posterior border of the cephalon and the pygidium of Lonche- domas carinatus. Pygidium (PI. 32, figs. 18, 19, 21, 22) three times as wide as long, anterior margin straight, transverse, posterolateral mar- vins rounded; gently convex axis tapering and extending back to inner margin of border, inner part of pleural regions hori- zontal, outer parts bent down to form a steeply sloping border, this border projecting in a gentle convexity behind the axis. Deep articulating furrow and short (sag.) articulating half ring, first axial ring indicated by its slight convexity; behind this ring axis subdivided only by the paired muscle areas; there are two pairs of muscle areas in each ring furrow, a larger outer transversely ovate area, and a smaller inner area, the inner pair being joined across the midline by a narrow band; up to nine ring furrows are indicated by these pairs of muscle areas. Pleural furrows narrow, deep, with a forwardly concave outline ; first pleural furrow visible on all specimens, commencing a short distance outside the axis and extending to the inner margin of the border; on some specimens the second, third, and fourth pleural furrow may be seen, each furrow directed outwards and slightly more strongly backwards. Small facet. Outer edge of border thickened, rounded, doublure not extending beneath border; in posterior view the outline of the border having a deep median notch. Axis of pygidium and adjacent part of pleural regions pitted in some specimens, fine, anastomosing raised lines on the outer parts of the thoracic pleurae and the border of the pygidium. Discussion. Lonchodomas carinatus was described by Cooper (1953, pp. 17-18, pl. 7, figs. 12-17, 19-23), the holotype coming from the present locality 2. Material from the other localities mentioned here appears to be identical with that from locality 2. The cranidium, thorax and pygidium of L. carinatus are ex- tremely like those of the type species L. rostratus (Whittington, 1950b, pp. 556-557, pl. 74, figs. 11-15, text-fig. 7), which comes from beds in Norway of the same age as the lower Edinburg formation. DL. carinatus cannot be compared with L. sublaevis Raymond (1925, pp. 40-41, pl. 2, figs. 11, 12,) because the type material is too poor for any comparison, nor can it be compared WHITTINGTON : ORDOVICIAN TRILOBITES A479 with L. punctatus Cooper (1953, p. 18, pl. 4, figs. 20-21) because only the pygidium of this species has been described in detail. The type material of L. politus Raymond (1925, pp. 39-40, pl. 2, figs. 8-10), consisting of two eranidia and a pygidium, cannot be distinguished from specimens of the same size in the present material. However, the free cheek of ZL. politus is not known, hence it is uncertain whether or not the librigenal spine had the hooked form characteristic of L. carinatus. I hesitate to regard L. carinatus as a synonym of L. politus without this information. Cranidia and pygidia from Tennessee identified by Cooper (1953, Pl. 6, figs. 1, 2, 5-8, 10) as L. politus differ from L. carinatus in that the carina is absent on the anterior part of the glabella, there are no pits in the external surface of the fixed cheek, and the pleural regions of the pygidium slope in toward the axial furrows and the distal parts of the external surface exhibit more pleural furrows. In species of Lonchodomas from Scotland and Oklahoma (Whittington, 1950b, p. 557) the librigenal spine is long and gently curved, rather like the corresponding spine in Ampyz. Through the kindness of Professor Marshall Kay I have examined material attributed to Lonchodomas halli (Billings, 1861) from the Youngman formation, Highgate Springs, Vermont (Kay, 1958, p. 83). Cranidia and pygidia of this Middle Ordovician species are associated with free cheeks bearing a librigenal spine having much the same form as that of L. carinatus. An associated hypostome is also like that of the Virginia species. Hvidentiy L. halli from Highgate Springs, Vt. (one of Billings’ type localities) is a species closely related to L. carinatus, and these two species are the only ones so far known with these peculiar hooked hbrigenal spines. According to Cooper (1956, pp. 31-52, Chart 1) the Youngman is of the same age as the early Edinburg limestone. Development One enrolled degree 1 exoskeleton (PI. 33, figs. 1-4, 7), laek- ing free cheeks, and isolated cranidia of similar size (Pl. 33, figs. 5, 6, 8) are known. Length of glabella of latter (excluding frontal spine) 0.65 mm.; width of cranidium 1.35 mm. Glabella subdiamond-shaped in outline, moderately convex, with a 450 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY rounded crest from which the flattened sides slope steeply, occipi- tal ring gently convex, backwardly projecting, occipital furrow shallow medially, represented distally by faint depressions; anterior part of glabella extended into a long, gently upwardly curving frontal g@labellar spine, this spine suboval in ecross-see- tion; in lateral view two raised longitudinal ridges on the proxi- mal part of the frontal glabellar spine; these ridges unite distally, proximally they run on to the anterior part of the glabella and die out. Fixed cheeks gently convex, triangular in outline, posterior border defined only in the outer part by a shallow pos- terior border furrow that is deepest just inside the facial suture ; in lateral view facial suture only slightly bowed inwards along lateral margin of fixed cheek, anteriorly two branches confluent and bounding anterior margin of glabella. Thoracic segment relatively long (sag. and exs.), convex axial ring and pleural furrow becoming deepest distally; outer part of pleura short (tr.), bent down vertically, fulerum running in a curve outward and backward so that the anterolateral margin of the segment fits closely beneath the bent-down posterolateral portion of the fixed cheek (PI. 33, fig. 3). Transitory pygidium three times as wide as long, axis gently convex, divided obscurely into two rings and a wider (tr.) rounded, gently convex, posterior part. Pleural region horizontal beside anterior part of axis, postero- lateral and posterior parts curved downwards so that close to, and behind, the axis they slope almost vertically, and in this region there is a shallow median notch in the outline; first pleural furrow shallow, faint interpleural groove, second and third pleural furrows extremely faint; narrow raised rim. Larger ecranidia (original of Pl. 33, figs. 9, 11, 12, of width 1.8 mm., of Pl. 33, figs. 14, 16, 17, of width 2.4 mm.) have been associated with free cheeks of appropriate size. The frontal elabellar spine is now subsquare in cross-section, each side excavated and a ridge running longitudinally along the angle, the spine itself with a reduced upward curvature, almost straight. The surface of the frontal lobe of the glabella below the base of the spine is now flat rather than convex, and where it is bounded by the suture it has assumed the characteristic outline of the hol- aspis. The free cheek and doublure are thus much like those of larger specimens, except that in the smaller specimen (PI. 33, WHITTINGTON : ORDOVICIAN TRILOBITES 481 fig. 11) that part of the free cheek outside the suture is relatively narrower (tr.), and the librigenal spine is bent, not hooked, so that the proximal portion runs outward and backward, the distal portion slightly inward and backward. In the larger specimen (Pl. 33, fig. 16) the free cheek has approximately the form of the adult, being wider (tr.), and the librigenal spine has assumed the hooked form and subsquare cross-section, though the hook is not quite so acute as in the larger specimens. A transitory pygidium of width 1.65 mm. (PI. 33, figs. 10, 13, 15) has the axis tapering back, six rings outlined by ring furrows which become progressively fainter posteriorly, and three pleural furrows. The rim is less pronounced than at degree 1 (PI. 338, fig. 7), the outline posterolaterally smoothly curved; in posterior view the border shows the deep and acute posterior notch. An enrolled exoskeleton of degree 4, lacking the free cheeks (Pl. 33, figs. 19, 22, 23, 26, 29, 30; the second thoracic segment lies edgewise between the first and third, the extremity of the left pleura being visible in the ventral view — Pl. 33, fig. 29) has the cranidium much like that of larger specimens. The four thoracic segments decrease in length (sag. and exs.) posteriorly, and in the last three segments the fulcrum is directed progressively more strongly inward as well as backward. The transitory pygidium is partly concealed in this specimen, but an isolated example of similar size (Pl. 33, figs. 18, 21) has four axial rings and three pairs of pleural furrows visible. Cephala up to the largest size obtained (PI. 32, figs. 1, 4, 5, 7, 16, 20) show little further change, the glabella becoming slightly more convex in longitudinal profile, and the lateral coneavity in the facial suture slightly more pronounced. Pygidia of larger size (Pl. 33, figs. 20, 25; Pl. 32, figs. 18, 19, 21, 22) likewise show little change, the pleural furrows become decidedly concavely- curved forward, and the median notch in the border slightly shallower. Genus AMPYXINA Ulrich, 1922 Type Species. Endymionia bellatula Savage, 1917. Redescribed by Whit- tington, 1950b, p. 557, pl. 73, figs. 7-8. At that time it was not realized that the type specimens were in the U. S. National Museum, numbered 72140. The original of Plate 34, figure 1, an external mould in which the posterior part of the cranidium conceals much of the first thoracic segment, is the holotype. 482 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY Discussion. Besides the type and the two species described below, others placed here come from England, Scotland and Sweden (Whittard, 1955, p. 25). Because of the different ways in which these specimens are preserved it is difficult to make com- parisons, or to be sure of such features as the convexity and loba- tion of the glabella, and nature and disposition of the lateral elabellar furrows. The glabella in the type species is rather wider than that of other species, the basal glabellar lobes are represented by shallow oval pits, and the alae merge at the inner posterior corner into the extremity of the occipital ring. In A. powelli (Pl. 34, figs. 3-13), A. wothertonensis Whittard, 1955 (pl. 3, figs. 1-4), and A. lanceola nsp. (PI. 35, fig. 32) three pairs of lateral glabellar furrows appear as subeireular shallow pits. The alae in A. powelli and A. lanceola (Pl. 34, figs. 15, 16) are like those of the type species, whereas in A. wothertonensis and A. aldonensis (P1. 28, fig. 13) they appear to be completely cut off from the occipital ring and posterior border by a furrow, perhaps because the latter specimens are internal moulds. In the pygidium the nature of the ribs between the pleural furrows appears to vary — in the type species and A. wothertonensis these ridges are broad and flat-topped, whereas in A. powelli and A. lanceola n.sp. (Pl. 34, figs. 25-27) they are narrow with steeply sloping sides. It is shown here (PI. 35) that a frontal glabellar spine is present in the development of Ampyzina. There is no sign of this spine in the largest specimens of some species, but in A. lanceola n.sp. there is a short frontal glabellar spine in the largest specimens, as there is in A. aldonen- sis (Reed, 1935) (this paper, Pl. 28, figs. 11, 13). AMPYXINA POWELLI (Raymond, 1920) Plate 34, figures 3-13; Plate 35, figures 1-25. Raphiophorus powelli Raymond, 1920, pp. 276-277; Raymond, 1925, pp. 32-335 pl. i2, fies: 1,2: Ampyzxina scarabeus Butts, 1941, p. 75, pl. 82, figs. 1-3; Cooper and Cooper, 1946, pl. 3, figs. 11, 12. Ampyzina powelli, Cooper, 1953, pp. 18-14, pl. 4, figs. 1-9, 12(?). Holotype. Museum of Comparative Zoology, no. 1598, from the Edinburg formation, Liberty Hall facies, Catawba Valley, Va. Figured Material. USNM, from Lower Edinburg limestone, localities 5 and 14. WHITTINGTON : ORDOVICIAN TRILOBITES 483 Description. The new material makes possible the following additions to Cooper’s deseription : — Occipital ring narrow (sag. and exs.), strongly convex (PI. 34, figs. 10, 12, 13), in dorsal view curved convexly backward. Oc- cipital furrow deep, curving forward distally and ending against the side of the deep, subcireular, basal glabellar furrow (on the inner surface, Plate 34, figure 6, this lateral glabellar furrow appears as a low, circular, raised area). In front of occipital ring, glabella expands forward to maximum width adjacent. to the anterolateral cephalic border, frontal lobe gently rounded; anterior part of glabella moderately convex, that part in front of occipital ring strongly convex, sloping steeply to occipital furrow at basal glabellar furrow. Second and third (from the posterior) lateral glabellar furrows appear as small, suboval, slightly depressed areas over which the exoskeleton is thinner (these thin areas are broken through on right side of original of Plate 34, figure 3) ; the second situated opposite the anterior end of the ala, the third a short distance behind the anterior pit, both adjacent to the shallow axial furrow. Anterior pit deep (Pl. 34, figs. 6, 10, 13). Frontal lobe of glabella overhangs anter- ior cephalic border, and is separated from this border by a shal- low preglabellar furrow (Pl. 34, fig. 12). Ala (referred to by Cooper, 1953, p. 13, as basal lateral lobe of glabella) gently convex, of length (sag.) about one third that of glabella (Pl. 34, figs. 3, 4, 6, 12, 13), separated from the glabella by a shallow axial furrow and from the cheek by a broad shallow alar furrow; inner posterior corner of ala merges into the posterior border adjacent to the occipital ring; outer, posterior part of ala separated from posterior border by a shal- low posterior border furrow. Cheek (Pl. 34, figs. 7, 8, 10-13) moderately convex, narrow anterolateral border defined by a deep border furrow which dies out, both as it approaches the anterior branch of the suture and the base of the librigenal spine; posterior border furrow curves forward distally so that the posterior border widens, furrow expands into shallow pit just inside facial suture; cheeks united in front of glabella by nar- row (sag. and exs.) convex frontal area. Facial suture curves around pit at extremity of posterior border furrow and runs in- ward and forward across the cheek with a slight sinuosity, the 484 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY two branches meeting in a smooth curve along the anterior border. Librigenal spine arises from outer and upper surface of cheek at genal angle, is long and tapering, and curves to run directly backward beyond the pygidium. A narrow doublure (Pl. 34, figs. 7, 8), not crossed by median or connective sutures, unites the free cheeks anteriorly; this doublure is horizontal anterolaterally and anteriorly, and a shallow groove runs just inside the smoothly curved inner margin; beneath the base of the librigenal spine the doublure narrows and there is no doub- lure along the inner part of the posterior border. Three specimens of a hypostome (PI. 34, figs. 5, 9) attributed to this species occur with the abundant material at locality 14. Outline subrectangular, slightly wider than long, anterolateral portion projecting slightly laterally; middle body moderately convex, most strongly so anteromedially. Narrow convex border becomes defined at outer part of anterior margin and curves around to follow lateral and posterior margins. An oval, longi- tudinally elongate, gently convex area situated inside the postero- lateral border. Margins of border bent up vertically but not eurled under to form a doublure. Small projection from postero- lateral part of this margin forms posterior wing. Margin of border between wings excavated in shallow notch. Thorax (PI. 34, figs. 3, 4) of five segments, the anterior mark- edly longer (sag. and exs.) than the succeeding four, which are progressively shorter backward. Axis about one-quarter the width, axial rings narrow (sag.), convex, curved arcuately back- ward, articulating furrow deep, with deep appendiferal pit situated a short distance inside the axial furrow. Articulating half ring absent on first segment, on succeeding four segments about as long (sag.) as the axial ring. Pleurae extend out hori- zontally, are flexed down distally, and at the fulerum a narrow (tr.) portion is bent down vertically and curled under to form an extremely narrow doublure. Pleural furrow of first segment eurved convexly forward, on succeeding segments directed trans- versely, becoming deeper on outward sloping portion. Outer parts of pleurae facetted, facet received by depression in doub- lure of preceding segment. Pygidium (Pl. 34, figs. 3, 4) with axis reaching back to margin of inner part of pleural regions, horder bent down steeply, in posterior view a median notch in the margin. Four to six axial rings outlined by ring furrows WHITTINGTON : ORDOVICIAN TRILOBITES 485 which are deepest distally; five pairs of pleural furrows which deepen distally are successively directed more strongly backward and have a distal flexure. External surface smooth, except for anastomosing raised lines on border of cephalon and outer parts of pleurae of thorax and pygidium. Development Smallest known specimens are isolated eranidia (PI. 35, figs. 1-3), of width (tr.) 0.69 mm. Glabella suboval in outline, moder- ately convex, the flattened sides sloping from the rounded keel ; narrow occipital ring, large deep subcireular basal furrow: frontal lobe prolonged into a thick frontal glabellar spine. Posterior border of convex fixed cheek defined only distally by a pit; ala cannot be distinguished. Next largest specimen is a complete exoskeleton (lacking the hypostome) of degree 2 (PI. 35, figs. 5, 6, 10, 11). The glabella has a long frontal spine, the alae are well defined by axial and alar furrows, and merge with the innermost part of the posterior border; outside the ala is the posterior border furrow; librigenal spine long and directed backward. First thoracic segment longer than the second and having the characteristic forward curve of the pleural furrow. On the pygidium three axial rings and three pairs of pleural furrows may be distinguished, the pleural furrows straight, lacking the distal backward curve. External surface granulate. a row of short spines along the cephalic doublure, similar spines on the posterior border just inside the base of the librigenal spine. In exoskeletons of degree 3 (Pl. 35, figs. 4, 7-9) the glabella is sharply keeled, but in exoskeletons of degree 4 (Pl. 35, fig. 12) the keel is less sharp. In small holaspides (PI. 35, figs. 13, 14) the frontal glabellar spine is about one-quarter the length (sag.) of the remainder of the glabella, the pleural furrows of the pygi- dium have the characteristic distal backward curve, the external surface is granulate, and there are still short spines on the poster- ior border just inside the base of the librigenal spine. In the largest holaspides (Pl. 34, figs. 3, 10-13) the frontal glabellar spine may be absent or may be represented by a small tubercle; the external surface appears smooth and there is no trace of the short spines. 486 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY AMPYXINA LANCEOLA D.sp. Plate 34, figures 14-28; Plate 35, figures 26-35. Ampyxina elegans Cooper, 1953 (part), USNM 116443a,b, originals of Cooper’s plate 4, figures 10, 11 (reproduced at X8, not X4, as stated by Cooper). Holotype. USNM 137692 (PI. 34, figs. 14-17, 22), locality 16. Other Material. Paratypes USNM 137698a-e; all figured material in U. S. National Museum. Geological Horizon and Localities. Lower Edinburg limestone, locality 16; Effna limestone, four miles southwest of Bland Court House, Va. Description. This species occurs quite commonly at locality 16 and may be distinguished from Ampyzina powelli by the following characters: Glabella (compare Pl. 34, figs. 14-17, 22, with Pl. 34, figs. 10, 12, 13) more convex, in dorsal aspect projecting farther in front of the cephalic border, and in the largest specimens bearing a short frontal glabellar spine about one-quarter the length of the rest of the glabella; portion of glabella in front of the occipital ring stands relatively higher and slopes down more steeply to the occipital furrow. Basal lateral glabellar furrow represented by a deep sub-cireular pit; in front of this two further furrows (Pl. 35, fig. 32) are faintly discernible as oval areas over which the exoskeleton is thinner, and are situated a short distance up the side of the glabella; the anterior is the smaller. Alar furrow broader and deeper, and the ala is more inflated, separated from the posterior border by the posterior border furrow; this furrow becomes extremely faint behind the inner, posterior corner of the ala, so that here the ala merges into the extremity of the occipital ring. Lateral border of the cheek less strongly curved outside and just in front of the librigenal spine, consequently the free cheek is relatively narrower (tr.) (compare PI. 34, figs. 14, 15 with Pl. 34, figs. 10, 13). Lateral and anterior cephalic borders narrow, convex, curled under in the narrow doublure which is convex ventrally (Pl. 34, fig. 16). Inner margin of anterior part of doublure faces upward and is rounded, not having the appearance of being a sutural junction. The well preserved speci- mens show clearly the anterolateral border furrow dying out as it approaches the facial suture (PI. 34, fig. 17), only an extremely WHITTINGTON : ORDOVICIAN TRILOBITES 487 shallow border furrow crosses the anterior part of the fixed cheek, and the preglabellar furrow runs into the axial furrow and sepa- rates the glabella from the anterior border. Pygidium (compare PI. 34, figs. 25, 27, with Pl. 34, figs. 3, 4) is somewhat more triangular in outline, for the axis and adjacent part of the pleural regions project shghtly posteriorly ; pleural furrows deeper and broader, especially distally, so that they are separated by narrower, sharper ridges. External surface finely granulate, in some specimens slightly coarser granules on the posterior part of the glabella and a row of tiny spines (PI. 34, fig. 21) along the posterior margin of the occipital ring and posterior border. The small cranidia of A. lanceola (Pl. 35, figs. 26-29, 32, 35) differ from those of comparable size of A. powelli (Pl. 35, figs. 17-24) mainly in the higher glabella and stouter frontal spine. Small pygidia (PI. 35, figs. 30, 31, 33, 84) differ in outline and in the width of the distal part of the pleural furrows. Discussion. A eranidium and pygidium placed in Raymond- ella elegans by Cooper (1953, pl. 4, figs. 10, 11) are here placed in Ampyxina lanceola n.sp. The type material of Ampyxina aldonensis (Reed, 1935) (Pl. 28, figs. 10, 11, 13) from strata of comparable age in Scotland, shows that this species is extremely similar to A. lanceola, particularly in the outline of glabella, the presence of the frontal glabellar spine, the convexity of the alae, and the curvature and width of the pleural furrows of the pygidium. Genus RAYMONDELLA Reed, 1935 Discussion. The type species, Raymondella macconochier (Nicholson and Etheridge, 1879), has been redeseribed (Whit- tington, 1950b, pp. 558-559, pl. 74, fig. 10; pl. 75, figs. 1, 2), and R. elegans Cooper, 1953 is remarkably similar to it. Among raphiophorids Raymondella (P1. 36) seems most closely related to Ampyzina (Pls. 34, 35), as comparison between illustrations will show. Raymondella differs from Ampyxina in form of the glabella (compare Pl. 36, figs. 1-5 with Pl. 34, figs. 1-17, 22) in front of the occipital ring — the basal part is narrow, sloping upwards, and expanding rapidly into a spherical frontal portion which projects and overhangs well in front of the anterior 488 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY cephalic border; the basal lateral glabellar furrow is a subcir- cular, deeply impressed pit lying immediately in front of the outer part of the occipital ring; in front of this furrow the glabella expands rapidly. The ala is smaller, subcireular in out- line and defined by a deep, broad, alar furrow which runs back- ward and slightly outward from the axial furrow to the posterior border furrow. The convex cheek of Raymondella, inside the border furrows, is crossed by raised anastomosing ridges, a type of sculpture that is not known in Ampyxina. Axial rings of thorax and pygidium (compare PI. 36, figs. 24-26, 29-32, 33, 34, 37, with Pl. 34, figs. 1, 3, 4, 18-20, 25-27) more convex, articulat- ing furrows and appendiferal pits deeper, pleural furrows deeper, and that of the first thoracic segment (PI. 36, figs. 24-26) lacking the curve convex forward of Ampyxina (PI. 34, figs. 3, 18), pleural furrows of pygidium wider distally and sep- arated by narrow, high ridges. RAYMONDELLA ELEGANS (Cooper, 1953) Plate 36 Ampyxina elegans Cooper, 1953 (part), p. 14, pl. 4, figs. 14, 15, 16, 18, and 19. The originals of Cooper’s figures 10 and 11, plate 4, are here referred to Ampyxina lanceola n. sp., and those of his figures 13 and 17, plate 4, to the pygidium of Tretaspis sagenosus n. sp. Holotype. USNM 116445C, a cranidium from the present locality 7. Figured Material. USNM 116446, original of Cooper, 1953, pl. 4, fig. 16; additional material figured here, in U. S. National Museum. Geological Horizon and Localities. Lower Edinburg formation, localities 3, 7, 16; Cherry Grove, 4 miles northwest of Linville Station, Rockingham Co., Va: Description. Largest cranidium here figured (Pl. 36, figs. 1-5) of maximum width 3.25 mm. Occipital ring narrow (sag. and exs.), convex, gently arcuate backward ; deep, wide, occipital furrow passes distally into the subcircular pit of the basal lateral glabellar furrow ; between these furrows glabella narrow, convex, stalklike, in front of these furrows expanding rapidly into the subspherical median and frontal portions, which project forward and downward over the anterior border; maximum width at about two-fifths the length. Short, blunt, frontal glabellar spine situated on the vertical anterior slope, axial furrow and anterior pit deep. WHITTINGTON : ORDOVICIAN TRILOBITES 489 Alar furrow diverges from the axial furrow at a point about opposite two-thirds the length of the glabella, and runs backward and slightly outward to the posterior border furrow, outer slope steep. Between the alar and axial furrows a triangular area is defined which is depressed below the fixed cheek, and is gently convex only at the inner posterior corner; this convex portion is subeireular in outline and is here termed the ala. The posterior border furrow becomes shallower as it passes inward behind the ala, and thus the inner posterior corner of the ala is separated by only a slight depression from the extremity of the occipital ring. Cheek convex, sloping steeply anterolaterally to the narrow convex border, which is defined by a deep border furrow; border furrow becomes shallow as it approaches the facial suture, nar- row convex anterior border separated from the glabella by the deep preglabellar furrow. Two branches of facial suture con- fluent anteriorly along the margin of the border, laterally having a sinuous course across the cheek. Librigenal spine originates from the posterolateral border and upper posterolateral surface of the cheek, and curves to run directly backward. Cephalic doublure narrow laterally and anteriorly, convex ventrally, absent along posterior border and beneath occipital ring. Hypostome unknown. Posterior border convex, bent down dis- tally, posterior border furrow deep, a distal pit just inside the point where it is crossed by the facial suture. Only isolated thoracic segments known (Pl. 36, figs. 24-26, 29-32). The anterior segment can be recognized by the curved outline of the anterolateral part of the pleura where it fits beneath the posterior cephalic border. Other segments have a rectangular outline to the distal part of the pleura. Axial ring narrow (sag. and exs.) and convex, articulating furrow deep, appendiferal pit deep and elongate, situated a short distance in from the axial furrow, appendifer a thick triangular projection. Pleural furrow deep and broad, running out transversely, flanked by convex ridges. Outer part of pleura bent down gently, the outermost part flexed vertically and curled under to form a nar- row doublure. Anterolateral angle of pleura facetted, and doub- lure with a depression to receive the facet of the following segment during enrollment. Pygidium (PI. 36, figs. 33, 34, 37) four times as wide as long (sag.), convex axis divided by deep furrows into four rings. Inner part of pleural region horizontal, 490 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY deeply excavated by three pleural furrows that widen rapidly distally, these furrows separated by strong interpleural ridges which run outward and backward to join the rim. Border bent down almost vertically, outline of outer edge in posterior view sinuous, with a median notch. Doublure extremely narrow. External surface of glabella, axial rings, and ridges of thorax and pygidium granulate, the granules coarsest in a row along the occipital and axial rings, the posterior cephalic border, and the posterior ridge of the thoracic pleurae. Cheek inside border furrows crossed by raised, anastomosing lines which run sub- parallel to the anterolateral margin. On the cephalic doublure, the outer parts of the pleurae, and on the border of the pygidium, raised lines run subparallel to the margin. Discussion. The pygidium attributed to Raymondella elegans by Cooper (1953, p. 14, pl. 4, figs. 138, 17) is considered here to be that of Tretaspis sagenosus n.sp. These two species occur together at localities 3, 7, and 16, Tretaspis sagenosus occurring rarely at a few other localities. The associations of exoskeletal parts made here are based on morphological comparisons between occipital ring and posterior border of cephalon, and thoracic segments and pygidia. They seem to be upheld by the likeness between the pygidium of Ff. elegans and the type species, and the likeness between the pygidium of Tretaspis sagenosus n.sp. and that of other species of this genus. Cooper (1953, p. 14) states that the width of the holotype cranidium is 4.8 mm. This figure may have been printed in error, since, using the magnifications given in his plate 4, figures 14, 15, 18 and 19, the width of the holotype cranidium appears to be about 3.5 mm. The largest specimens I have seen in both Cooper’s and my material are of about this size. Another species of Raymondella, from the Oranda formation, Chambersburg, Pa., is that described by Cooper (1953, pp. 14-15) as “‘Ampywxina sp.’ The cranidium is typical, but the glabella (Pl. 28, figs. 15, 16, 19) appears to differ from that of R. elegans in having flattened sides and a rounded median keel. Development A size series of cranidia from a width of about 1.0 mm. to 3.5 mm. is known. The smallest cranidia (PI. 36, figs. 18, 19, 22, WHITTINGTON : ORDOVICIAN TRILOBITES 49] 23) are like those of comparable size of other raphiophorids (Pl. 31, figs. 7-9; Pl. 35, figs. 5, 6, 10, 11), the glabella clavate, pinched in in front of the occipital ring by the basal glabellar furrows, frontal lobe projecting beyond the anterior border and continued by a thick, straight, frontal glabellar spine. A rela- tively large triangular area is enclosed between the alar and axial furrows, and the posterior border furrow is shallow. As size increases (Pl. 36, figs. 16, 17, 21; 10, 15, 20; 12-14; 8, 9; 6, 7, 11), the most conspicuous changes occur in the glabella. In the early stages the glabella is keeled, with flat, steeply sloping sides. As the part in front of the basal glabellar furrows swells up to become subspherical, the median portion becomes higher and rounded in transverse section, the transverse profile changing from triangular to circular ; the frontal part of the glabella bulges forward and downward over the anterior border, and the frontal glabellar spine is progressively reduced to the short, blunt-tipped spine of the largest cranidia. The posterior border furrow be- comes deeper with increasing size, and at all times it seems to separate the inner posterior convex portion of the ala from the posterior border. The smallest known pygidia (Pl. 36, figs. 27, 28) differ from the largest in not exhibiting the fourth axial ring. External surface of cephalon in the small stages granulate, row of tiny spines around the anterior and lateral margin. In the larger specimens the surface is similar, except that on the cheek granulation has fused to form the anastomosing lines. Family ENDYMIONIIDAE Raymond, 1920 Genus SALTERIA Thomson, 1864 SALTERIA AMERICANA Cooper, 1953 Plate 28, figures 9, 12, 14, 17, 18 Discussion. Cooper (1953, pp. 12-13, pl. 3, figs. 8-10) based this species on material from the lower part of the Liberty Hall facies of the Edinburg limestone in Rockingham and Pulaski counties, Va. A few additional silicified specimens have been obtained from the present locality 14. 492 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY Occipital ring is extremely narrow (sag. and exs.) ; the ‘‘raised posterior carina’’ is not evident. In front of the preglabellar furrow is a narrow (sag. and exs.) preglabellar field, separated by a shallow anterior border furrow from the narrow anterior border. No trace of genal caeca. Cranidium bounded by facial suture which runs forward in a curve convex outward across the posterior border, and then forward and slightly inward across the cheek in a sinuous course, reaching the anterior border out- side the projected line of the axial furrow and running along the margin of this border. Free cheek and cephalic doublure unknown. Thorax of 7 segments, the anterior segment the longest (sag. and exs.). Pleurae inflated, inner part horizontal, crossed by a deep, narrow, pleural furrow which runs in a curve coneave for- ward and deepens distally. Distal part of pleurae bent down in two stages, the first sloping gently outward, the outermost part, beyond the end of the pleural furrow, narrow (tr.) and bent down vertically. Pygidium more than twice as wide as long (sag.). Pleural regions with a broad, shallow, border furrow outside which is a convex border, the outermost part of this border narrow and sloping almost vertically. Six pairs of deep, narrow pleural furrows running in eurve coneave forwards, dying out at the inner margin of the border furrow. Doublure of thoracic pleurae and pleural regions extremely narrow, being merely the tightly rolled-under edge of the exoskeleton. Sculp- ture of the external surface of the exoskeleton not well preserved, but fine raised lines run along the doublure of the pygidium. As Cooper (1953, p. 13) pointed out, this species is exceedingly like the type species Salteria primaeva, from the Balelatchie beds, lower Middle Ordovician, Scotland (Reed, 1903, pp. 43-44, pl. 4, figs. 18, 14). I have placed this genus in the family Endymioniidae, but the cephalon of Salteria resembles that of Seleneceme (Whittington, 1952a, p. 4, pl. 2, fig. 13) in the sub- circular outline of the glabella, narrowness (sag.) of the occipital ring, breadth of the posterior cephalie border, and course of the facial suture. REFERENCES Burts, C. 1941. Geology of the Appalachian Valley in Virginia, Part 2, fossil plates and explanations. Virginia Geol. Surv., Bull. 52, pp. 1-271, pls. 1-135. WHITTINGTON : ORDOVICIAN TRILOBITES 493 Cooper, B. N. 1953. Trilobites from the Lower Champlainian formations of the Ap- palachian Valley. Geol. Soc. Am., Mem. 55, pp. 1-69, pls. 1-19, 2 figs., table. Cooper, B. N. and G. A. COOPER 1946. Lower Middle Ordovician stratigraphy of the Shenandoah Valley, Virginia. Geol. Soc. Am., Bull., vol. 57, pp. 35-114, pls. 1-3, 9 figs. Cooper, G. A. 1956. Chazyan and related brachiopods. Smithsonian Mise. Coll., vol. 127 (in two parts), pp. 1-1245, pls. 1-269. Evirr, W. R. 1951. Some Middle Ordovician trilobites of the families Cheiruridae, Harpidae and Lichidae. Jour. Paleont., vol. 25, pp. 587-616, pls. 85-88, 1 fig. 1953. Observations on the trilobite Ceraurus. Jour. Paleont., vol. 27, pp. 33-48, pls. 5-8, 1 fig. Evirr, W. R. and H. B. WHITTINGTON 1953. The exoskeleton of Flexicalymene (Trilobita). Jour. Paleont., vol. 27, pp. 49-55, pls. 9, 10, 1 fig. Hupk, P. 1953. Classe des Trilobites. Jn Traité de Paléontologie, vol. 3, pp. 44- 246, Ed. J. Piveteau. Paris. Kay, M. 1958. Ordovician Highgate Springs sequence of Vermont and Quebec and Ordovician classification. Amer. Jour. Sci., vol. 256, pp. 65-96, figs. 1-11. KOBAYASHI, T. 1951. On the Ordovician trilobites in Central China. Jour, Fac. Sci. Univ. Tokyo, sec. 2, vol. 8, part 1, pp. 1-87, pls. 1-5. LINDSTROM, G, 1901. Researches on the visual organs of the trilobites. K. Svenska Vet.-Akad. Handl., vol. 34, no. 8, pp. 1-86, pls. 1-6. lia Ye He 1957. Trilobita. Jn Index Fossils of China, Invertebrate, part 3, pp. 249-294, pls. 137-155. Geological Press, Peking. OPIK, A. 1926. Beitrige zur Kenntnis der Kukruse-(C2) Stufe in Eesti. I. Pub. Geol. Inst. Tartu, no. 4, pp. 1-18, pls. 1, 2. 1937. Trilobiten aus Estland. Pub. Geol. Inst. Univ. Tartu, no. 52, pp. 1-163, pls. 1-27. 494 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY RAYMOND, P. E. 1920. Some new Ordovician trilobites. Mus. Comp. Zool., Bull., vol. 64, no. 2, pp. 273-296. 1925. Some trilobites of the Lower Middle Ordovician of eastern North America. Mus. Comp. Zool., Bull., vol. 67, no. 1, pp. 1-180, pls. 1-10. REED, F. R. C. 1903. The Lower Palaeozoic trilobites of the Girvan District, Ayrshire, Part 1. Paleontogr. Soe., London, pp. 1-48, pls. 1-6. 1935. The Lower Palaeozoic trilobites of the Girvan District, Ayrshire, Supp. No. 3. Paleontogr. Soe., London, pp. 1-64, pls. 1-4. Ross, R. J. 1948. Revisions in the terminology of trilobites. Amer. Jour. Sci., vol. 246, pp. 573-577. 1951. Ontogenies of three Garden City (early Ordovician) trilobites. Jour. Paleont., vol. 25, pp. 578-586, pls. 81-84. 1953. Additional Garden City (early Ordovician) trilobites. Jour. Paleont., vol. 27, pp. 633-646, pls. 62-64. RUEDEMANN, R. 1901. Trenton conglomerate of Rysedorph hill and its fauna. N. Y. State Mus., Bull. 49, pp. 3-114, pls. A, B, 1-7. 1916. The cephalic suture lines of Cryptolithus (Trinucleus auct.). N. Y. State Mus., Bull. 189, pp. 144-148, pl. 35, figs. 43-46. SavaeE, T. E. 1917. The Thebes Sandstone and Orchard Creek shale and their faunas in Illinois. Trans. Illinois Acad. Sci., vol. 10, pp. 261-275, pls. 12s SKJESETH, 8. 1955. The Middle Ordovician of the Oslo Region, Norway, 5, the trilo- bite family Styginidae. Norsk. Geol. Tids., vol. 35, pp. 9-28, pls. 1-5; STAUBLE, A. 1953. Two new species of the family Cryptolithidae. Natur. Canadien, vol. 80, pp. 85-119, 201-220, figs. 1-24. STeRMER, L. 1930. Scandinavian Trinucleidae with special reference to Norwegian species and varieties. Vid.-Akad. Oslo, Skr. I, Mat.-Naturv. Klasse, no. 4, pp. 1-111, pls. 1-14, 47 figs. 1940. Early descriptions of Norwegian trilobites. Norsk. Geol. Tids., vol. 20, pp. 113-151, pls. 1-3. WHITTINGTON : ORDOVICIAN TRILOBITES 495 1949. Classe des Trilobites. Jn Traité de Zoologie. Directed by P.-P. Grasse, Masson et cie, Paris, vol. 6, pp. 160-197, figs. 1-24. THORSLUND, P. 1940. On the Chasmops series of Jemtland and Séddermanland (Tvaren). Sveriges Geol. Under., ser. C, no. 436, pp. 1-191, pls. 1-12, 56 figs., map. TRIPP, R. P. 1954. Caradocian trilobites from mudstones at Craighead Quarry, near Girvan, Ayrshire. Royal Soe. Edinburgh, Trans., vol. 62, part 3 (no. 16), pp. 655-693, pls. 1-4. WARBURG, E. 1925. The trilobites of the Leptaena limestone in Dalarne. Geol. Instit. Upsala, Bull., vol. 17, pp. 1-446, pls. 1-11, 23 figs. W HITTARD, W. 10, 1955. The Ordovician trilobites of the Shelve Inlier, West Shropshire, Part 1. Paleontogr. Soe., London, pp. 1-40, pls. 1-4. WHITTINGTON, H. B. 1941a. The Trinculeidae — with special reference to North American genera and species. Jour. Paleont., vol. 15, pp. 21-41, pls. 5, 6, 1 fig. 1941b. Silicified Trenton trilobites. Jour. Paleont., vol. 15, pp. 492-522, pls. 72-75, 13 figs. 1949. Dolichoharpes and the origin of the harpid fringe. Amer. Jour. Sci., vol. 247, pp. 276-285, pls. 1, 2. 1950a. British trilobites of the family Harpidae. Paleontogr. Soc., London, pp. 1-55, pls. 1-7, figs. 1-16. 1950b. Sixteen Ordovician genotype trilobites. Jour. Paleont., vol. 24, pp. 531-565, pls. 68-75, figs. 1-9. 1952a. The trilobite family Dionididae. Jour. Paleont., vol. 26, pp. 1-11, pls. 1-25 figs 1. 1952b. A unique remopleuridid trilobite. Breviora, Mus. Comp. Zool., no. 4, pp. 1-9, pl. 1. 1956. Silicified Middle Ordovician trilobites: the Odontopleuridae. Mus. Comp. Zool., Bull., vol. 114, no. 5, pp. 155-288, pls. 1-24, figs. 1-25. 1957. The ontogeny of trilobites. Biol. Rev., vol. 32, pp. 421-469, figs. 1-29. 1959. Trilobita. Jn Treatise on Invertebrate Paleontology, Part O, Arthropoda 1. Direct. and Ed. Raymond C. Moore, Geol. Soc. Amer. and Univ. Kansas Press, pp. 038-0560, figs. 27-415. 496 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY WHITTINGTON, H. B. and W. R. Evirt 1954. Silicified Middle Ordovician trilobites. Geol. Soc. Am., Mem. 59, pp. 1-137, pls. 1-33, 27 figs. WHITTINGTON, H. B. and ALLEN 8S. Hunt 1958. Growth of the cephalon of Cryptolithus (Trilobita) (Abstract). Geol. Soe. Amer., Bull., vol. 69, p. 1662. EXPLANATION OF PLATES 1-36 To make the photographs, the larger specimens were mounted on insect pins with gum arabic, the smaller specimens laid on a black surface. A light coating of ammonium chloride was used. Stereoscopic pairs of views were made by tilting the specimen, and only the left-hand view of a pair is numbered. The heads and shafts of the pins have been blacked out; otherwise the photographs have not been retouched. It is arbitrarily decided that the plane passing through the margin of the occipital or axial ring shall run in the dorsoventral direction, and views are described accordingly. Exterior or interior views have been taken in a direction lying in the sagittal plane to give the fullest possible view of the exoskeletal surface. Oblique exterior or in- terior views have been taken in directions at an angle to the sagittal plane. The initials USNM refer to the U.S. National Museum, Washington, D.C., the initials HM to the Hunterian Museum, Glasgow, Scotland, and the initials MCZ to the Museum of Comparative Zoology. These initials are followed by the cata- logue numbers. i im 1 ght ¥ oe ft ih een re aww ah chives Te ST ED 4 inlet Moll wd wee! yl abe yavee peste te worn i ee Lvvtidoranie’ wt. -fiaas fa erste. Beene! ot vin Tins, tenant bai ernting tain Fels wh ow'ml oop aciey allt Tig.evh a boast ievite @ 4 dimes bei} ba a etn labicyivad, aly te on Seah i¥i myrict Fete, any Hig wil Pogy ‘De Prk, Cae heer lardae niephl: Gh? ni y0)% Viele 4 real bran avale TiO aim $k on Ae ie we ow -wheothiiadruyiy erdpeli tire al ai pels inion 6 ‘aes fn Ptotearhe sa hoon tii Trip tones fate tie arity ilies : wis | elon i be oben e-s4i 3 Hines] Habit, seal ied ' batihee, 2 sien | shad Yors |) otetlion oath audi ee daiaicasattD welt shi Wied atbiipd any Tt tery ita oe ape od? eh SY adit GA irate? erga oben Bear oth) Genwi ay Neg Ay ite. A' ilk Omg matt h i = — . . or) done ene - = - ' - : oe ¥ > e 7 =— , PLATES PLATE 1 Remopleurides caelatus n.sp. Lower Edinburg formation Holotype cranidium, dorsal, anterior views, X 15. USNM 137675. Locality 2. Cranidium: anterior, dorsal, ventral, posterior, right lateral views, X 6. Locality 2. Paratype free cheek: dorsal, oblique exterior views, X 6. USNM 137676a. Locality 6. Hypostome: right lateral view, X9. Locality 2. Free cheek and paratype hypostome: ventral, anteroventral, dorsal views, X 9. Locality 2. Paratype hypostome: oblique interior, posterior views, X 9; posterodorsal view, X 15. USNM 1387676b. Locality 2. PLATE 1 nN & Ss =< re Ay Figure TONS 455 6, 7, 8, 9, 10 11, 12,13 PLATE 2 Remopleurides caelatus n.sp. Lower Edinburg formation Paratype thoracic segment: anterior, dorsal, posterior, ventral, left lateral views, X 9. USNM 137676c. Locality 15. Paratype, two articulated thoracic segments: dorsal, anterior, ventral, posterior, right lateral views, X 9. USNM 137676d. Locality 15. Paratype, posterior four thoracic segments and pygidium, articulated: right lateral, dorsal views, X 9; ventral view, X 7. USNM 137676e. Locality 6. Pygidium: dorsal, posterior views, X 9. Locality 2. Pygidium: dorsal, posterior views, X 9. Locality 2. Posterior four thoracic segments and pygidium, articulated: dorsal view, X 6. Locality 6. Posterior thoracic segment and pygidium, articulated: dorsal view, X 15; posterior view, X 9. Locality 2. Pygidium: dorsal view, X 15. Locality 2. Part of paratype free cheek: exterior view, X 30. USNM 137676a. Locality 6. Cranidium and left free cheek: oblique interior view, X 9. Locality 2. Detail of left free cheek, original of figure 23: interior view showing cone extending inward from pit in external surface, X 30. Locality 2. Detail of tongue of cranidium, original of figure 23: interior view showing pit in anterior boss, X 30. Loeality 2. Figure (or) chy JETS 1105) 10, 12, 13, 14 16 Vi l8 PLATE 3 Remopleurides caelatus n.sp. Lower Edinburg formation Protaspis: dorsal, posterior, ventral, lateral views; oblique exterior stereograph, X 30. Locality 6. Broken posterior portion of protaspis: oblique exterior view, X 30. Locality 3. Degree 0 cranidium: ventral, dorsal stereographs, X 30. Lo- cality 6. Degree 0 transitory pyigidium: ventral, right lateral, dorsal views, X 30. Locality 6. Degree 0 transitory pygidium: right lateral, dorsal, posterior, ventral views, X 30. Locality 2. Meraspid cranidium, locality 2: free cheek, locality 3: dorsal stereograph, X 30. Small free cheek: dorsal, ventral views, X 15. Locality 16. iA ae aby ; fai, PLATE 4 Figure 9 ,- Or fe CO 6,13 7,8 9, 10 11,12 14, 15 16, 17,18 19, 20 21 99 are 99 pars) 24 25 PLATE 4 Remopleurides caelatus n.sp. Lower Edinburg formation Small eranidium: dorsal, anterior views, X 15. Locality 6. Small hypostome: exterior view, X 15. Locality 6. Small hypostome: exterior view, X 15. Locality 2. Degree 5? transitory pygidium: ventral view, X 30. Locality 3. Degree 5? transitory pygidium: dorsal, left lateral views, X 30. Loeality 3. Small ecranidium: dorsal, anterior views, X 15. Locality 6. Small eranidium: dorsal, anterior views, X 15. Locality 2. Small free cheek: ventral, dorsal views, X 15. Locality 15. Small cranidium: dorsal, anterior views, X 9. Locality 6. Degree 6? transitory pygidium: ventral, right lateral, dorsal views, X 30. Loeality 3. Small cranidium: anterior, dorsal views, X 9. Locality 6. Small holaspid pygidium: dorsal view, X 30. Locality 2. Degree 7? transitory pygidium: anterolateral view, X 30. Lo- cality 3. Transitory pygidium: dorsal view, X 30. Locality 3. Degree 9? transitory pygidium: dorsal view, X 30. Locality 3. Transitory pygidium: dorsal view, X 30. Locality 3. Remopleurides plaesiourus n.sp. Lower Martinsburg limestone, locality 9 Paratype hypostome: exterior, left lateral, posterior views, X 6. USNM 137678b. Paratype free cheek: ventral view, X 6. USNM 137678d. Part of paratype free cheek, oblique exterior view showing eye surface, X 30. Figure 11 12, 13,14 15 16,17 18, 19, 20, 26 21, 22, 23 24, 25 PLATE 5 Remopleurides plaesiourus n.sp. Lower Martinsburg shale Cranidium: dorsal, ventral (photographed under alcohol to show thinner exoskeleton over muscle areas) views, X 6. Loeality 9. Holotype cranidium: anterior, right lateral, dorsal views, X 6. USNM 137677. Locality 9. Paratype free cheek: dorsal, oblique exterior views, X 6. USNM 137678d. Locality 9. Pygidium: dorsal view, X 9. Locality 10. Pygidium: dorsal view, X 9. Locality 10. Paratype eighth (?) thoracic segment, dorsal view, X 6. USNM 137678a. Locality 9. Paratype thoracic segments: exterior view, X 6. USNM 137678e. Locality 9. Paratype posterior thoracic segment and pygidium: right lateral, dorsal, ventral views, X 6. USNM 137678c. Locality 10. Pygidium: dorsal view, X 15. Locality 10. Degree 0 cranidium: exterior, interior views, X 30. Locality 10. Protaspis: dorsal, posterior, left lateral, oblique ventral views, X 30. Loeality 10. Degree 0 transitory pygidium: exterior, right lateral, posterior views, X 30. Locality 10. Transitory pygidium: ventral, dorsal views, X 30. Locality 10. PLATE 5 PLATE 6 Figure 16, 17 18 19 20, 21, 22 23 24 25 26, 28, 33 30 27 29, 30, 31, 32 PLATE 6 Remopleurides plaesiourus u.sp. Lower Martinsburg shale, locality 10 Degree 0 cranidium: exterior view, X 30. Small meraspid cranidium: exterior view, X 30. Meraspid cranidium and free cheek: exterior view, X 30. Meraspid eranidium: dorsal, anterior views, X 15. Meraspid free cheek: dorsal view, X 15. Small ecranidium: dorsal, anterior views, X 9. Cranidium: anterior, dorsal views, X 9. Cranidium: dorsal, anterior views, X 6. Small holaspid pygidium: dorsal view, X 30. Hypostome: exterior view, X 15. Robergiella sagittalis n.gen., n.sp. Lower Edinburg formation (Liberty Hall facies ) Holotype, incomplete cranidium, free cheek, first thoracic seg- ment: exterior, interior views, X 6. USNM 137679. Locality 14. Cranidium: dorsal view, X 6. Locality 14. Thoracie segment: exterior view, X 6. Locality 16. Cranidium: dorsal, anterior, right lateral views, X 9. Locality 16. Free cheek: interior view, X 6. Locality 16. Thoracic segment: dorsal view, X 6. Locality 16. Small pygidium: dorsal view, X 9. Locality 16. Free cheek: dorsal, interior views, X 10; lateral view, X 30. Locality 16. Free cheek: exterior view, X 6. Locality 14. Paratype pygidium: dorsal, right lateral, posterior, ventral views, X 6. USNM 137680. Locality 14. PLATE 7 Kemopleurides caphyroides n.sp. Lower Edinburg formation Figure Wey 1G Holotype cranidium, locality 3, USNM 137681; paratype free cheek, locality 2, USNM 137682a: anterior, dorsal views, X 6; oblique exterior stereograph, X 4. 2,8 Holotype cranidium: left lateral, ventral views, X 6. Locality Bio Ce (ig) Paratype hypostome (USNM 137682b) and free cheek in juxta- position: interior, exterior, oblique anterior, oblique interior views, X 414. Locality 3. 6 Paratype free cheek: Ventral view, X 6. Locality 2. - a Ss