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ISSN 0097-4463
AN NALS
0/ CARNEGIE MUSEUM
CARNEGIE MUSEUM OF NATURAL HISTORY
4400 FORBES AVENUE ® PITTSBURGH, PENNSYLVANIA 15213
ARTICLE 1
23 MAY 1986
VOLUME 55
CAPTORHINID REPTILES FROM THE EARLY PERMIAN OF NEW MEXICO, WITH DESCRIPTION OF A NEW GENUS AND SPECIES
David S Berman
Associate Curator, Section of Vertebrate Fossils
Robert R. Reisz^
Research Associate, Section of Vertebrate Fossils
Abstract
A new genus and species of single-tooth-rowed captorhinid, Rhiodenticulatus heatoni, is based on two skulls and partial postcranial skeletons collected from the Lower Permian Cutler Formation near Arroyo de Agua, north-central New Mexico. A cladistic analysis of its relationships to other single-tooth-rowed captorhinids suggests that it is a primitive sister taxon to Labidosaurus and Eocaptorhinus. The dentition of R. heatoni, however, exhibits several unique derived features which are interpreted as representing an adap- tation to a specialized diet.
Puercosaurus obtiisidens Williston, 1916, the only previously described captorhinid from New Mexico, is declared a nomen dubium because the holotypic left dentary is indeterminate, and there is no basis for accepting that it and the two poorly preserved captorhinid skulls found at a different locality and referred to the species by Williston (1916) are conspecific. Additional captorhinid remains have been collected recently from the Lower Permian Cutler, Abo, and Sangre de Cristo formations at widely scattered localities in central and northern New Mexico. Though these specimens, as well as the skulls referred to ^‘‘Puercosaurus obtusidens,"’’ are too poorly preserved to be assigned to existing or new taxa, they do indicate that the Captorhinidae was diverse and widely distributed in the Lower Permian of New Mexico.
‘ Address: Department of Biology, Erindale Campus, Univer Ontario L5L 1C6, Canada.
Submitted 2 August 1985.
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Introduction
Published accounts of captorhinid reptiles from the late Paleozoic of New Mexico have been limited to two reports (Williston, 1916; Langston, 1953). Williston (1916) described a small captorhinid, Puer~ cosaurus obtusidens, on the basis of three poorly preserved and incom- plete specimens, a left dentary and two skulls, collected from the Cutler Formation in the Rio Puerco drainage in the north-central part of the state. Further discoveries of captorhinids were not made until 1934- 1935, when collecting was resumed by field parties from the University of California, Berkeley. While conducting extensive field work in the Lower Permian Cutler Formation of the same area, three moderately well preserved specimens, including two skulls with jaws and articu- lated postcranial materials were found at the well known Camp quarry near the small village of Arroyo de Agua (see Langston, 1953, for histories and vertebrate assemblages of well known localities of the area). The only published report of these specimens was a brief reference to them by Langston (1953) in a discussion of the age of the late Paleozoic vertebrate-bearing strata of New Mexico. Here he notes (1953: 410) “a small romeriid cotylosaur possibly referable to Puercosaurus obtusidens is more primitive than Romeria texana of the middle Wich- ita (Putnam)” of the Lower Permian of Texas. Extensive collecting by the authors during the past several years in the Lower Permian deposits throughout New Mexico has resulted in the discovery of additional captorhinid remains from the Cutler, Abo, and Sangre de Cristo for- mations.
Taxonomic evaluation of the undescribed captorhinid materials of New Mexico has necessitated a reexamination of the type specimens of Puercosaurus obtusidens Williston (1916). The partial left dentary, designated by Williston as the holotype, is not only indeterminate, but also provides no basis for considering it conspecific with the two partial, crushed skulls referred by him to the species. Even though the two referred skulls are undoubtedly captorhinids, they are too poorly pre- served to be assigned to an established or new taxon. Under these circumstances P. obtusidens is judged a nomen dubium. On the other hand, the specimens collected by the University of California, Berkeley, are sufficiently well preserved and unique to be referred to a new genus and species, Rhiodenticulatus heatoni. With the exception of the types of this species, all other Lower Permian captorhinid specimens from New Mexico are too incomplete to recommend assignment to existing or new taxa. Yet, they exhibit sufficient variation to indicate that the group was probably quite diverse and widely distributed in New Mexico during the Early Permian.
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Throughout the text the abbreviations CM, FMNH, and UCMP are used to refer to collections of the Carnegie Museum of Natural History, Field Museum, Chicago, and the Museum of Paleontology, University of California, Berkeley, respectively.
Systematic Paleontology
Class Reptilia Order Cotylosauria Suborder Captorhinomorpha Family Captorhinidae Genus Puercosaurus Williston, 1916 Puercosaurus obtusidens Williston, 1916, nomen dubium
Puercosaurus obtusidens Williston, 1916:189-192, fig. 37A~D.
Remarks. -—The original description of Puercosaurus obtusidens Wil- liston (1916) was based on poorly preserved and incomplete speci- mens—an incomplete dentigerous left dentary, FMNH 743, designated as the holotype and two severely crushed skulls, FMNH 745, referred to the species (Fig. 6; only one of the skulls is figured). Williston (1916) illustrated the mandible and one of the two skulls, but a partial recon- struction of the skull was based on both skulls. Although the specimens were collected from the Lower Permian Cutler Formation near Arroyo de Agua in the Rio Puerco drainage area, north-central New Mexico, the holotypic dentary is from the well known Miller bonebed (see Langston, 1953, for description of locality), whereas the referred skulls were apparently found at least several kilometers away along the Rio Puerco (Williston, 1916). The holotypic dentary is too poorly preserved and incomplete to be reasonably certain that it belongs to that family. Further, the holotypic dentary and referred skulls do not exhibit any unique features in common which would demonstrate that they are conspecific. In view of these circumstances P. obtusidens is declared here a nomen dubium. Though the skulls FMNH 745 are sufficiently complete to recognize their captorhinid affinities, assignment to either a known or new species is not possible.
Genus Rhiodenticulatus^ new genus
Type species. — Rhiodenticulatus heatoni, new species.
Etymology.— Prom the Greek rhio, nose, and denticulatus, with small teeth, referring to the relatively small teeth of the premaxilla.
Diagnosis. —Small captorhinid that differs from all other single-tooth- rowed captorhinids in the following features: 1) premaxillary dentition reduced to three teeth which are subequal in size and equal to or smaller than precanine maxillary teeth; 2) reduction of maxillary dentition to 1 1 teeth; 3) number of precanines reduced to two; 4) extremely large
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A
Fig. \.—Rhiodenticulatus heatoni, holotype, UCMP 35757. Skull in A, lateral, B, dorsal, and C, ventral views. Abbreviations: a, angular; art, articular; bo, basioccipital; d, den- tary; f, frontal; j, jugal; 1, lacrimal; m, maxilla; n, nasal; p, parietal; pa, prearticular; pf, postfrontal; po, postorbital; pp, postparietal; prf, prefrontal; pt, pterygoid; q, quadrate; qj, quadratojugal; s, stapes; sp, splenial; sq, squamosal. Scale = 1 cm.
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single canine with basal diameter as much as twice that of any post- canine; 5) very broad lacrimal with a height (measured at the highest level of the dorsal expansion of the maxilla) to length (shortest distance between orbit and naris) ratio of .65 to .73; and 6) prefrontal extends far anteriorly to a level about 84 to 90% of the distance from the orbit to the naris. Distinguished from Labidosaurus and Eocaptorhinus by its proportionately narrower skull postorbitally. Straight occipital mar- gin of skull table separates it from Romeria which has a bilateral parietal embayment and from Labidosaurus and Eocaptorhinus which have a median embayment. Differs from Romeria and Protocaptorhinus in having a long, low rectangular quadratojugal with a longitudinal length that is approximately four times the height. Pointed postcanine teeth of Rhiodenticulatus heatoni are distinguishable from the blunt teeth of Labidosaurus and Eocaptorhinus.
Rhiodenticulatus heatoni, new species
Etymology.— ISlsLmQd. in honor of the late Malcolm J. Heaton in recognition of his significant contributions to our understanding of the morphology and phylogenetics of the Captorhinidae.
Holotype. — UCMP 35757: partial, articulated skeleton that includes skull with closely joined lower jaw, vertebral series with ribs from the axis to the seventh caudal, pectoral and pelvic girdles, right humerus and proximal ends of ulna and radius, femora, left tibia, fibulae, and tarsi; skull not attached to postcranial skeleton.
Paratypes. —UCMP 40209: skull with closely joined lower jaw, miss- ing left postorbital cheek region and posterior half of left mandible.
UCMP 40210: partial, articulated postcranial skeleton preserved in three small segments: 1) a series of seven postaxial cervical and dorsal vertebrae with ribs, essentially complete pectoral girdle, and proximal ends of humeri; 2) series of six vertebrae that includes the last two presacrals, two sacrals with ribs, and the first two caudals, and pelvis; and 3) portion of the left hindlimb, including proximal two thirds of femur and nearly complete tibia. It is quite likely that UCMP 40209 and UCMP 40210 belong to the same individual.
Horizon and locality. — All specimens are from the Cutler Formation exposures of the Rio Puerco drainage, Rio Arriba County, north-central New Mexico. An Early Permian Wolfcampian age is generally accepted for these exposures. Although the holotype and paratypes are listed as coming from UCMP Camp quarry locality V-2814, Langston (1952: 98) notes that they were probably not found in the main bone level of the quarry, but rather as float on the slope of Loma Salazar a few feet away and presumably at or just above the quarry bone level. The Camp quarry is located in SW1/4NE1/4NE1/4 sec. 8, T. 22 N,, R. 3 E., about
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Fig. 2,—Rhiodenticulatus heatoni, paratype, UCMP 40209. Skull in A, lateral, B, dorsal, and C, ventral views. Scale = 1 cm.
1 . 1 km southeast of Arroyo de Agua. All three specimens are preserved in red, indurated concretionary nodules.
Description
Skw//. —Specimens of Rhiodenticulatus heatoni exhibit the general structural pattern seen in all captorhinids and, therefore, aside from a
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few structures, Figs. 1 , 2 eliminate the need for a detailed description of its anatomy. The skulls of the holotype UCMP 35757 and paratype UCMP 40209 have suffered little distortion, but most of the superficial features of the skulls, such as sculpturing, have been lost due to weath- ering and excessive preparation performed prior to this study. In UCMP 40209 the left postorbital region was removed in the late 1930s in an attempt to study the braincase in thin section. The extent of ossification of the appendicular and axial portions of the holotypic skeleton suggests that it is a mature individual. The skulls are triangular, with the post- orbital width being only about 80 to 82% of the midline length. The occipital margin of the skull stable is straight.
The downtumed premaxilla possesses three teeth. In the paratype UCMP 40209 (Fig. 2A) the anterior end of the right maxilla greatly overlaps the lateral surface of the maxillary process of the premaxilla, making it appear as though the third premaxillary tooth originates from the anterior end of the maxilla. Although imperfectly preserved, the premaxillary teeth obviously had the shape of sharply pointed pegs, were subequal in size, and were approximately the same size as, or even possibly slightly smaller than, the precanine maxillary teeth. An- teriorly the maxilla forms the ventral rim of the naris, gradually expands to a moderate midlength dorsal swelling, and then tapers to a posterior terminus at, or just short of, the level of the posterior margin of the orbit. The right and left maxillae of the holotype possess 1 0 and 1 1 teeth, respectively, whereas both maxillae of UCMP 40209 possess 1 1. In both skulls the third tooth forms an extremely large canine relative to any of the other marginal teeth, with a basal diameter equal to, or greater than, twice that of any of the postcanines. In the holotype the precanines are slightly larger than the largest postcanines. The post- canines exhibit a steady decrease in size posteriorly. As in the pre- maxilla, the maxillary teeth have the form of sharply pointed pegs. In neither skull is it possible to observe directly that only a single row of marginal maxillary teeth is present. Indirect evidence for a single row is present, however, in that the teeth form a straight row along the outermost margin of the jaw, the postcanines exhibit a steady decrease in size, and there does not appear to be sufficient space for an additional tooth row on the alveolar shelf of the maxilla.
The lacrimal is unusual in being very broad. The ratio of its height (measured at the level of the dorsalmost expansion of the maxilla) to length (measured as the shortest distance between the orbit and naris) is about .65 in the holotype and about .73 in UCMP 40209. There is a correspondingly narrower lateral exposure of the prefrontal as a result of the expanded height of the lacrimal. The prefrontal is also very long and extends anteriorly along the dorsal margin of the lacrimal to a level that is about 90 and 84% of the distance between the orbit and the naris in the holotype and UCMP 40209, respectively. A long ventral
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process of the prefrontal can be seen in the holotype extending along the medial margin of the lacrimal on the anterior orbital rim. The prefrontal and postfrontal are separated by only a small lateral process of the frontal on the dorsal rim of the orbit. The frontals have a long, narrow rectangular outline. Measured from the level of their orbital contribution, the length of the anterior portion of the frontal is almost one and one half times that of the posterior portion. The pineal opening in both skulls is large and positioned anterior of the midlength of the union of the parietals. The supratemporals are not preserved in either skull. The presence of the postparietal is indicated only in the holotype and then only as an impression of its ventral surface; its suture with the parietal is therefore uncertain. The anterior ends of the right jugals of both skulls appear to wedge between the lacrimal and maxilla, rather than forming the step-like sutural encroachment onto the lateral surface of the dorsal margin of the maxilla seen in other captorhinids (Heaton, 1979). This is undoubtedly due to imperfect preservation, however, inasmuch as the standard condition is present on the nonfigured left side of the holotypic skull. The quadratojugal has the outline of a long, low rectangle, with the length exceeding the height by about four times.
Description of the palate is limited by the attached jaws. As in all captorhinids there is no ectopterygoid, and the rectangular palatine probably extends posteriorly to the subtemporal fossa. The presence of a medial jugal process cannot be determined. The denticle fields of the palate are preserved only in the paratype UCMP 40209. There is a scattering of denticles along the posterior border of the transverse flange of the pterygoid. There are also two faint, denticle bearing ridges; one extends along the medial border of the palatal ramus of the pter- ygoid, and a second extends obliquely anterolaterally across the palatal ramus of the pterygoid and onto the palatine. The three columns of irregularly arranged denticles converge toward the basicranial articu- lation. Denticles also appear to be present on the parasphenoid.
The braincases of the holotype and UCMP 40209 are exposed in ventral and occipital views and, though poorly preserved for the most part, do not appear to exhibit any noteworthy differences from the standard captorhinid construction. Both stapes of the holotype and the right of UCMP 40209 are exposed in ventral view and are well enough preserved to deserve comment. Though the footplates are not fully exposed, they appear to conform closely to those of Ecocaptorhinus (Heaton, 1979) and Captorhinus (Fox and Bowman, 1966). It has the form of a broadly oval disk that thins toward its periphery. The disk is drawn out posterolaterally into a cone-like structure, with the apex being smoothly continuous with the columella. The cross-sectional shape of columella, which remains unchanged throughout its short length, is that of a mediolaterally flattened blade having a vertical height
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Berman and Reisz— Permian Captorhinid Reptiles
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about three times its horizontal width. A large stapedial foramen pierces the proximal end of the columella at a slightly anteromedial angle from the vertical. Occipital view of the holotypic skull (not drawn) clearly reveals the dorsal process of the left stapes just distal to the stapedial foramen. It is very narrow, tapers to a point distally, and curves slightly medially.
The mandibles of both skulls are visible in partial lateral view and in ventral view; their sutural pattern and shape show no deviation from those of other captorhinids. The posterior ends of the mandibles are too damaged to determine whether or not a retroarticular process was present. Dentary teeth are visible only in the holotype, but unfortu- nately only the anterior half of the series is visible, and these are only partially exposed. The first three teeth exhibit a marked increase in size posteriorly, with the third tooth probably being the largest of the entire series. On the basis of basal diameter, the fourth and fifth teeth are slightly smaller than the third, whereas the sixth appears to be equal to the third in size. The seventh and eighth decrease further in size, as undoubtedly does the remaining unexposed portion of the series. It is estimated that the dentary of the holotype held 14 or 15 teeth.
Postcranial skeleton. — Whereas the skull of Rhiodenticulatus heatoni exhibits notable differences from those of other captorhinids, the op- posite appears to be true of the postcranial skeleton; this is not unex- pected inasmuch as this characterizes the history of captorhinids (Hea- ton and Reisz, 1980).
The holotype appears to possess a complete, articulated vertebral column from the axis to the sixth caudal vertebra (Fig. 3). Unfortu- nately, the column is exposed only in ventral view, and small segments of the series are hidden by the pectoral and pelvic girdles. Despite this, it can be safely estimated that the entire presacral column consisted of 25 vertebrae. The centra are slightly pinched laterally, and except for what is believed to be the axial centrum the ventral midlines are still broadly rounded in transverse section; the axial centrum has a distinct keel-like ventral midline. The wing-like transverse processes exhibit a gradual reduction in their lateral extent posteriorly in the column. The ventral surface of the processes slope anteroventrally, and the lateral width narrows as the processes extend to the anterior rim of the cen- trum. Both ends of the centra are slightly beveled to accommodate the intercentra, giving them a slightly keystone appearance in lateral view. The intercentra are variably displaced dorsally into the notochordal canals of the centra, where attempts to fully expose them would result in damage to the centra. As a result, many of the intercentra appear to be absent, whereas those that are partially exposed vary in size and have a lozenge-shaped outline. The first chevron occurs between cau- dals three and four.
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The string of seven postaxial cervical and dorsal vertebrae of UCMP 40210 are exposed in dorsal view only (Fig. 4 A) and undoubtedly include postaxial cervicals. Although poorly preserved, the neural arch- es exhibit the swollen appearance so typical of captorhinids. The neural spines are barely developed and appear as mere nubbins. The zyg- apophyses are widely spaced from the midline, giving the neural arches the typical lateral expansion of captorhinids. The transverse processes extend laterally beyond the zygapophyses. Only the badly weathered neural arches are exposed in the UCMP 40210 vertebral series which includes the second to last presacral to the second caudal (not figured), and they reveal no important differences from the far anterior presacral of the same specimen.
The ribs of the holotype and paratype UCMP 40210 are moderately well preserved, but the expansion of the heads is rarely visible, and the shafts frequently appear as narrow rods. The heads of the postaxial cervical ribs appear to be holocephalous and articulate in part with the intercentra. The rib shafts of the cervicals of UCMP 40210 are ex- panded into blade-like structures, whereas the more posterior rib shafts of the holotype are subcircular in cross-section. The ribs of the anterior half of the presacral column are more strongly curved posteroventrally than those of the posterior half The sacral ribs are straight, thick, and greatly expanded distally. The anterior caudal ribs of the holotype are fused to the centra, curve strongly posteriorly, are thicker than the presacral ribs, and quickly decrease in length more posteriorly in the column.
The greater portions of the pectoral girdles are preserved in both the holotype and paratype UCMP 40210, and together they exhibit most of the important features of this structure (Figs. 3A, B, 4B). The head of the interclavicle is roughly diamond-shaped, and the long, thin stem is nearly complete in UCMP 40210, missing only a small part of the distal end. The ventral plates of the clavicles are not complete, but impressions on the interclavicles indicate that they were broad and met medially; there is also no indication of a prominent, thumb-like posterior process diverging from the main body of the ventral plate as has been described in Labidosaurus (Williston, 1917) and Captorhinus (Holmes, 1977). The narrow dorsal stem is directed abruptly dorsally
Fig. 3.~Rhiodenticulatus heatoni, holotype, UCMP 35757. A, ventral view of postcranial skeleton, B, right lateral view of shoulder region, and C, lateral view of left hindlimb. Abbreviations: as, astragalus; ax, axis; cal, calcaneum; cl, clavicle; cor, coracoid; cr, caudal rib; cth, cleithrum; f, femur; fi, fibula; h, humerus; id, interclavicle; Ic, lateral centrale; of, obturator foramen; r, radius; sc, scapula; sr, sacral rib; t, tibia; u, ulna; 2, 4, 5, distal tarsals; iii, iv, v, metatarsals. Scales = 1 cm.
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Fig. A.—Rhiodenticulatus heatoni, paratypc, UCMP 40210. A, dorsal view of series of seven far anterior presacral vertebrae with ribs, and B, ventral view of pectoral girdle with proximal ends of humeri preserved in a single nodule.
at nearly a right angle to the ventral plate. As in Captorhinus (Holmes, 1977), a distinct, posteriorly directed flange-like expansion of the ven- tral half of the dorsal stem for the clavicular deltoid muscle is clearly seen in the holotype. What may be a portion of the cleithrum is present on the distal end of the dorsal stem of the right clavicle of the holotype. There are no visible sutural divisions of the endochondral portion of the pectoral girdle. The scapular blade curves dorsally rather abruptly from the essentially horizontal coracoid plate. The anterior and pos- terior margins of the scapular blade are essentially straight and parallel to each other except for the anterodorsal comer being broadly curved. The anterior coracoid portion expands a short distance anteriorly be- yond the scapular blade as a smoothly rounded plate. A coracoid fo- ramen located ventromedially to the anterior buttress of the glenoid and a supraglenoid foramen on the posterior margin of the lateral surface of the scapular blade just above the supraglenoid buttress are clearly visible in the holotype and UCMP 40210.
Essentially all that is visible of the pelves of the holotype and UCMP 40210 is the worn ventral surface of the puboischiadic plate (Fig. 3 A); the less complete pelvis of UCMP 40210 is not figured. In both spec- imens osssification along the puboischiadic suture appears to be com- plete in that there are no open spaces. The sutural division between
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the pubis and ischium is barely discemable in the holotype. The anterior border of the puboischiadic plate is moderately concave. The ischium is slightly longer and narrower than the pubis. A short distance from the ventral rim of the acetabulum the pubis is perforated by the ob- turator foramen.
The humerus is best represented in the holotype (Fig. 3A, B). It is poorly preserved, but exhibits the same general configuration as those of Captorhinus and Eocaptorhinus except that the shaft and distal head have a more slender appearance. Its length, about 1.8 cm, is approx- imately 90% of that of the femur. All other forelimb elements are either too incomplete to comment on or are absent. The hindlimb and pes are preserved only in the holotype (Fig. 3A, C). The preservation of the femora allows recognition of only some of the major features of this element. Except in being considerably more slender, particularly the shaft, the femur is very similar to that of Captorhinus. It is about 2.0 cm long, has a minimum shaft diameter of about 1.3 mm, and a maximum width of the distal head of 4.3 mm. The head appears rather massive, with a well developed intertrochanteric fossa. The popliteal area is a smooth, broadly concave depression. Though the internal trochanter is well developed, there appears to no distinct step or notch between it and the head. The tibiae and fibulae of the holotype are present, but only those of the left limb are well preserved. The tibia and fibula appear to be identical to those of Captorhinus except in being noticeably more slender. In typical primitive reptilian fashion the tibia is much shorter, 1 1.0 mm, than the femur, roughly 55% of its length. The mediolateral width of the massive proximal end is about 45% of the length, whereas anteroposterior width of the distal end is about 36% of the length; the narrowest mediolateral width of the shaft is about 0.8 mm. A deep groove divides the anterior face and articular surface of the expanded proximal end; the groove is bounded medially by a prominent cnemial crest. The lateral margin of the tibia is bowed slightly medially away from the fibula. The left fibula is about 12.3 mm long and the mediolaterally expanded proximal and distal ends are about 2.5 and 3.5 mm wide, respectively; the narrowest mediola- teral width of the shaft is about 1.1 mm. The medial margin of the fibula is strongly concave and the lateral margin only very slightly convex, giving it the appearance of being bowed laterally away from the tibia.
The tarsi of the holotype are well ossified. The right, exposed in ventral view (Fig. 3A), is nearly complete, missing only the first distal tarsal, whereas the left is represented only by the dorsally exposed calcaneum and astragalus (Fig. 3C). The tarsal elements conform closely to the pattern seen in Captorhinus (Peabody, 1951) except for two apparent deviations; the fourth distal tarsal is relatively smaller and
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the fifth which is relatively larger than in Captorhinus. The typical pattern in primitive reptiles is for the fourth distal to be considerably larger than the other distal tarsals. In Rhiodenticulatus, however, the fourth distal is roughly equal in size to the fifth. The extreme proximal ends of the third, fourth, and fifth metatarsals are all that remains of the rest of the right pes.
Discussion
Placement of Rhiodenticulatus heatoni within the Captorhinidae of the suborder Captorhinomorpha is unquestionable. It should be made clear, however, that we follow Heaton (1979), Gaffney and McKenna (1979), Reisz (1980), and Heaton and Reisz (in press) in the assignment of genera in the two recognized captorhinomorph families, the Early Pennsylvanian to Early Permian Protorothyrididae (=Romeriidae of many authors) and Early to Late Permian Captorhinidae. The capto- rhinids are differentiated from the protorothyridids by their low, wide, massive skull, hooked premaxillae, loss of tabulars and ectopterygoids, fully ossified paroccipital processes, stoutly built postcranial skeleton, 25 presacral vertebrae with swollen neural arches and low neural spines, absence of cleithra, thumb-like process on the ventral plate of clavicle, short stoutly built limbs, absence of a supinator process of humerus, and wide manus and pes. Presently, about 1 4 genera of captorhinids are recognized. Among these, however, only four genera, Romeria, Protocaptorhinus, Eocaptorhinus, and Labidosaurus, could conceiv- ably be confused with Rhiodenticulatus, because they possess single- rowed, marginal dentitions.
Clark and Carroll (1973) and Heaton (1979) presented nearly iden- tical phylogenies in which the series of successively later occurring, single-tooth-rowed captorhinids Romeria, Protocaptorhinus, and Eocaptorhinus forms a single, continuous, phylogenetic lineage de- picting transitional morphological stages that links the protorothyridids with the later occurring, multiple-tooth-rowed Captorhinus. Among the captorhinids with multiple-rowed marginal dentitions, Captorhinus is the only genus known in great detail and is also generally accepted as the most primitive. Labidosaurus, the least understood of the single- tooth-rowed captorhinids, is not included in Heaton’s (1979) phylo- genetic scheme, but is depicted in Clark and Caroll’s (1973) phyloge- netic tree as the end member of an offshoot from Protocaptorhinus. Gaffney and McKenna (1979:7) criticized the systematic methodology used by Clark and Carroll, and Heaton as being “stratophenetic” {sensu Gingerich, 1976) in which “similar morphologies are arranged strati- graphically and connected using usually implicit rather than explicit criteria, to form what are interpreted as ancestor-descendant lineages.”
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Gaffney and McKenna, without altering the basic phylogenies of Clark and Carroll, and Heaton, reexpressed them in the form of a cladogram and, thus, as a testable hypothesis. Our only serious reservation of their cladogram of the Captorhinidae is the position of Labidosaurus as a member of the clade containing Protocaptorhinus. As brought out be- low, Labidosaurus shares with Eocaptorhinus and Captorhinus several derived features of the skull not seen in Protocaptorhinus. Further, restudy of Labidosaurus is greatly needed before its phylogenetic rel- tionships can be accurately evaluated. Despite this, the cladogram of Gaffney and McKenna presents a reasonable understanding of the evo- lutionary relationships of the captorhinids and, along with the detailed morphological studies of the known captorhinomorphs by Carroll and Baird (1972), Clark and Carroll (1973), Heaton (1979), and Olson (1984), provides a basis for assessing the polarity of several character states of Rhiodenticulatus heatoni.
The maxillary dentitions of the holotype and paratype UCMP 40209 of Rhiodenticulatus are unique among the single-tooth-rowed capto- rhinids in possessing: 1) a single, extremely large canine whose basal diameter is equal to, or greater than, twice that of any of the postcanines; 2) 1 1 teeth; and 3) two precanines. It can also be noted that among the protorothyridids only the Pennsylvanian Cephalerpeton exhibits a sim- ilar specialization toward a greatly reduced number (16) of maxillary teeth that includes a low number (four) of precanines (Reisz and Baird, 1983). Protorothyridids typically possess, as does Romeria, a pair of prominent, subequal canines, yet their basal diameters are far less than twice that of the largest postcanines. Although a single tooth may be designated as a canine in Protocaptorhinus and Labidosaurus, it is not as prominent as either of the paired canines of Romeria. Eocaptorhinus also exhibits a single, prominent canine, and although the first through third postcanines may be noticeably shorter, their basal diameters are only slightly smaller than that of the canine. In the holotypic skull of Rhiodenticulatus, having a midline length of about 38 mm, the basal diameter of the canine is about 2.2 mm. This is larger in both absolute and relative measurements than the canines of Romeria and Proto- captorhinus, in which the basal diameters range from roughly 1.2 to 1.7 mm for skulls 50 to 53 mm in midline length. On the other hand, though the maximum basal diameter of the canines in Eocaptorhinus and Labidosaurus may be as much as 2.6 and 3.0 mm, respectively, their midline skull lengths are as much as two and four times greater than that of Rhiodenticulatus.
Previous authors (Clark and Carroll, 1973; Heaton, 1979) have noted that there is a general reduction in the number of maxillary teeth in successively later occurring, single-tooth-rowed captorhinids. Approx-
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imate maxillary tooth counts for Protorothyris, Romeria, Protocapto- rhinus, Labidosaurus, mid Eocaptorhinus are 24-30, 22-23, 18-22, 14- 18, and 17-22, respectively. The maximum of 11 maxillary teeth in Rhiodenticulatus can only be interpreted as a unique derived character. Probably related to this trend is the unique occurrence in Rhiodentic- ulatus of only two precanines. Protorothyridids typically possess five precanines, but as many as seven or eight have been described in Paleothyris (Carroll, 1969). A further slight reduction in the number of precanines occurs in the successively later occurring captorhinids; Romeria prima possesses six precanines, R. texana, Protocaptorhinus, and Labidosaurus four or five, Eocaptorhinus three or occasionally four, and Captorhinus three or rarely four.
The lacrimal of Rhiodenticulatus may be unique among all capto- rhinomorphs in having an unusually large height to length ratio. The height was measured at the level of the dorsalmost expansion of the maxilla, whereas the length was taken as the shortest distance between the orbit and naris. Despite the small errors expected in making these sorts of measurements, the height to length ratios of .65 and .73 for the holotype UCMP 35757 and paratype UCMP 40209, respectively, are considerably greater than those of other single-tooth-rowed cap- torhinids, which range from about .25 to .40. In the protorothyridids Paleothyris and Protorothyris, the lacrimals are very long and narrow, and have a height to length ratio of about .17. The fact that in Rhio- denticulatus the ratio is smaller for the larger holotype than for the paratype, suggests that the ratio decreases somewhat with growth or increase in size. This notion is reinforced in Romeria texana, where the ratios for an adult and juvenile described by Clark and Carroll (1973) are .27 and .40, respectively.
The extreme anterior extent of the prefrontal along the dorsal margin of the lacrimal in Rhiodenticulatus also sets it apart from all other captorhinids. In the holotype UCMP 35757 and paratype UCMP 40209 the prefrontal extends anteriorly to a level that is 90 and 84% of the distance from the orbit to the naris, respectively, whereas in other captorhinids and in Protorothyris this measurement ranges from ap- proximately 43 to 58%. It might be suspected that the greater anterior extension of the prefrontal in Rhiodenticulatus is due to removal, either as a result of weathering or mechanical preparation, of that portion of the nasal overlying its anterior end. In Eocaptorhinus, for example, where additional exposure of the prefrontal could conceivably increase its preorbital length by as much as 28% (Heaton, 1979), the anterior extension of the prefrontal would increase from about 44 to 56% of the distance between the orbit and naris. As pointed out by Heaton (1979), in Clark and Carroll’s (1973) illustration and reconstruction of
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Berman and Reisz— Permian Captorhinid Reptiles
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the holotype of Romeria texana the prefrontals appear unusually long due to the exposure of that portion of their anterior ends normally overlapped by the nasals. For this reason we used Heaton’s (1979) reconstruction of Romeria texana in calculating the relative anterior extension of the prefrontal. In the holotype and paratype UCMP 40209 of Rhiodenticulatus both pairs of prefrontals have identical lateral ex- posure configurations, strongly suggesting that they have not been sig- nificantly distorted in this way by weathering or excessive preparation.
Rhiodenticulatus exhibits several shared derived features with other advanced single-tooth-rowed captorhinids. Its possession of only three premaxillary teeth is considered advanced among the captorhinids in view of the general trend within the captorhinomorphs toward reduc- tion in the number of premaxillary teeth. Protorothyridids typically have five or six premaxillary teeth, although Protorothyris archeri ap- pears to have four and Cephalerpeton only three (Reisz and Baird, 1983). Premaxillary tooth counts for Romeria prima, R. texana, Pro- tocaptorhinus, Labidosaurus, and Eocaptorhinus are 4, 5, 4 or 5, 3, and 4 or 5, respectively. The premaxilla of Captorhinus typically pos- sesses four teeth and rarely three or five. Rhiodenticulatus is also similar to the more derived captorhinids Labidosaurus, Eocaptorhinus, and Captorhinus in having a long, narrow, rectangular quadratojugal in which the longitudinal length exceeds by almost four times the height, and the dorsal margin tends to be straight. As pointed out by Heaton (1979), in the more primitive Romeria and Protocaptorhinus the dorsal margin of the quadratojugal tends to be more convex. Heaton also noted that in the reconstruction of Romeria prima by Clark and Carroll (1973) this feature is erroneously exaggerated and is actually not sig- nificantly different from that of R. texana and Protocaptorhinus. More notable, however, is the shorter length of the quadratojugals of Romeria and Protocaptorhinus, so that the length exceeds the height by no more than two and one half times. The quadratojugals of the protorothyridids tend to be more like those of the more primitive captorhinids. The straight occipital margin of the skull table of Rhiodenticulatus, seen also in Protocaptorhinus, is a derived feature with respect to the bi- lateral parietal embayment of the occipital margin of Romeria and the protorothyridids. On the other hand, Rhiodenticulatus is viewed as primitive with respect to the median embayment of the occipital mar- gins of Labidosaurus, Eocaptorhinus, and Captorhinus.
Rhiodenticulatus exhibits at least two characters that link it with the more primitive captorhinids Romeria and Protocaptorhinus, and ex- clude it from the more advanced Labidosaurus, Eocaptorhinus, and Captorhinus. It has been noted by several authors (Clark and Carroll, 1973; Heaton, 1979) that in the evolution of the captorhinids there is
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a marked trend toward relative widening of the postorbital region of the skull. In Labidosaurus, Eocaptorhinus, and Captorhinus, the post- orbital lateral expansion of the skull becomes so pronounced that the lateral margin of the skull in dorsal view is noticeably concave, whereas in Romeria, Protocaptorhinus, and Rhiodenticulatus it is essentially straight. Labidosaurus, Eocaptorhinus, and Captorhinus are advanced over Romeria, Protocaptorhinus, and Rhiodenticulatus in exhibiting the shared derived feature of blunt (rather than sharply pointed) post- canine maxillary teeth (Olson, 1984).
Finally, there is one unique feature of Rhiodenticulatus with respect to all other single-tooth-rowed captorhinids which on first consider- ation seems unquestionably primitive, its possession of small premax- illary teeth of subequal size. In all captorhinids the premaxillary teeth exhibit a steady but dramatic increase in size anteriorly, with the an- terior teeth reaching sizes equal to, or greater than, the maxillary canine. Though Rhiodenticulatus is like its protorothyridid predecessors in this character, implying a primitive state, the alternative interpretation that it represents an evolutionary reversal is argued below.
On the basis of the above character state analysis we conclude that the most plausible relationship of Rhiodenticulatus heatoni to other captorhinids is that depicted by the cladogram of Fig. 5 in which it is the primitive sister taxon to Labidosaurus, Eocaptorhinus, and Cap- torhinus (plus all other multiple-tooth-rowed forms). We recognize, however, that the cladogram possesses a few weaknesses. First, several of the nodes are defined by only a single character. Second, there are at least two notable contradictions between the cladogram and the character state analysis presented. Perhaps the most obvious is the possession by Rhiodenticulatus of small, subequal premaxillary teeth. The cladogram requires that this character be interpreted as the result of a secondary reduction in tooth size, or an evolutionary reversal, rather than more simply, as our character analysis implies, a primitive character. The likelihood that such an event occurred, however, seems very reasonable in light of the several derived modifications of the dentition of Rhiodenticulatus noted: 1) a single, extremely large canine, 2) reduction of the maxillary dentition to 1 1 teeth, 3) reduction in the number of precanines to two, and 4) reduction of the premaxillary dentition to three teeth. Of these, the first three are judged unique to Rhiodenticulatus among the single-tooth-rowed captorhinids, whereas the last also occurs in Labidosaurus. It should be noted here, however, that in our opinion it seems quite likely that the reduction in the number of premaxillary teeth to three in Rhiodenticulatus and Labidosaurus was achieved independently given the otherwise marked differences between their dentitions. A second possible inconsistency between the
Romeria Protocaptorhinus Rhiodenticulatus Labidosaurus Eocaptorhinus Captorhinus
19
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Berman and Reisz— Permian Captorhinid Reptiles
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VOL. 55
placement of Rhiodenticulatus in the cladogram and our character state analysis concerns the unique derived features of its dentition. If, as suggested above, the extremely large, single canine, 1 1 maxillary teeth, and two precanines of Rhiodenticulatus represent the most advanced stages of general trends within the single-tooth-rowed captorhinids, then it could be argued that these features indicate an advanced sister taxon relationship with Labidosaurus and Eocaptorhinus as well. This interpretation is rejected, however, in favor of the alternative argument that these unique features of the dentition of Rhiodenticulatus, as well as its relatively small, few premaxillary teeth, probably reflect an ad- aptation to a specialized diet not present in the other single-tooth- rowed captorhinids.
Other New Mexico Captorhinids
In recent years the authors have collected additional captorhinid remains from the Lower Permian Cutler, Abo, and Sangre de Cristo formations at widely scattered localities in northern and central New Mexico. Although these specimens, as well as the two crushed and incomplete skulls referred to ''Puercosaurus obtusidens^^ by Williston (1916), are too poorly preserved to be assigned safely to an existing taxon or made the basis of a new one, they permit the recognition of at least three possible morphotypes, one each from the Cutler, Abo, and Sangre de Cristo formations. These specmiens are, therefore, im- portant as indicators of the diversity and spatial range of the capto- rhinids in the Lower Permian of New Mexico.
Indeterminate Cutler Captorhinid
All the indeterminate captorhinid specimens from Cutler Formation of the Rio Puerco drainage, Rio Arriba County, in the north-central part of the state are considered together as though pertaining to a single form distinct from Rhiodenticulatus heatoni of the same area. This is done despite the fact that the indeterminate specimens exhibit some differences from each other. It is realized that future discoveries may indicate that the differences between them may be due to either the presence of more than one undescribed species, or distinct growth stages of the same species, or both. If conspecificity is being masked by onto- genetic growth stages, then it is also conceivable that one or more of the indeterminate Cutler specimens may prove to be conspecific or congeneric with R. heatoni. This possibility is given some support by the presence in a few of the unassigned Cutler specimens of at least one feature considered derived in R. heatoni, the single, greatly enlarged canine. The unassigned Cutler specimens include:
FMNH 745, two crushed and very incomplete skulls referred to '' Puercosaurus ob- tusidens” by Williston (1916), who illustrated only one, the same skull shown here in
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Berman and Reisz— Permian Captorhinid Reptiles
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Fig. 6.—'‘'Puercosaurus obtusidens'' Williston (1916). A, dorsal, and B, ventral views of referred skull FMNH 745. C, lateral view of holotypic dentary FMNH 743. Abbrevia- tions: d, dentary; f, frontal; j, jugal; m, maxilla; pf, postfrontal; po, postorbital; prf, prefrontal; pt, pterygoid; qj, quadratojugal; sq, squamosal. Scale = 1 cm.
Fig. 6A, B. Their exact locality is unknown, and according to Williston (1916) they were found by Mr. Miller in 1911 on the Rio Puerco a few miles below Arroyo de Agua. The holotypic left dentary of “P. obtusidens'" (Fig. 6C) is too incomplete to assign to the Captorhinidae with reasonable certainty.
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Fig. 7.— Indeterminate captorhinid from the Cutler Formation. A, lateral view of partial left maxilla, B, lateral view of posterior portion of left dentary, and C, lateral and dorsal views of anterior portion of right dentary of CM 28592. D, partial skull CM 28591 showing mainly paired frontals in dorsal view, dentaries in ventral view, and small portion of left maxilla in both medial and lateral views. Abbreviations: d, dentary; f, frontal; m, maxilla. Scale = 1 cm.
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CM 28591, a partial skull (Fig. 7C, D); CM 28589, fourteen dorsal vertebrae, most of which are articulated in strings of two or three, and associated fragments of ribs and appendicular elements (Fig. 8A). These vertebrae are indistinguishable from those of R. heatoni, as are those of most captorhinids, but are included here because they were found in very close proximity to CM 28591 in NEV4SWV4NEy4 sec. 5, T. 22 N., R. 3 E. about 1.5 km northeast of Arroyo de Agua.
CM 28592, partial left maxilla (Fig. 7A), small portion of both dentaries (Fig. 7B), presacral vertebra, and left humerus (Fig. 8C). These elements undoubtedly belong to a single individual and were collected in N^ASW'ASE^A sec. 8, T. 22 N., R. 3 E. about 1.6 km southeast of Arroyo de Agua.
The left premaxilla of the figured skull of FMNH 745 (Fig. 6) appears to have held four teeth as Williston (1916) described; this estimate takes into account an unoccupied space. The premaxillary teeth, as in Rhiodenticulatus, are very small relative to the pre- and postcanines of the maxilla. Accounting for spaces, the maxilla of FMNH 745 held approximately 13 to 15 teeth, including two or possibly three preca- nines, one extremely large canine, and 10 or 11 postcanines that de- crease gradually in size posteriorly. As in Rhiodenticulatus, the basal diameter of the canine is about twice that of any of the postcanines. The dentition of the partial left maxilla of CM 28592 (Fig. 7 A) is considerably different, however, in that the canine is relatively smaller when compared to the postcanines, and the third or posteriormost precanine is nearly as large as the canine, producing a double canine appearance. A segment of the right maxilla of CM 28591 (Fig. 7D) shows the canine as dominating the postcanines in size, though not as greatly as in Rhiodenticulatus. The maxillary dentitions of FMNH 745, CM 28591, and CM 28592 are single rowed, and the teeth appear as simple, sharply pointed pegs except for a slight, posterior curvature of the tips. The frontals of FMNH 745 and CM 28591 (Figs. 6B, 7D) are complete, and their very narrow contribution to the orbital rim is clearly discemable. As in Rhiodenticulatus, the portion of frontal an- terior to its contribution to the orbital rim is considerably larger than that which is posterior. In FMNH 745 the pineal foramen appears to be more centrally positioned along the median parietal suture than in Rhiodenticulatus. The dentary dentition is well preserved in CM 2859 1 except for most of the teeth lacking their tips; the more complete right dentary is estimated to have held about 18 teeth. The first tooth is extremely small in typical captorhinid fashion, the second and third are subequal in size and much larger than the others of the series, and the following teeth do not exhibit an obvious size pattern except for the last three being greatly reduced. The anterior seven teeth preserved on the fragment of right dentary of CM 28592 (Fig. 1C) exhibit the same size relationships as in CM 28591. In contrast, the first five teeth of the left dentary of the FMNH 745 are of subequal, moderate size. The dentary teeth also have the form of simple, sharply pointed pegs.
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Fig. 8,— Indeterminate captorhinid from Cutler Formation, Presacral vertebrae of A, CM 28589, and B, CM 28592, C, dorsal proximal and distal ventral views of left humerus CM 28592. Scale = 1 cm.
The presacral vertebrae of CM 28589 and CM 28592 (Fig. 8 A, B) are alike and as far as comparisons will allow like those of Rhioden- ticulatus. The neural spine is small, triangular in lateral view, and distinctly set off from the neural arch, which has the expected swollen appearance. The zygapophyses extend slightly beyond the lateral mar- gins of the centra, and their articular facets are essentially horizontal. There is no evidence of a suture between the neural arch and centrum. The transverse process is positioned on the anterodorsal quadrant of the lateral surface of the centrum. In lateral view the process is a thin, ridge-like structure whose base extends anteroventrally to the centrum
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Fig. 9, —Indeterminate captorhinid CM 41707 from the Abo formation. A, medial view of left maxilla, B, lateral view of left jugal, C, dorsal view of distal half of right femur, and D, distal ventral view of left humerus. Scale = 1 cm.
rim. In anterior view its lateral projection diminishes as it extends to the centrum rim, giving it a wing-like appearance. The ends of the centra are beveled slightly so as to give them a slightly keystone ap- pearance in lateral view. The lateral surfaces of the centra are mod- erately concave in horizontal section, producing a spool-shaped ap- pearance. The only clearly visible intercentrum is seen in the vertebra of CM 28592 (Fig. 8B); it has a low, narrowly triangular outline in lateral view and a crescent- shaped outline in anterior view.
The only appendicular element of the indeterminate specimens from the Cutler Formation worthy of description is the well preserved left humerus of CM 28592 (Fig. 8C). It differs from those of Eocaptorhinus and Captorhinus (Holmes, 1977) mainly in having a more gracile form, but in this feature is also like that of Rhiodenticulatus. The proximal and distal ends are relatively narrower, and the entepicondyle extends far more distally beyond the radial condyle than in Eocaptorhinus or Captorhinus.
Indeterminate Abo Captorhinid
A second possible New Mexico captorhinid form for which there is insufficient morphological information to assign to either an existing or a new taxon is based on a single specimen, CM 41707, collected from the Abo Formation about 20 km northeast of Socorro in the central part of the state in SEV4NEV4Wy4 of sec. 14, T. 2 S., R. 3 E. CM
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41707 consists of disarticulated elements of the skull and postcranial skeleton of an individual that are randomly associated and densely concentrated in a small, strongly indurated, red concretion. Only those elements close to the outer surface of the concretion were prepared and include: a left maxilla, left jugal, anterior half of the right mandible, a presacral vertebra, ribs, greater part of the left humerus, distal half of right femur, and several unidentified fragments. Only a few of the above elements are figured here (Fig. 9). The left maxilla (Fig. 9A), although poorly preserved, retains an accurate outline of its dentition, which consists of 14 teeth and at least one unoccupied space; in this feature it is like the maxilla of the indeterminate Cutler specimen FMNH 745. As in Rhiodenticulatus, there are two moderate sized precanines. Al- though the third tooth is the largest of the series and should be con- sidered a canine, the fourth tooth is nearly as large, giving CM 41707 a distinctly double canine appearance like that in the partial left maxilla of the indeterminate Cutler specimen CM 28592. The basal diameter of neither canine of CM 41707, however, exceeds that of the largest postcanine as greatly as does the single, enlarged canine of Rhioden- ticulatus. The teeth gradually decrease in size from the first canine to the seventh tooth; this is followed first by four somewhat larger, sub- equal teeth and then by the last three and smallest teeth of the series. The jugal (Fig. 9B) is like that of other captorhinids. A smooth flange on the dorsal margin of the posterior plate clearly indicates the position of the overlaping postorbital, and the spike-like projection on the pos- terior margin marks the point of separation between the jugal-squa- mosal and jugal-quadratojugal contacts.
The anterior half of the right mandible (not figured) is exposed in lateral and dorsal view, and the first 1 6 teeth are present, though many are represented by only their bases. As in the captorhinids Eocapto- rhinus and Captorhinus, the first tooth is extremely small, the second moderate sized, and the third is greatly enlarged and dominates the entire series, having a basal diameter of about 2 mm and a height of about 5 mm. The fourth tooth is the second largest of the series, with a basal diameter of about 1.5 mm and an estimated height of 2.5 mm, whereas the fifth is greatly reduced and about equal to the second in size. Teeth 6, 7, and 8 are of subequal, moderate size, the larger ninth tooth appears to have been about the size of the third tooth, and the remaining seven teeth steadily decrease in size posteriorly. All the dentary teeth appear to have the form of simple pointed pegs and are aligned in a single row. As in Eocaptorhinus and Captorhinus, the first three teeth lean obliquely forward and the fourth is nearly vertical. The one partial vertebra appears to be typical of captorhinids. The left humerus of CM 41707 (Fig. 9D) is nearly complete, missing only a
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B
Fig. 10. —Indeterminate captorhinid from the Sangre de Cristo Formation. Lateral and ventral views of maxillary fragments A, CM 28594, and B, CM 28595. Scale = 1 cm.
portion of its proximal end. The shaft is more slender and the entepi- condyle possibly less expanded than those of Eocaptorhinus or Cap- torhinus. The entepicondyle extends distally only slightly beyond the radial condyle. In contrast, the distal half of the right femur (Fig. 9C) is very stoutly constructed, especially in comparison with Eocaptor- hinus and Captorhinus.
Indeterminate Sangre de Cristo Captorhinid
Numerous fragments of captorhind maxillae and dentaries have been collected from the Lower Permian Sangre de Cristo Formation ap- proximately 50 km southeast of Santa Fe in the northeastern part of the state in NEV4 sec. 36, T. 14 N., R. 13 E. Two of the maxillary fragments are figured (Fig. 10), and they clearly indicate that the cap- torhinid from this locality had at least two rows of teeth. In one (CM 28594, Fig. lOA) the broken edges of a second row of teeth can be seen lateral to the posterior end of the main row. Medial wear facets of the teeth, which are more evident on the other fragment (CM 28595, Fig. lOB), give them the same blunt, peg-like outlines seen in Eocaptorhinus and multiple-tooth-rowed forms such as Captorhinus. These two fea- tures of the dentition indicate clearly that the Sangre de Cristo cap- torhinid remains are of a distinct and more advanced taxon than the other representatives of the family in New Mexico.
Acknowledgments
Support for this research was provided by grants from the New Mexico Bureau of Mines and Mineral Resources (to D.S B. and R.R.R.), the M, Graham Netting Research Fund through a grant from the Cordelia Scaife May Charitable Trust (to D.S B.), and the Natural Sciences and Engineering Research Council of Canada (to R.R.R.). We are grateful to Ms. Diane Scott, Erindale Campus, University of Toronto, for the preparation of the specimens and the drawing of the illustrations. We are obliged to the University of California, Berkeley, and the Field Museum of Natural History for the loan of spec- imens. Special thanks are extended to Dr. Wann Langston, Jr., of the University of Texas, Austin, for bringing these specimens to our attention and expediting their loan to us. Thanks are due Drs. Robert L. Carroll of McGill University, Montreal, and E. C. Olson of the University of California, Berkeley, for critical reading of the manuscript.
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Literature Cited
Carroll, R. L, 1969. A Middle Pennsylvanian captorhinomorph, and the interrela- tionships of primitive reptiles. J. Paleont., 43:151-170.
Carroll, R. L., and D. Baird. 1972. Carboniferous stem-reptiles of the Family Ro- meriidae. Bull. Mus. Comp. ZooL, Harvard Univ., 143:321-363.
Clark, J., and R. L. Carroll. 1973. Romeriid reptiles from the Lower Permian. Bull. Mus. Comp. Zool., Harvard Univ., 144:353-407.
Fox, R. C., AND M. C. Bowman. 1966. Osteology and relationships of Captorhinus aquti (Cope) (Reptilia: Captorhinomorpha). Univ. Kansas Paleont. Contrib., Ver- tebrata, 11:1-79.
Gaffney, E. S., AND M. C. McKenna. 1979. A Late Permian captorhinid from Rho- desia, American Mus, Novit, 2688:1-15.
Gingerich, P, D. 1976. Cranial anatomy and evolution of early Tertiary Plesiadapidae (Mammalia, Primates). Univ. Michigan, Papers Paleont., 15:1-141.
Heaton, M. J. 1979. Cranial anatomy of primitive captorhinid reptiles from the Later Pennsylvanian and Early Permian OWahoma and Texas. Bull. Oklahoma Geol. Survey, 127:1-84.
Heaton, M. J., and R. R. Reisz, 1980. A skeletal reconstruction of the Early Permian captorhinid reptile Eocatorhinus laticeps (Williston). J. Paleont., 54:136-143.
. In press. The interrelationships of captorhinomorph reptiles. Canadian J. Earth
Sci.
Holmes, R. 1977. The osteology and musculature of the pectoral limb of small cap- torhinids. J. Morph., 152:101-140.
Langston, W., Jr. 1952. Permian vertebrates of New Mexico. Unpublished Ph.D. dissert., Univ. California, Berkeley, 212 pp.
. 1953. Permian amphibians of New Mexico, Univ, California Publ., Geol. Sci.,
Bull., 29:349-416.
Olson, E. C. 1 984. The taxonomic status and morphology of Pleuristion brachyocoelus Case; referred to Protocaptorhinus pricei Clark and Carroll (Reptilia: Captorhino- morpha). J. Paleont., 58:1282-1295.
Peabody, F. E. 1951. The origin of the astragalus of reptiles. Evolution, 5:339-344.
Reisz, R. R. 1 980. A protorothyridid captorhinomorph reptile from the Lower Permian of Oklahoma. Royal Ontario Mus., Life Sciences Contrib., 121:1-16.
Reisz, R. R., and D. Baird. 1983. Captorhinomorph “stem” reptiles from the Penn- sylvanian coal-swamp deposit of Linton, Ohio. Ann, Carnegie Mus., 52:393-411.
Williston, S. W. 1916. The osteology of some American Permian vertebrates, 11. Univ. Chicago, Walker Mus. Contrib., 1:165-192.
. 1917. Labidosaurus Cope, a lower Permian cotylosaur reptile from Texas. J.
Geol., 25:309-321.
NPi
ISSN 0097-4463
ANNALS
0/ CARNEGIE MUSEUM
CARNEGIE MUSEUM OF NATURAL HISTORY
4400 FORBES AVENUE ® PITTSBURGH, PENNSYLVANIA 15213 VOLUME 55 23 MAY 1986 ARTICLE 2
DESCRIPTION OF THE LOWER JAW OF STEGOSAURUS (REPTILIA, ORNITHISCHIA)
David S Berman
Associate Curator, Section of Vertebrate Fossils John S. McIntosh^
Research Associate, Section of Vertebrate Fossils
Abstract
Description of a well preserved lower jaw of Stegosaurus reveals previously unknown structural details. Although comparisons between the lower jaw of Stegosaurus and those of other stegosaurs are greatly limited, they strongly suggest a close comformity. The lower Jaw of Stegosaurus exhibits the most primitive grade of organization among the omithischians with one noticeable exception, the retention of an intercoronoid in cer- atopsians.
Introduction
During the past several years there has been a renewed effort by the Carnegie Museum of Natural History to prepare the remaining dinosaur materials collected by that institution over 60 years ago (1909-1922) at what is now Dinosaur National Monument. During the routine preparation of one of the blocks, field no. 83, an almost complete, excellently preserved left mandible of Stegosaurus was unexpectedly discovered. Earl Douglass, who was in charge of the quarrying oper- ations, had identified the contents of field no. 83 as ""Dinosaur, vertebra, rib, etc.,” and preparation of the block was, therefore, subsequently
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postponed because it was thought to contain little of importance. Al- though Stegosaurus is one of the most common of the Morrison di- nosaurs, skull material is extremely rare. To date, only one complete skull and lower jaw have been found, that of the type specimen of Stegosaurus stenops (USNM 4934) from the Felch quarry (YPM quarry 1; see Ostrom and McIntosh, 1966) at the entrance of Garden Park, about 13 km north of Canyon City, Colorado. It was originally de- scribed briefly by Marsh (1887) and later in much greater detail by Gilmore (1914). Several braincases of Stegosaurus have been found, but the anterior half of a right dentary (USNM 4935) from the “stego- saur” quarry (YPM quarry 13; see Ostrom and McIntosh, 1966), about 6.5 km east of Como Bluff, Wyoming, is otherwise the only other mandible known. The mandible described herein lacks only the pre- dentary and teeth, and is noteworthy in exhibiting structural details heretofore unknown in other specimens.
The following abbreviations are used to refer to repositories of specimens: CM, Car- negie Museum of Natural History; USNM, National Museum of Natural History; and YPM, Yale Peabody Museum.
Systematic Paleontology Class Reptilia
Order Omithischia Seeley, 1888 Suborder Stegosauria Marsh, 1877 Family Stegosauridae Marsh, 1880 Genus Stegosaurus Marsh, 1877 Stegosaurus sp.
Specimen.— CM 41681, left mandible lacking predentary and den- tition.
Horizon.— Morrison Formation, Late Jurassic.
Locality.— 'Dinosnnr National Monument (originally called the Car- negie quarry), about 10 km north of Jensen, Uintah County, Utah. CM 41681 was found under the fourteeth cervical vertebra of the type skeleton of Apatosaurus louisae Holland, CM 30 1 8, at the co-ordinates F to G- 5 4 on the quarry map published by Gilmore (1936).
Collector.— EdcrX Douglass and party, 1910.
Assignment.— T>oscrrpXions and illustrations by Marsh (1887) and Gilmore ( 1 9 1 4) of the holotypic lower jaw of Stegosaurus stenops USNM 4934, although containing several errors that will be brought out in the text below, and by Gilmore (1914) of the right dentary of Stego- saurus sp. USNM 4935 leave little or no doubt about the assignation of CM 41681 to this genus. It can also be noted that there exists no known dinosaur from the Late Jurassic whose lower jaw could be confused with that of Stegosaurus.
1986
Berman and McIntosh— ^'recFOS'^c/Rt/^ Jaw
31
Description
As Figs. 1-3 indicate, the left mandible CM 4 1 68 1 is complete except for the absence of the one unpaired element of the lower jaw, the median predentary, and the dentition. The mandible is well preserved and the sutures delineating the seven elements that comprise it, except those of the articular, are clearly discemable. The mandible is narrowly com- pressed in transverse section. In lateral or medial view the mandible is shallow, exhibiting a gradual dorsal and ventral expansion posteriorly from the symphysis to a maximum depth at about three-fifths its length, then gradually narrows at about the same rate posteriorly to the end of the retroarticular process. Whereas the expansion of the dorsal mar- gin is somewhat angular in outline, that of the ventral margin is smooth- ly convex. There is no development of a distinct coronoid process, but rather there is a low coronoid eminence whose apex is formed by the surangular. The plate-like anterior end of the mandible turns abruptly medially and slightly ventrally, and with the symphysis inclined pos- teroventrally the joined mandibles would have a scoop-like appearance. It has not been possible to identify the small rectangular plate of bone adhering to the medial surface of the anterior end of the dentary; it might be part of the adjoining left dentary.
The sutures of the lateral surface of the mandible (Fig. 1) are, for the most part, clearly defined and need little comment. The articular-sur- angular suture is not evident, and a portion of the angular-surangular suture is restored on the basis of impression. The dentary, surangular, and angular border a very large, horizontally elongate, oval mandibular foramen. A very thin, short strip of the splenial is visible on the mid- ventral margin of the mandible. Three shallow, parallel grooves occur shortly posterior of the point where the anterior end of the dentary curves medially. Each deepens slightly as it extends 1 to 2 cm antero- dorsally from the inferior jaw margin and ends at a small foramen that pierces the dentary at a very low angle. Three relatively small foramina are located near the dorsal border of the surangular; two closely spaced smaller ones are located a considerable distance posterior to a larger one. They undoubtedly penetrate to the medial side of the surangular. Gallon (1974) has described similar foramina in the omithischian di- nosaur Hypsilophodon and suggested that they may have transmitted cutaneous branches of the inferior alveolar nerve as in modem lizards (Oelrich, 1956).
In medial view (Fig. 2) all seven elements of the mandible are visible. The dentary is the largest of the jaw elements. Its upper half, above the Meckelian canal, is remarkably thick, having a triangular cross- sectional outline with the apex directed medially to form a dorsal, shelf-like surface (Fig. 3). Posteriorly much of the medial edge of the
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Berman and McIntosh— (75 Jaw
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dorsal shelf is occupied by a series of 20 closely spaced alveoli. The remaining anterior portion of the medial edge of the shelf takes the form of a thin, knife-like ridge that continues to the symphysis. The slightly concave dorsal shelf of the dentary faces dorsomedially at its posterior end, but because of a gradual twisting as it is followed an- teriorly, the shelf faces directly dorsally just beyond its midlength and then anteriorly at the symphysis. Along most of the length of the dorsal shelf, mainly at the level of the alveoli, is a series of about nine foramina of varied sizes, which might be interpreted as mental foramina. The posteriormost pair of foramina are located at the anterior end of a prominent, 3 cm long groove that deepens anteriorly. The orientation of the alveolar series suggests that the teeth were directed mainly me- dially and only slightly dorsally. Directly below the series of alveoli is a series of foramina; they are apparently arranged so that each foramen lies opposite an alveolus, but they do not occur the entire length of the alveolar series.
The Meckelian canal of the dentary is exposed medially as it emerges from beneath the anterior margin of the splenial, becoming progres- sively shallower as it extends nearly to the symphysis. The large, sub- rectangular, plate-like splenial has a sinuous, overlapping suture with the dentary anteriorly. There is a deep, narrow emargination of the anterior border of the splenial at the level of the Meckelian canal. The splenial-prearticular contact is also very sinuous; a moderate sized inframandibular foramen is located on this suture. The posterodorsal comer of the splenial appears to enter narrowly the rim of the adductor fossa. Its ventral margin contacts the dentary along the ventral edge of the mandible, whereas more posteriorly it wraps a short distance around the ventral edge to contact the angular on the lateral surface of the mandible. A large foramen of unknown function penetrates the splenial in a posterodorsal direction near its anteroventral border. What may be a large foramen is located near the middorsal margin of the splenial. A small portion of the angular wraps around the ventral edge of the mandible and is visible in medial view. Although a sutural contact between the prearticular and articular cannot be discerned, the prear- ticular undoubtedly formed the greater part of the ventromedial border and the articular the posterior border of the large, oval adductor fossa. The coronoid is roughly triangular in medial view, with its longer side forming a substantial portion of the dorsal margin of the mandible. It also contributes greatly to the anteromedial border of the adductor
Fig. 1.— Photograph and illustration of left mandible Stegosaurus CM 41681 in lateral view. Abbreviations: A, angular; C, coronoid; D, dentary; SA, surangular; SP, splenial.
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Berman and McIntosh— ^rE^GO^^c/i? c/5 Jaw
35
fossa. There is no indication that an intercoronoid was present. Just posterior to the coronoid there is a small, triangular, thickened area on the medial surface of the surangular apex of the coronoid eminence. This thickened area undoubtedly marks the site of concentrated at- tachment of jaw adductor musculature. The slightly concave articular surface of the articular is subcircular in outline. At its posteromedial border is a stout, dorsally directed triangular process. The retroarticular process is short and bluntly rounded.
Comparison and Discussion
The Stegosaurus mandible CM 41681 exhibits three features of par- ticular interest that are not noted or shown in earlier descriptions or illustrations of the genus (Marsh, 1887; Gilmore, 1 9 14): a large, external mandibular foramen; presence of a large coronoid bone; and a very weakly developed coronoid process with the apex formed by the sur- angular.
Several orders of reptiles have small fenestrae on the external surface of the mandible, but the presence of a large external mandibular fo- ramen located at the intersection of the dentary, angular, and suran- gular, as in Stegosaurus, is rare. Such a foramen is found in crocodiles, prosauropods, theropods, and some thecodonts, but among omithis- chians it has been reported only in the primitive omithopod Fabro- saurus (Thulbom, 1970) and the stegosaur Huayangosaurus (Dong et al., 1982; Zhou, 1983). The external mandibular foramen of Fabro- saurus and CM 41681 are very similar except that in the former the dentary forms the anterior half or more of its margin. Huayangosaurus, from the Middle Jurassic of China, is the only stegosaur in which an external mandibular foramen has previously been described (Dong et al., 1982; Zhou, 1983). Although the formaen in Huayangosaurus is relatively smaller than that in CM 4 1 68 1 , it has the same general outline shape and precisely the same position and relationships to the sur- rounding dentary, surangular, and angular. Owen (1863) did not find evidence of the mandibular foramen in the primitive armored omith- ischian Scelidosaurus, and recent preparation using modem chemical techniques has clearly indicated the absence of the foramen (Charig, 1979). Scelidosaurus has often been considered a stegosaur, but Charig (1979:126), who is currently restudying this animal, has remarked that
Fig. 2.— Photograph and illustration of left mandible of Stegosaurus CM 4168 1 in dorsal view. Abbreviations: AL, alveoli; AR, articular; C, coronoid; D, dentary; PA, preartic- ular; SA, surangular.
1986
Berman and McIntosh— Jaw
37
“It has often been regarded as an ancestral stegosaur; it might be an ancestral ankylosaur; it could be neither.”
Gilmore (1914) presumed that a separate coronoid bone was present in Stegosaurus, but was unable to verify this on the basis of the then only known complete jaw of the genus, that of the holotype of S. stenops (USNM 4934). It is, therefore, gratifying that relatively large coronoid is clearly evident in CM 41681. Owen (1863) stated that the lower jaw of SceUdosaurus includes a coronoid, but did not describe it or indicate it in his figures (pis. 46, 47). Although the presence of a coronoid has not been demonstrated in other stegosaurs, there is no basis for sug- gesting its absence. On the contrary, there is increasing evidence that a coronoid element was present in all omithischians except the had- rosaurs. It has been explicitly reported in the omithopods Hypsilopho- don (Gallon, 1974), Camptosaurus (Gilmore, 1909), Iguanodon (Dol- lo, 1883), and Ouranosaurus (Taquet, 1976), in the ceratopsians Protoceratops, Montanoceratops, Centrosaurus, and Triceratops (Brown and Schlaikjer, 1940), in the Edmontonia (Gilmore, 1930;
Russell, 1940) (=Panoplosaurus of Coombs, 1978), and in the pachy- cephalosaur Stegoceras (Gilmore, 1924). The presence of a second coronoid element, the intercoronoid, has also been found by Brown and Schlaikjer (1940) in the Protoceratopsidae and Ceratopsidae. The intercoronoid, which apparently occurs in all saurischians (for example, Plateosaurus, Brachiosaurus, Camarasaurus, AUosaurus, Tyrannosau- rus, and others), has not been reported in any other omithischian group, and we are unable to find any evidence for its presence in CM 41681.
Among the omithischians the coronoid of the ankylosaur Edmon- tonia (Gilmore, 1930; Russell, 1940) resembles most closely that of Stegosaurus in shape, size, and position, but important differences are evident. The anterior extension of the coronoid in Edmontonia lies medial to the posterior end of the tooth row, whereas in Stegosaurus it lies lateral to the tooth row. In Edmontonia, as well as in other ankylosaurs, and the pachycephalosaur Stegoceras (Gilmore, 1 924) the surangular is expanded dorsally into a broadly rounded but prominent coronoid process; only in Edmontonia, however, is the coronoid clearly shown to curve upward onto the leading edge of the process. In Stego- saurus, on the other hand, the dorsal margin of the mandible rises gradually and evenly to the apex of the very low coronoid eminence whose apex is formed by the surangular. With the exception of Fabro-
Fig. 3. —Photograph and illustration of left mandible of Stegosaurus CM 4 1 68 1 in medial view. Abbreviations: A, angular; C, coronoid; D, dentary; PA, prearticular; SA, suran- gular; SP, spleniaL
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saurus (Thulbom, 1970), omithopods possess a large, shaft-like cor- onoid process formed mainly by the dentary, that extends dorsally from the main body of the bone; the summit of the process may even curve anteriorly. There is a suboval coronoid on the medial surface of the process except in hadrosaurs, which appear to lack this element (Lull and Wright, 1942; Ostrom, 1961). In Fabrosaurus the dentary forms the apex of a low coronoid eminence; there is no distinct coronoid process. Within the omithopods there is an apparent trend toward reduction and loss of the coronoid. The coronoid is relatively large in the primitive Fabrosaurus and Hypsilophodon, but in advanced forms it is greatly reduced, as in Ouranosaurus, and apparently lost in had- rosaurs.
Only one difference between CM 41681 and previous descriptions of the lower jaw of Stegosaurus is possibly noteworthy. The Stegosaurus right dentary USNM 4935 possesses 23 alveoli (Gilmore, 1914), where- as that of CM 41681 has 20. Whether this should be regarded as an individual variation or as having taxonomic significance is not known. It can be noted, however, that the USNM 4935 right dentary is ap- proximately 20% larger than that of CM 41681, and the difference in their tooth counts may be related to size. What little is known about the jaws of other stegosaurs suggests that they compare closely with that of Stegosaurus. Though the descriptions of the jaw of Huayang- osaurus (Dong et al., 1982; Zhou, 1983) are very brief, the accompa- nying figures of its lateral surface suggest that it probably differs in only minor ways from that of Stegosaurus— i\ may have a slightly more prominent coronoid process, but with the apex still apparently formed by the surangular; the teeth appear to be vertically oriented; a lateral exposure of the coronoid is not indicated; and the anterior end of the mandible curves more strongly ventrally. The stegosaurs Kentrurosau- rus (Hennig, 1936), Tuojiangosaurus (Dong et al., 1983), and Chung- kingosaurus (Dong et al., 1983) are known only from incomplete den- taries. The dentary of the Kentrurosaurus resembles very closely that of Stegosaurus in the position, shape, and development of the Meck- elian canal, but the slightly larger alveoli suggest that it may have held fewer teeth. All that can be said of the dentaries of the latter two genera is that they were long and low, as in Stegosaurus.
The possession of a large external mandibular fenestra, large coro- noid, and very weakly developed coronoid process in which the apex is formed by the surangular can be justifiably interpreted as primitive omithischian features, inasmuch as they occur in a wide range of the- codonts. The above comparisons, therefore, indicate that the lower jaw of Stegosaurus, as is probably true of other stegosaurs, exhibits the most primitive grade of organization among the omithischians with
1986
Berman and McIntosh -■5'r£'GD5’^i7R (75 Jaw
39
one conspicuous exception, the retention of the intercoronoid in the ceratopsians.
Acknowledgments
Support for this project was provided by a grant from the M. Graham Netting Research Fund through a grant from the Cordelia Scaife May Charitable Trust (to D.S B.). We are especially grateful to Ms. Amy Henrici for the difficult task of preparing the specimen and to Ms. Nancy Perkins for drawing the illustrations; both are of Carnegie Museum of Natural History. We also wish to thank Drs. Peter M. Galton of the University of Bridgeport, Connecticut, John H. Ostrom of the Yale Peabody Museum of Natural History, and Mary R. Dawson of the Carnegie Museum of Natural History for critically reviewing the manuscript.
Literature Cited
Brown, B., and E. M. Schlaikjer. 1940. A new element in the ceratopsian jaw with additional notes on the mandible. Amer. Mus. Novit., 1092: l-l 3.
Charig, a. J. 1979. A new look at dinosaurs. Mayflower Books, New York, 160 pp.
Coombs, W. P., Jr. 1978. The families of the omithischian dinosaur order Ankylo- sauria. Palaeontology, 21:143-170.
Dollo, L. 1883. Quatrieme note sur les dinosauriens de Bemissart. Bull. Mus. Hist. Nat. Belgique, 11:223-252.
Dong Zm, Tang Zl, and Zhou Sw. 1982. Note on the new mid-Jurassic stegosaur from Sichuan Basin. Vertebrata Palasiatica, 20:85-87.
Dong Zm, Zhou Sw, AND Zhang Yh. 1983. The dinosaurian remains from Sichuan Basin, China. Palaeont. Sin., New C Ser., 162:1-145.
Galton, P. M. 1974. The omithischian dinosaur Hypsilophodon from the Wealden of the Isle of Wight. Bull. British Mus. (Nat. Hist.), 25:1-152.
Gilmore, C. W. 1909. Osteology of the Jurassic reptile Camptosaurus, with a revision of the species of the genus and description of two new species. Proc. U.S. Nat. Mus., 36:197-302.
— — 1914. Osteology of the armored dinosauria in the U.S. National Museum with special reference to the genus Stegosaurus. Bull. U.S. Nat. Mus., 89:1-140.
■ 1924. On Troodon validus, an omithopod dinosaur from the Belly River Cre- taceous of Alberta, Canada. Bull. Univ. Alberta, 1:1-43.
. 1930. On dinosaurian reptiles from the Two Medicine Formation of Montana.
Proc. U.S. Nat. Mus., 77:1-39.
1936. Osteology of Apatosaurus with special reference to specimens in the
Carnegie Museum. Mem. Carnegie Mus., 1 1:175-300.
Hennig, E. 1936. Ein dentale von Kentrurosaurus aethiopicus Hennig. Palaeontogr., Suppl. 7, 1:309-312.
Lull, R. S., and N. E. Wright. 1942. Hadrosaurian dinosaurs of North America. Spec. Pap. Geol. Soc. Amer., 40:1-242.
Marsh, O. C. 1887. Principal characters of American Jursassic dinosaurs. Part IX, the skull and dermal armor of Stegosaurus. Amer, J. Sci., 43:413-417.
Oelrich, T. M, 1956. The anatomy of the head of Ctenosaura pectinata (Iguanidae). Misc. Pubis. Mus. Zook, Univ. Michigan, 94:1-122,
Ostrom, J, H. 1961. Cranial morphology of the hadrosaurian dinosaurs of North America. Bull. Amer. Mus. Nat. Hist., 122:1-186.
Ostrom, J. H,, and J. S. McIntosh. 1 966. Marsh’s dinosaurs. Yale Univ. Press, 388 pp.
Owen, R. 1863. A monograph of the fossil Reptilia of the Liassic formations. Part II. Palaeontogr. Soc. Monogr., 13: 1-26.
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Russell, L. S. 1940. Edmontonia rugosidens (Gilmore), an armoured dinosaur from the Belly River series of Alberta. Univ. Toronto Stud. Geol. Ser., 43:1-28.
Taquet, P. 1976. Geologie et Paleontologie du Gisemement de gadoufaoua (Aptien du Niger). Cahiers de Paleontologie, 191 pp.
Thulborn, R. a. 1970. The skull of Fabrosaurus australis, a Triassic omithischian dinosaur. Palaeontology, 13:416-432.
Zhou Sw. 1983. A nearly complete skeleton of stegosaur from Middle Jurassic of Dashanpu, Zigong, Sichuan. J. Chengdu College Geol., SuppL 1:15-26.
^ ISSN 0097-4463
ANNALS
of CARNEGIE MUSEUM
CARNEGIE MUSEUM OF NATURAL HISTORY 4400 FORBES AVENUE » PITTSBURGH, PENNSYLVANIA 15213 VOLUME 55 23 MAY 1986 ARTICLE 3
BIOCHEMICAL AND MORPHOLOGICAL DIFFERENTIATION IN SPANISH AND MOROCCAN POPULATIONS OF DISCOGLOSSUS AND THE DESCRIPTION OF A NEW SPECIES FROM SOUTHERN SPAIN (AMPHIBIA, ANURA, DISCOGLOSSIDAE)
Stephen D. Busack^’^
Research Associate, Section of Amphibians and Reptiles
Abstract
Biochemical and morphological divergence among Moroccan and Iberian populations suggests that populations of Discoglossus inhabiting these regions are not conspecific. Northern Moroccan Discoglossus are assigned to D. pictus’, Discoglossus galganoi Capula et al, 1985 inhabits the Iberian peninsula to the north of the Guadalquivir River basin, and the Discoglossus population residing to the south of the Guadalquivir River basin on Iberia is described as a new species. An evolutionary scenario for Iberian and Mo- roccan populations, consistent with the biochemical and morphological data, is inferred from the physiogeographic history of the region.
Introduction
Until recently only two species of Discoglossus were thought to in- habit Europe. Spanish, French, and Sicilian populations were called D. pictus, and populations inhabiting Corsica, Sardinia, Elba, and Monte Argentario, Italy, were called D. sardus (Knoepffler, 1961a, 1961Z?, 1 962). Recent electrophoretic examinations of Discoglossus have, how-
^ Address: Fellow in Herpetology, California San Francisco, California 94118.
^ Present address: Department of Genetics j Urbana, Illinois 61801.
Submitted 1 May 1985.
Academy of Sciences, Golden Gate Park,
41
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Fig. I.™ Localities of examined specimens of Discoglossus galganoi (squares), D. pictus (triangles), and D. jeanneae (circle). Dotted line traces the presumed northern limit of D. jeanneae', see Discussion for further explanation.
ever, revealed two additional species, D. montalentii from Corsica (Lanza et al., 1984) and D. galganoi from north of the Guadalquivir River basin in Spain and Portugal (Capula et aL, 1985).
My interest in Discoglossus was initially focused on the amount of genetic differentiation that accumulated between European and African populations after the formation, 5-7 million years ago, of the Strait of Gibraltar (Busack, 1986). Because my electrophoretic study revealed substantial genetic differentiation between Spanish and Moroccan sam- pies, the morphologies of these populations were also compared. Bio- chemical and morphological data indicate that Iberian populations to the north and to the south of the Guadalquivir River basin are not conspecific, and that neither is conspecific with populations inhabiting northern Morocco. The data supporting this conclusion are presented in the following pages, together with a description of a second species from Spain, and a hypothetical reconstruction of the evolutionary his- tory of Iberian and Moroccan forms of Discoglossus.
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Busack— Spanish and Moroccan Discoglossus
43
Table l.~~ Protein systems" examined by electrophoresis; enzymes are arranged by En- zyme Commission number.
|
Protein (abbreviation) |
Enzyme Commission number |
Electro- phoretic conditions |
|
Albumin (Ab) |
B |
|
|
(Oxidoreductases) |
||
|
Alcohol dehydrogenase (Adh) |
1.1. 1.1 |
A |
|
Glycerol- 3 -phosphate dehydrogenase (Gpd) |
1.1. 1.8 |
D |
|
L-Lactate dehydrogenase (Ldh) |
1.1.1.27 |
F |
|
Malate dehydrogenase (Mdh) |
1.1.1.37 |
F |
|
Malate dehydrogenase (Me) |
1.1.1.40 |
F |
|
Isocitrate dehydrogenase (led) |
1.1.1.42 |
E |
|
Phosphogluconate dehydrogenase (Pgd) |
1.1.1.44 |
E |
|
Glutamate dehydrogenase (Glud) |
L4.1.3 |
D |
|
Superoxide dismutase (Sod) |
1.15.1.1 |
D |
|
(Transferases) |
||
|
Aspartate aminotransferase (Aat) |
2.6.1. 1 |
D |
|
Hexokinase (Hk) |
2.7.1. 1 |
G |
|
Creatine kinase (Ck) |
2.7.3.2 |
G |
|
(Hydrolases) |
||
|
Esterase (Est) |
3.1. 1.1 |
B |
|
Esterase-D (Est-D) |
3.1. 1.1 |
B |
|
Acid phosphatase (Aep) |
3.1.3.2 |
G |
|
Fmetose-bisphosphatase (Hdp) |
3.1.3.11 |
D |
|
N - Acetyl-Beta-glucosaminidase (Hex) |
3.2.1.30 |
G |
|
Dipeptidase I, L-Leucyl-L-Alanine (La) |
3.4.11 |
B |
|
Dipeptidase III, L-Leucylglycyl-glycine (Lgg) |
3.4.11 |
C |
|
Dipeptidase IV, L-Phenylalanyl-L-Proline (Pap) |
3.4.11 |
B |
|
Adenosine deaminase (Ada) |
3.5.4.4 |
A |
|
(Lyases) |
||
|
Fmctose-bisphosphate-aldolase (Aid) |
4.1.2.13 |
H |
|
Aconitate hydratase (Aeon) |
4.2.1.3 |
E |
|
(Isomerases) |
||
|
Mannose-6-phosphate isomerase (Mpi) |
5.3.1.8 |
E |
|
Glucose-6-phosphate isomerase (Gpi) |
5.3.1.9 |
F |
|
Phosphoglucomutase (Pgm) |
5.4.2.2 |
E |
A = Histidine, pH 7.8 gel and electrode buffer (Harris and Hopkinson, 1976), 150v/3h.
B = LiOH A + B, pH 8.2 gel and LiOH A, pH 8.1 electrode buffer (Selander et al., 1971), 300v/3h.
C = Poulik, pH 8.7 gel and borate, pH 8.2 electrode buffer (Selander et al., 1971),
250v/3h.
D = Tris citrate II, pH 8.0 gel and electrode buffer (Selander et al., 1971), 130v/4h.
E = Tris citrate II, pH 8.0 + NADP gel and tris citrate II, pH 8.0 electrode buffer (Selander et al., 1971), 130v/4h.
F = Tris citrate III, pH 7.0 gel and electrode buffer (Ayala et al., 1972), 180v/3h.
G == Tris citrate III, pH 7.0 + 1 5% glycerine gel and tris citrate III, pH 7.0 electrode buffer (Ayala et al., 1972), 180v/3h.
H ^ Tris citrate III, pH 7.0 + NAD + 2-mercaptoethanol gel and tiis citrate III, pH 7.0 electrode buffer (Ayala et al., 1972), 180v/3h.
44
Annals of Carnegie Museum
VOL. 55
Materials and Methods
Electrophoresis. —OnQ individual was collected from north of the Guadalquivir River basin in Avila Province, Spain, at San Martin del Pimpollar (40®22'N, 5®03'W; Fig. 1, square C). Seven specimens were collected south of the Quadalquivir River basin in Cadiz Province, Spain (Fig. 1, circle), near the towns of Facinas (36®08'N, 5®42'W, 5 specimens) and Benalup de Sidonia (36®20'N, 5M9'W, 2 specimens). Specimens collected in Tetouan Prefecture at Chechaouene (35®10'N, 5®16'W; Fig. 1, open triangle; 5 spec- imens) and in the vicinity of Tleta Tarhremt (35®47'N, 5®28'W; Fig. 1, closed triangle; 5 specimens) represent northern Moroccan populations.
Specimens were sacrificed in the field and samples of heart and liver were removed, frozen, and stored in liquid nitrogen (- 196®C). In the laboratory, tissues were transferred to a freezer (-76®C) until used in electrophoresis two to 12 months later. Tissue samples were pooled for each animal. Proteins were separated electrophoretically in horizontal starch gels (11.5% hydrolyzed starch, Sigma Chemical Co.) and localized by standard histochemical staining procedures (Ayala et al., 1972; Harris and Hopkinson, 1976; Selander et al., 1971; Table 1). Genetic interpretations of allozymic data were based on criteria developed by Selander et al. (1971). Multiple loci within a protein system were numbered with “ 1 ” designating the most anodally migrating set of allelic products. Alleles of a locus were lettered, with “a” representing the most anodally migrating product. Data resulting from the electrophoretic analysis are summarized in Table 2.
Two methods were used to analyze genetic relationships among populations of Dis- coglossus. The first distinguishes patterns of allele distribution among populations that occur as a result of chance association from those that occur too frequently to be chance phenomena. If all patterns of allele distribution are equally probable, the probability, P, that a particular pattern will occur is 1/S„ where S„ is the sum of the number of possible patterns of allele distribution. In the case of the three populations of Discoglossus being compared, S„ = 7 for three items combined three at a time, two at a time, and one at a time. The probability of observing a specific pattern of allele distribution, r, two or more times is given by summing the terms of the binomial expansion:
b(r) = (7)/>'(l - Pr-,
where r is the number of replications seen for a given pattern of allele distribution, m is the total number of alleles in the data set, and is the number of possible combi- nations of m alleles taken r at a time (Straney, 1980; Patton and Smith, 1981).
The second method is the computation of estimates of, and standard errors for, the unbiased minimum genetic distance {D\ Nei, 1978, 1971, respectively) between Disco- glossus populations. Allele frequency data (Table 2) were used directly for the compu- tation of genetic distances and their standard errors.
Morphology.— CB.puhL et al. (1985:tables 2, 4) published comprehensive tables of mor- phological measurements for D. galganoi. I have used the data in these tables to elucidate the morphological features of D. galganoi and to make direct assessments of morpho- logical differentiation between D. galganoi and other taxa.
Spanish specimens I personally examined were representative of the same Cadiz Prov- ince populations as my electrophoretic samples, but collected earlier (between 1969 and 1972). Moroccan specimens included those used for electrophoresis as well as additional material from the same sites. Straight-line measurements of snout-urostyle (SUL), snout (anterior comer of the eye to the tip of the snout), head (posterior angle of the jaw to the tip of the snout), eye (horizontal diameter from posterior comer to anterior comer), tibia, femur, hand (proximal aspect of the central metacarpal tubercle to the tip of the third digit), and foot (proximal aspect of the metatarsal tubercle to the tip of the third digit) lengths were taken to 0.1 mm with dial calipers. Head width (angle of jaws).
1986
Busack— Spanish and Moroccan Discoglossus
45
interorbital (between the anterior comer of the eyes), and intemarial (center to center) distances were also recorded.
Frogs are sometimes sexually dimorphic in many characters and sexes were analyzed separately. The effect of having limited numbers of individuals of each size representing each sex of each population was minimized by converting each character measurement to natural logarithms; the variance of In-transformed data estimates intrinsic variability and is unaffected by size (Lewontin, 1966; Moriarty, 1977). Transformed data repre- senting each character were subjected to covariance analysis in which snout-urostyle length was selected as the independent variable. Although allometry is correctly assessed only from the study of growth of an individual, I used individuals of different sizes from a population to obtain estimates of allometric coefficients. Identification of dissimilarities in the allometric growth influence by using transformed data is acceptable for comparing populations (Thorpe, 1976).
Linear regression analysis, in which the measurement data were left untransformed, was then performed for variables demonstrating significant differences in allometric growth. For ease of presentation and interpretation, only the resulting slope and intercept values are reported in Table 4. Significance levels for all statistical tests were set (a priori) at 0.05 and probabilities are those for committing a Type I error in a two-tailed test.
Results
Biochemical Comparisons
Aatl, Aat2, Acp2, Ada, Est-D, Hdp, Me, Pgm, and Sod were mon-
omorphically expressed among all three populations I examined. Table 2 summarizes the distribution of allozymes at the 25 polymorphic loci I was able to score unambiguously.
Populations of Discoglossus residing to the north and to the south of the Guadalquivir River basin in Spain and those inhabiting northern Morocco are genetically well differentiated. Fifteen of 87 alleles iden- tified among these Discoglossus samples are shared among all popu- lations. Spanish Discoglossus residing north of the Quadalquivir River basin share 17 alleles with individuals from northern Morocco and 23 alleles with Discoglossus residing south of the Guadalquivir River basin in Spain. Populations inhabiting the area south of the Guadalquivir River basin share 29 alleles with Discoglossus inhabiting northern Mo- rocco (Table 3).
Seventy alleles, however, differentiate between individuals of Dis- coglossus from north of the Guadalquivir River basin in Spain and those from northern Morocco, 64 alleles differentiate individuals from north of the Guadalquivir River basin and those from south of the Guadalquivir River basin in Spain, and 58 alleles differentiate indi- viduals living to the south of the Guadalquivir River basin in Spain from those inhabiting northern Morocco (Tables 2 and 3). The distri- bution of these alleles contributes to a genetic distance {D) of 0.74 ± 0.18 between the Spanish sample from north of the Guadalquivir River basin and the northern Moroccan samples, 0.39 ± 0.12 between Span- ish samples residing to the north and to the south of the Guadalquivir
46
Annals of Carnegie Museum
VOL. 55
Table 2.— Genic variation within and among samples o/Discoglossus.
|
D. jeanneae |
D. galganoi |
D. pictiis |
||
|
Number of specimens |
1 |
1 |
10 |
|
|
Mean heterozygosity per lo- cus |
0.16 |
0.03 |
0.16 |
|
|
Percentage of loci polymor- phic |
38.2 |
2.9 |
44.1 |
|
|
Locus and alleles |
||||
|
Abl |
a |
0.86 |
1.00 |
0.95 |
|
b |
0.14 |
0.00 |
0.00 |
|
|
c |
0.00 |
0.00 |
0.05 |
|
|
Ab2 |
a |
1.00 |
1.00 |
0.00 |
|
b |
0.00 |
0.00 |
1.00 |
|
|
Aeon |
a |
0.57 |
0.00 |
1.00 |
|
b |
0.29 |
0.00 |
0.00 |
|
|
c |
0.14 |
0.00 |
0.00 |
|
|
d |
0.00 |
1.00 |
0.00 |
|
|
Acpl |
a |
0.00 |
0.00 |
0.10 |
|
b |
1.00 |
1.00 |
0.90 |
|
|
Adh |
a |
1.00 |
1.00 |
0.80 |
|
b |
0.00 |
0.00 |
0.20 |
|
|
Aid |
a |
0.00 |
1.00 |
0.00 |
|
b |
0.86 |
0,00 |
1.00 |
|
|
c |
0.14 |
0.00 |
0.00 |
|
|
Ck |
a |
0.57 |
0.00 |
0.10 |
|
b |
0.00 |
0.00 |
0.10 |
|
|
c |
0.43 |
0.00 |
0.75 |
|
|
d |
0.00 |
0.00 |
0.05 |
|
|
e |
0.00 |
1.00 |
0.00 |
|
|
Est |
a |
0.00 |
0.00 |
0.20 |
|
b |
0.00 |
0.00 |
0.40 |
|
|
c |
0.29 |
0.00 |
0,40 |
|
|
d |
0.14 |
0.00 |
0.00 |
|
|
e |
0.57 |
1.00 |
0.00 |
|
|
Glud |
a |
0.00 |
1.00 |
0.20 |
|
b |
1.00 |
0.00 |
0.80 |
|
|
Gpd |
a |
0.00 |
LOO |
0.00 |
|
b |
0.07 |
0.00 |
0.00 |
|
|
c |
0.93 |
0.00 |
0.50 |
|
|
d |
0.00 |
0.00 |
0.50 |
|
|
Gpi |
a |
1.00 |
1.00 |
0.90 |
|
b |
0.00 |
0,00 |
0.10 |
|
|
Hex |
a |
0.00 |
0.00 |
0.10 |
|
b |
1.00 |
1.00 |
0.00 |
|
|
c |
0.00 |
0.00 |
0.90 |
|
|
Hkl |
a |
1.00 |
1.00 |
0.00 |
|
b |
0.00 |
0.00 |
1,00 |
|
|
Hk2 |
a |
0.00 |
1.00 |
0.00 |
1986
Bus ACK— Spanish and Moroccan Discoglossus
47
Table 2.— Continued.
|
D. jeanneae |
D. galganoi |
D. pictus |
|||
|
b |
1.00 |
0.00 |
0.00 |
||
|
c |
0.00 |
0.00 |
1.00 |
||
|
Icdl |
a |
0.00 |
0.00 |
0.10 |
|
|
b |
0.86 |
0.00 |
0.85 |
||
|
c |
0.14 |
0.00 |
0.05 |
||
|
d |
0.00 |
1.00 |
0.00 |
||
|
Icd2 |
a |
0.00 |
0.00 |
1.00 |
|
|
b |
0.00 |
0.50 |
0.00 |
||
|
c |
1.00 |
0.50 |
0.00 |
||
|
La |
a |
0.00 |
0.00 |
0.10 |
|
|
b |
0.57 |
1.00 |
0.90 |
||
|
c |
0.43 |
0.00 |
0.00 |
||
|
Ldhl |
a |
1.00 |
1.00 |
0.00 |
|
|
b |
0.00 |
0.00 |
1.00 |
||
|
Ldh2 |
a |
1.00 |
0.00 |
1.00 |
|
|
b |
0.00 |
1.00 |
0.00 |
||
|
Lgg |
a |
0.00 |
0.00 |
1.00 |
|
|
b |
1.00 |
1.00 |
0.00 |
||
|
Mdhl |
a |
0.14 |
0.00 |
0.00 |
|
|
b |
0.29 |
0.00 |
0.00 |
||
|
c |
0.43 |
0.00 |
0.00 |
||
|
d |
0.14 |
0.00 |
0.00 |
||
|
e |
0.00 |
1.00 |
0.60 |
||
|
f |
0.00 |
0.00 |
0.30 |
||
|
g |
0.00 |
0.00 |
0.10 |
||
|
Mdh2 |
a |
0.71 |
0.00 |
0.90 |
|
|
b |
0.29 |
0.00 |
0.10 |
||
|
c |
0.00 |
1.00 |
0.00 |
||
|
Mpi |
a |
0.00 |
0.00 |
0.50 |
|
|
b |
0.71 |
1.00 |
0.50 |
||
|
c |
0.29 |
0.00 |
0.00 |
||
|
Pap |
a |
0.29 |
0.00 |
0.00 |
|
|
b |
0.71 |
1.00 |
0.00 |
||
|
c |
0.00 |
0.00 |
1.00 |
||
|
Pgd |
a |
0.86 |
0.00 |
0.50 |
|
|
b |
0.00 |
0.00 |
0.40 |
||
|
c |
0.14 |
0.00 |
0.10 |
||
|
d |
0.00 |
1.00 |
0.00 |
River basin, and 0.39 ± 0.12 between Spanish samples from south of the Guadalquivir River basin and those from northern Morocco.
Morphological Comparisons
Between sexes. — The 19 male and 8 female specimens in the sample of D. galganoi examined by Capula et al. (1985) do not demonstrate
48
Annals of Carnegie Museum
VOL. 55
Table 3.— Number of replications of the seven possible patterns of allele distribution present among the three populations o/Discoglossus. Patterns replicated eight times or more do not occur as a result of chance association (b < 0.05, see Materials and Methods) but only alleles shared between two or more populations are informative.
D. galganoi D. jeanneae D. pictus Number
|
X |
X |
X |
15 |
|
X |
14 |
||
|
X |
10 |
||
|
X |
24 |
||
|
X |
X |
2 |
|
|
X |
X |
8 |
|
|
X |
X |
14 |
sexual dimorphism » There were no significant differences in allometry identified by the covariance analysis.
While no significant difference was found in the distribution of SUL between 14 male and 16 female Discoglossus from south of the Gua- dalquivir River basin in Spain, the allometric relationship between SUL and foot length is significantly different between the sexes of this population (F ^ 6.61, R < 0.05). The foot length of a female with an SUL of 45 mm (an intermediate size in the sample of adults of either sex) is approximately 95% the length of the foot of an equivalent-sized male.
Five male and six female Discoglossus from northern Morocco dem- onstrated sexual dimorphism in the allometric relationship between SUL and snout length. While no significant differences were identified between male and female SUL in these samples, the relationship be- tween SUL and snout length is significantly different between sexes (F = 16.39, P ^ 0.05). At an SUL of 45 mm, the snout length of a female specimen is approximately 86% of the snout length of an equiv- alent-sized male.
Between populations.— M.^\q specimens of D. galganoi differ from male Discoglossus from northern Morocco in the allometric growth relationship between SUL and snout (F = 68.01, P c 0.05), head (F = 63,57, P ^ 0.05), tibia (F = 10.66, P < 0.05), femur (F = 16.90, P <c 0.05) and hand lengths (F = 55.07, P ^ 0.05) and in the growth re- lationships of SUL and head width (F = 5,84, P < 0.05) and SUL and intemarial distance (F = 8.05, P < 0.05). Female specimens repre- senting these populations differ in the allometric relationship between SUL and head (F = 39.56, P <c 0.05) and eye (F = 28,73, P c 0,05) lengths,
Male specimens of D. galganoi differ from males from south of the Guadalquivir River basin in Spain in the allometric growth relationship
1986
Busack— Spanish and Moroccan Discoglossus
49
Fig. 2.— Spotted (upper) and striped (lower) color phases of Discoglossus jeanneae.
50
Annals of Carnegie Museum
VOL. 55
of SUL and snout (F 224.33, P c 0.05), head (F = 54.76, P c 0.05), tibia (F = 1 1.29, P c 0.05), femur (F - 7.08, P < 0.05), and hand lengths (F ^ 45.82, P <c; 0.05), and in the allometric relationship between SUL and intemarial distance (F ^ 5.63, P < 0.05). Female specimens representing these populations differ in the allometry be- tween SUL and head length (F = 72.34, P ^ 0.05), SUL and hand length (F = 77,57, P c 0.05), and between SUL and intemarial dis- tance (F= 12.62, P <c 0.05).
Male specimens from south of the Guadalquivir River basin in Spain and those from northern Morocco differ only in the allometric growth relationship between SUL and snout length (F = 16.47, P ^ 0.05). The allometric growth relationship between SUL and all examined morphological characters is not different in female specimens drawn from these populations.
Systematic Considerations
Northern Moroccan populations. — Lanza et al. (1984) demonstrated that specimens of Discoglossus from near Barika, Algeria, are geneti- cally very similar to those from Sicily, the type locality of D. pictus (Nei’s average genetic identity = 0.93, Nei’s D = 0.07). Although Ca- pula et al. (1985:table 7) do not provide a table of allele frequencies with which I might directly compare results, they do list 12 loci that distinguished Algerian and Tunisian D. pictus from Iberian D. galga- noi. Three of these 12 loci (Ada, Aatl, and Acp2) were found to be monomorphic among populations I compared, and seven were not considered in my study. Only two of these 12 loci (Icd2 and Ldhl) distinguished between Moroccan and Iberian populations in my study. While it is possible that not all Moroccan, Algerian, Tunisian, and Sicilian populations are conspecific, at this time it is zoogeographically and systematically conservative to consider populations of Discoglossus inhabiting North Africa and Sicily D. pictus Otth, 1837.
Iberian populations residing north of the Guadalquivir Basin. —Tht albumin immunological distance obtained when specimens from Vi- lla viciosa and Arenas de San Pedro, Spain (Fig. lA, B, respectively), were compared to those representing D. pictus from Tleta Tarhremt, Morocco (Fig. 1 , closed triangle), was 1 7 units (Maxson and Szymura, 1984:249). The unbiased genetic distance (Nei’s D) between one in- dividual I collected from San Martin del Pimpollar, Spain (Fig. 1C), and D. pictus from northern Morocco was 0.74 ± 0.18. Capula et al. (1985) reported a genetic distance (Nei’s D) of 0.58 between their pooled samples representing Portugal and central Spain and those from Algeria and Tunisia.
It is apparent from genetic (Capula et al., 1985:table 9) and mor-
1986
Busack— Spanish and Moroccan Discoglossus
51
Fig. 3.— Snout length regressed on snout-urostyle length in male Discoglossus galganoi (squares), D. pictus (triangles), and D. jeanneae (circles). See Table 4 for regression coefficients.
phological comparisons (this study, Results) that Iberian populations of Discoglossus residing north of the Guadalquivir River basin are dearly differentiated from those inhabiting northern Morocco and are deserving of the species status recently ascribed to them by Capula et al. (1985). Whether or not, however, the specimens Maxson and Szy- mura (1984) and I examined biochemically (Fig. 1A-~C) actually rep- resent D. galganoi is unclear. Until data with which to further assess the taxonomic status of Iberian populations residing to the north of the Guadalquivir River basin become available, these populations are best referred to D. galganoi Capula et aL, 1985.
Iberian populations residing south of the Guadalquivir Basin.— Dis- coglossus from south of the Guadalquivir River basin in Spain are morphologically and genetically different from both Moroccan D. pictus and. D. galganoi (Tables 2, 3, and Results). I consider the extent of these differences to be representative of species level differentiation and designate the new species herewith.
Table A.— Estimates of slope (b) and intercept (a) obtained when measurements derived from various morphological features (y) were regressed againts smui-urostyie length (x) in Discoglossus galganoi, D. jeanneae, and D. pictus. Linear regression results are reported
52
Annals of Carnegie Museum
VOL. 55
f.s 1 8
a
1^:
^ § a ^
8 I
sil
G g g
g
•a ?
§ ^ S « ^ ^ K ^ "a ^ -S S
.a
Q a K ^
.-<* '*«**
S’
g o “
s-s <» ail
fit
H
^ I
II
■Si
l“S
I'
VO ^ o m o
O r4 O
+1 +1 I
O Os ^ O CN
1 I !
I I I
8^1
+1 +1 T
2 2^''
I I I
I I I
^
O VO ^
^ ^ M
+1 +1 ^
o
fNj en o ^
I I I
I I I
W W WOT
+1 +1 g
^ liT
I I I
|
CN |
»n |
en |
«ri |
m |
||||
|
o |
P-- |
o |
O •O |
O |
O |
O |
o |
|
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d |
d |
d V |
d d |
d \/ |
d |
d |
d V |
d |
|
+1 |
+1 |
FI +1 |
V |
+1 |
+1 |
+1 |
||
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VO |
\o m |
en O |
oC |
Os |
VO |
oq |
||
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'"7 |
m |
os |
o |
oo |
o |
|||
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d |
d |
o |
d ■^' |
d |
d |
VO |
d |
|
X |
||||||||
|
tn q |
a |
.—4 |
m |
m o |
||||
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VO |
d |
s |
o |
o |
m |
d |
a |
|
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d |
¥ |
d |
d |
d |
¥ |
1 |
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+1 |
1 i |
1 1 |
+1 |
+1 |
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+i |
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F' |
IT) |
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|
o |
q |
r- |
o |
q |
in |
|||
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VO |
o© |
q |
Os |
d |
q |
|||
|
d |
VO |
d |
d |
|
m O |
«n |
m o |
in |
in o |
.—1 |
||||
|
o |
o |
a\ |
O |
o |
|||||
|
d |
d |
d u |
d |
d u |
d |
d |
d y |
d |
|
|
+1 |
+1 |
J |
+1 |
+! |
V |
+1 |
+1 |
V |
+1 |
|
o |
d |
m |
m |
fT |
os |
VO |
m |
||
|
m |
1—1 |
»n |
O |
q |
O |
||||
|
d |
d |
cn |
d |
d |
r>- |
d |
en |
vd |
d |
|
m w |
W |
m |
pq |
|
WOT |
W |
“ a. |
w |
|
+' +1 . |
+1 |
+1®: |
+1 |
|
,o |
a±SE 0.55 ± 0.59 0.76 ± 0.43 0.36 ± 0.19 0.42 ± 0.44 0.97 ± 0.4!
F„ P 17.46, cO.05 20.43, <0.05 158.89, c0.05 36.11, c0.05 26.70, <0.05
Table A. — Continued.
1986
Busack— Spanish and Moroccan Discoglossus
53
|
—1 m O Vi |
>ri O |
|||||||||||||||
|
d d |
d |
|||||||||||||||
|
•3 s |
+1 +1 |
V |
\ |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
|||||
|
r- Vi o — 1 |
so" oo |
|||||||||||||||
|
3 ts |
d d |
d |
||||||||||||||
|
s. |
||||||||||||||||
|
ci |
so |
•n |
o |
m |
n |
m |
m |
|||||||||
|
o |
r-- |
o |
o |
o |
o |
o |
oo |
o |
||||||||
|
s |
d |
d V |
d |
r4 |
d V |
d |
d |
d V |
||||||||
|
1 1 |
1 |
+1 |
+1 |
+1 |
+1 |
+1 |
+1 |
|||||||||
|
§ |
os |
o |
ri q |
oo |
o |
so" |
o |
in |
XJ-" q |
|||||||
|
CN |
m |
T |
fN |
|||||||||||||
|
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54
Annals of Carnegie Museum
VOL. 55
Discoglossus jeanneae, new species
(Fig. 2)
Holotype.— Carnegie Museum of Natural History (CM) 546575 an adult female, along highway C-440, 1 5 km ESE Alcala de los Gazules, CMiz Province, Spain, 6 April 1971, Stephen D. Busack.
Paratypes (21, all from Cadiz Province, Spain).— CM 52126, female, along highway C-440, 12.9 km ESE Alcala de los Gazules, 18 October 1969; CM 52128--52129, two females, along highway C-440, 13.8 km WNW Los Barrios, 18 October 1969; CM 52475- 52476, two males, along highway C-440, 6.6 km NW Algeciras, 10 January 1970; CM 53087, male, along highway C-440, between 7.6 and 7.7 km NW Los Barrios, 12 March 1970; CM 53119, male, along highway CA-221, 21.2 km ENE Facinas, 10 April 1970; CM 53324, male, along highway C-440, 9.01 km NW Los Barrios, 18 June 1970; CM 53884a--d, 4 females, along highway C-440, between 4.0 km WNW Casas del Castano and 5.0 km WNW Los Barrios, 8 October 1970; CM 54244, female, along highway C-440, between 1 9 km ESE Alcala de los Gazules and 1 km WNW Casas del Castano, 18 November 1970; CM 54581, female, and CM 54582, male, along highway C-440, between 3.7 and 1 1.4 km WNW Los Barrios, 19 March 1971; CM 54608-54610, three females, along highway CA-P-2 1 1 2, between 12.6 and 1 4.2 km NNE Benalup de Sidonia, 2 April 1971; CM 54704, male, along highway C-440, 0.3 km WNW Casas del Castano, 16 May 1971; CM 55742-55743, two males, and CM 55744, female, along highway C-440, between 18.4 and 22.7 km WNW Los Barrios, 15 January 1972.
Diagnosis.— Discoglossus jeanneae is similar in coloration and pat- tern to D. galganoi and D. pictus, but is distinguished from them by biochemical and morphological characters. Discoglossus jeanneae and D. pictus share no alleles at six electrophoretic loci (Ab2, Hkl, Hk2, Icd2, Ldhl, and Lgg), D. jeanneae and D. galganoi share no alleles at three loci (Glud, Hk2, and Ldh2).
Male D. jeanneae have a shorter snout than males of either D. pictus or D. galganoi of similar SUL (Fig. 3, Table 4). Male D. jeanneae also have a greater head length, a shorter intemarial distance, and shorter tibia, femur, and hand lengths than male D. galganoi of comparable SUL (Table 4). Female D. jeanneae have a shorter head and hand lengths, and shorter intemarial distances, than D. galganoi of similar SUL (Table 4).
Description of holotype. — An adult female with the following mea- surements (mm): SUL 47.9, snout length 5.1, head length 10.8, head width 15.1, horizontal diameter of eye 4.1, interorbital distance 7.5, intemarial distance 3.0, tibia length 23.4, femur length 21.7, hand length 5.9, foot length 15.1. Choanae oblong and relatively large; pre- vomerine dentigerous processes in two nearly straight series, each with 1 1 teeth, located just behind the choanae and separated by a distance less than half that of the diameter of a choana. Tongue roundish, thick, scarcely free behind. Nostrils dorsal, much closer to the tip of the snout than to the eye, horizontal diameter of the eye slightly greater than the distance from the naris to the eye. Tympanum indistinct. Fingers rel-
1986
Busack— Spanish and Moroccan Discoglossus
55
Fig. 4.-- A Lower Miocene reconstruction of the Iberian peninsula and North Africa; heavily blackened areas indicate marine incursions (after lilies, 1975),
atively short, unwebbed; III, IV, II, I in order of decreasing length. Three prominent palmar tubercles, the largest at the base of finger IV, the next largest at the base of finger I, and the smallest in the center. Toes slender, very slightly webbed, IV, III, V, II, I in order of decreasing length. No subarticular tubercles, no tarsal fold; small ellipsoidal meta- tarsal tubercle. Heels overlap slightly when femora are held at right angles to the body axis. Skin of dorsum with several irregularly posi- tioned diminutive, pustules, skin of venter smooth.
Coloration ofholotype (in alcohol).— The ground color of the top of the head is citrine drab from mid-eye to snout. Beginning at mid-eye, and continuing to just below the area of front limb insertion, a dark olive patch reminiscent of a large italicized “X” appears on a ground color of a lighter shade of dark olive. A prominent (6.5 mm in length) and elongated teardrop-shaped patch of chaetura drab angles ventro- posteriorly from the posterior comer of the eye. This patch, and the ground color of the “X,” are edged in citrine drab. Small faint patches shaped as triangles oriented with the base down appear along the upper lip; a light stripe extends from the front comer of the eye through the nostril.
Colored as the ‘"X,” a lightly-homed, heart-shaped, shield extends over the central dorsum. The upper half of the shield is edged in citrine drab on a ground color of lighter dark olive; intense asymmetrical patches of dark olive line the lower “V” and the upper and outer curves of this shield. A broken dorsolateral line of citrine drab is present.
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Annals of Carnegie Museum
VOL. 55
In lateral view, an ovoid dark olive blotch appears below the dorso- lateral line and just above the area of insertion of the front limb. The interaxillary region is light deep olive blending into dark olive buff towards the venter; the dark olive buff blends into buffy brown in the region of the groin. The venter is deep olive buff and unpattemed.
The upper surfaces of the hind legs are clove brown with irregular transverse blotches of fuscous black; upper surfaces of the front legs are citrine drab with three deep olive transverse blotches.
Pattern polymorphism {in alcohol). —Spoiled individuals are most common, striped specimens are less common (Fig. 2), and occasionally an unpattemed D. jeanneae is found. Unstriped individuals exhibit either a complete or broken anteriodorsal ‘"X.” All have a teardrop eye patch, triangular blotches on the upper labium, and, in varying degrees of distinction, a stripe extending from the front comer of the eye through the nostril. A broken dorsolateral line of citrine drab also appears, at times faint, at times pronounced, in all unstriped individ- uals.
Unstriped individuals have a ground color which varies from citrine drab to light dark olive to fuscous. Coloration on the posterior portion of the dorsum varies from a pattern in which prominent spots are connected to suggest a shield-like shape to a simple pattern of widely spaced and barely discemable spots. Ventral coloration is generally unpattemed deep olive buff; some specimens, however, demonstrate olive or clove brown pigmentation of varying intensity on the lower jaw and upper pectoral region.
In the striped paratype (CM 53119), the “X” and shield symmetry are broken by a medial stripe of citrine drab. The resultant halves of the “X” and shield merge on either side of the medial stripe to form two solid regions of dark olive, one on the right, one on the left, and each between the medial stripe and the dorsolateral region. The small ovoid blotch found below the dorsolateral ridge and above the area of limb insertion in spotted individuals is continuous with the teardrop eye patch and forms a single blotch in this specimen. The eye-nostril stripes and upper labial triangles are pronounced and, along the dor- solateral region, there is a prominent citrine drab stripe extending from the eye to the groin. The outer edge of this stripe exhibits the darker coloration reminiscent of the dark olive upper and outer curves of the heart-shaped shield described for the holotype.
Etymology. — Jeanne A. Visnaw accompanied me during four months of field work in Spain and Morocco during 1982; in spite of what she learned during her first trip abroad, Jeanne again accompanied me in 1983. While the husbands and wives of graduate students often contribute substantially to the success of their spouses, few routinely sacrifice as much or give as unselfishly as my wife has. I dedicate this new Spanish frog to her.
1986
Bus ACK— Spanish and Moroccan Discoglossus
57
Discussion
Physiogeographic changes have been extensive in this area of the western Mediterranean region in the last few million years. The Betic Cordillera (an alpine ridge located on the Spanish Meseta between CMiz in the west and Alicante in the east) was subjected to folding and extensively restructured during Miocene-Pliocene. Lower Miocene Atlantic waters flowed to the Mediterranean through what is now the Guadalquivir River basin (Figs. 4-5; Berggren and Van Couvering, 1974; lilies, 1975; Le Pichon et al., 1972), Neogene sedimentation filled the younger, western portion (Cordoba-Sevilla) of the Guadalquivir Basin and the northern portion began to emerge from considerable depth at the end of the Miocene (Tjalsma, 1971:1 20-125), and Pliocene events allowed Atlantic waters to form the Strait of Gibraltar (Femix et al, 1967; Hsu, 1983; and Mantura, 1977).
Geologic events such as these have a direct effect on sexually repro- ducing organisms. The rate at which genetic divergence accumulates between populations is believed to be a function of the time those populations have been separated (Zuckerkandl and Pauling, 1965). If this is true, the more genetically differentiated two populations of ter- restrial anurans are, the longer they have been separated. Of the three patterns of allele distribution that are phylogenetically informative among these three populations of Discoglossus, D. galganoi and Mo- roccan D. pictus share only two, D. galganoi and D. jeanneae share 8, and D. jeanneae and Moroccan D. pictus share 14 (Table 3). If unbiased genetic distance units {D) are considered, Moroccan D. pictus are sep- arated from D. galganoi by 0.74 ± 0.18 units and D. jeanneae is separated from both D. galganoi and Moroccan D. pictus by 0.39 ± 0.12 units (Fig. 5). Discoglossus galganoi and Moroccan D. pictus are probably not sister species.
Males of D. galganoi are more morphologically differentiated from males of D. jeanneae and males of D. pictus than are female D. galganoi from female D. jeanneae or D. pictus. The allometric growth relation- ship between SUL and 10 morphological characters were compared among males and females of these three species and male D. galganoi differ from Moroccan D. pictus in seven such relationships, females differ in only one. Male D. galganoi differ from D. jeanneae in six allometric relationships, females differ in three. Male D. jeanneae ap- pear to be little changed from Moroccan D. pictus, however, as only one allometric relationship is clearly different; female D. jeanneae, on the other hand, demonstrate no differences among any of the 10 al- lometric growth relationships when compared with female D. pictus.
The fossil record of Discoglossus is limited. Middle Miocene remains from Beni-Mellal, Morocco, referred to the genus Discoglossus by Ver- gnaud-Grazzini (1966), have been reassigned to the extinct discoglossid
58
Annals of Carnegie Museum
VOL, 55
Fig. 5.— A mid-Miocene reconstruction of the Gibraltar area (after Femix et al., 1967), Shaded areas represent emergent land; geographic and unbiased genetic distances between sampled populations of Discoglossus are indicated.
genus Latonia by Sanchiz and Alcover {in litt.); as a result, fossil rep- resentatives of Discoglossus are unknown in North Africa. The Lower Miocene Discoglossus troschelii from Rott, Germany, was considered conspecific with D. pictus (=? galganoi) from Spain (Boulenger, 1891). Neogene specimens from Escobosa de Calatahazor (Soria), Venta del Moro (Valencia), and Alcoy (Alicante), Pliocene samples from El Ar-
1986
Busack— Spanish and Moroccan Discoglossus
59
quillo II (Teruel), and mid-Pleistocene specimens from Arganda (Ma- drid), attest to the age, persistence without morphological change, and widespread nature of D. pictus (==? galganoi) throughout prehistoric Iberia (Sanchiz, 1977a, 1911 b).
The electrophoretic and morphological data coincide well with one evolutionary scenario that may be inferred from the geographic history of the region. The ancestral stock of D. galganoi, which once populated the Spanish Meseta, and that of D. pictus-D. jeanneae, which once populated an area now known as southern Spain and northern Morocco, suffered temporal and climatic separation dating from the Lower Mio- cene. The Pliocene formation of the Strait of Gibraltar then divided ancestral D. pictus- D. jeanneae populations into two populations which evolved to become D. jeanneae and D. pictus.
Additional research is clearly necessary before we can fully under- stand the evolutionary history, distributional limits, and taxonomy of Iberian and North African Discoglossus. Until additional data becomes available, however, conservative limits for the distribution of D. jean- neae may be drawn. These would include the northern edge of the Guadalquivir River basin (Fig. 1 , dotted line), regions inundated during Miocene flooding that currently lay to the east of the headwaters of the Guadalquivir River (Fig. 1 , dotted line with question marks), and the shores of the Atlantic Ocean and the Strait of Gibraltar.
Acknowledgments
I wish to extend my gratitude to Benedetto Lanza, who provided me with a manuscript copy of the description of D. galganoi, and to D. B. Wake and M. M. Frelow of the Museum of Vertebrate Zoology, University of California, Berkeley, who provided fa- cilities, financial support, and assistance during the electrophoretic analysis. L. R. Maxson graciously provided immunological data and a copy of her and Szymura’s unpublished manuscript, C. J. McCoy and E. J. Censky generously provided sustenance and lodging during a visit to Pittsburgh, D. B, Wake, L. R, Maxson, and T. Uzzell reviewed and improved earlier versions of the manuscript, Lezlie Skeetz assisted in translating the patterns and colors of D. jeanneae into words, G. M. Christman prepared Figure 5, and Alfredo Salvador assisted with a portion of the field work in Spain. Travel funds were provided by a National Science Foundation dissertation improvement grant (DEB 81- 20868) and a National Geographic Society research grant (2600-83). Specimens in Spain were collected under authority of permits 888 (1982) and 22061 (1983) issued by the Institute Nacional para la Conservacion de la Naturaleza, Madrid. Collecting in Morocco was authorized by letter from the Embassy of Morocco to the United States, Mohamed Benjelloun, economic counsellor.
Specimens Examined
D. (Electrophoretic Analysis): SDB 1556, 1691, 1905, 1906, 1930, 1949,
1954, and 1989 at the Universidad de Leon, Spain. (Morphological Analysis): Carnegie Museum of Natural History (CM) 52125-52129, 52475-52477, 52537, 52626, 53087- 53088, 53119-53120, 53324, 53884(4 specimens), 54244, 54581-54582, 54608-54610, 54657, 54704, and 55742-55743.
D. galganoi.— {p\QcXTOp\ioreX\c Analysis): SDB 1691 at the Universidad de Leon.
60
Annals of Carnegie Museum
VOL. 55
D. (Electrophoretic Analysis): SDB 1773 (2 specimens) and SDB 1774 (3
specimens) at the Museum of Vertebrate Zoology, University of California, Berkeley (MVZ); MVZ 186124-186125 and 186132-186134. (Morphological Analysis): MVZ 186124-186134.
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Busack, S. D. 1986. Biogeographic analysis of the herpetofauna separated by the formation of the Strait of Gibraltar. National Geographic Research 2(1): 17-36.
Capula, M., G. Nascetti, B. Lanza, L. Bullini, and E. G. Crespo. 1985. Morpho- logical and genetic differentiation between the Iberian and the other west Mediter- ranean Discoglossus species (Amphibia Salientia Discoglossoidae). Monitore ZooL Ital. (N.S.), 19:69-90.
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Lanza, B., G. Nascetti, M. Capula, and L. Bullini. 1984. Genetic relationships among west Mediterranean Discoglossus with the description of a new species (Am- phibia Salientia Discoglossidae). Monitore Zool. Ital. (N. S.), 18:133"“152.
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Back issues of many Annals of Carnegie Museum articles are available, and a few early complete volumes and parts are listed at half price. Orders and inquiries should be addressed to: Publications Secretary, Carnegie Museum, 4400 Forbes Avenue, Pittsburgh, Pa. 15213.
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CARNEGIE MUSEUM OF NATURAL HISTORY 4400 FORBES AVENUE ® PITTSBURGH, PENNSYLVANIA 15213 VOLUME 55 23 MAY 1986 ARTICLE 4
YANOMAMA MATERIAL CULTURE IN THE CARNEGIE MUSEUM OF NATURAL HISTORY. PART IL WEARING APPAREL AND FESTIVAL ARTIFACTS
Lorraine Couture-Brunette Collections Manager, Section of Anthropology
Abstract
The Carnegie Museum of Natural History has one of the largest, most inclusive, and best documented collections of Yanomama material culture in the world. The collection, consisting of two accessions comprising 572 specimens, spans a 5 year period from 1979 to 1984. This corpus of material documents not only traditional Yanomama material culture, but also shows the changes it has undergone due to the introduction of Western goods and materials. Part I dealt with material culture categories related to Food Pro- curement and Household Articles (Couture-Brunette, 1985). Part II deals with Wearing Apparel and Festival Artifacts; and Part III will deal with Foreign Influence and Mis- cellaneous Constructions.
Introduction
The Yanomama Indians inhabit an area of approximately 30,000 (Smole, 1976:3) to 100,000 (Migliazza, 1972:20) square miles in north- ern Brazil and southern Venezuela. They are one of the largest Indian populations in the Amazonian rain forest. In spite of their large pop- ulation, they have been able to remain isolated and unacculturated until the present due to their settlement locations olf major waterways in the Guyana Shield area. There is no of Yanomama. While Chagnon (1974:
Submitted 2 April 1985,
63
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Annals of Carnegie Museum
VOL. 55
of 10,000, Saffirio (1985:36) feels that 16,400 in 320 villages is closer to the true number.
All Yanomama speak four mutually intelligible languages and several dialects. The group has been called by several names; terms such as “Waica,” “Xiriana,” “Shamatari,” and “Yanoama,” have all been used to designate this family of languages. “Yanomamo,” “Yanam,” “San- uma,” and “Yanomam” actually denote only one dialect or language within the larger group. Thus “Yanomama” is used to refer to the entire family of languages (Migliazza, 1972:33). The Catrimani River Yanomama of Brazil, from whom the Carnegie Museum of Natural History collections have come, belong to the linguistic group “Yano- mam.” They inhabit areas of the Federal Territory of Roraima and the State of Amazonas.
Since the construction in the mid 1970s of Brazilian Highway BR 210, the Perimetral Norte, deep into Yanomama territory in Brazil, acculturation has been proceeding at a rapid rate. The introduction of Western goods, beliefs, and diseases has brought about profound and permanent changes in the Yanomama lifestyle (Saffirio, 1980; Saffirio et al., 1983). Like many South American groups before them the Yan- omama are substituting Western material culture in replacement of their traditional technology. This loss of native crafts is documented in the Carnegie Museum of Natural History collections.
Giovanni Saffirio, a member of the Consolata Society for Foreign Missions, arrived at a mission post on the Catrimani River in 1968. He was the collector of Yanomama material culture for Carnegie Mu- seum of Natural History. He has recently been awarded his Ph.D. in the Department of Anthropology at the University of Pittsburgh, and has since returned to the Catrimani Mission.
Collection I, accession number 32703, was made during 1979-80, while collection II, accession number 32735, took place in 1984. Every effort was made to secure as complete a representation of Yanomama material culture as possible, with a wide range in variation among specimens. As a result, the Carnegie Museum of Natural History has one of the largest, best documented, and most inclusive collections in the world.
The Catrimani River Yanomama Indians of South America, and the Carnegie Museum of Natural History collections of Yanomama ma- terial culture, have been presented in more detail in the Part I article (Couture-Brunette, 1985). Part II covers the Wearing Apparel and Fes- tival Artifacts segment of the collection.
The classification of an artifact as either an item of wearing apparel or a festival artifact is based on the function of the artifact, regardless of its method of construction. Some specimens which served the same
1986
Couture-Brunette— Yanomama Material Culture, II
65
Fig. L— Women’s aprons, an item of everyday apparel. Accession 32735.
purpose were constructed differently based on their use as an item of everyday wear or as a festival artifact. In these cases the entire group of specimens was placed in the category where it was most often used. In this manner, separation of like artifacts was avoided. Two examples, one from each of the Festival Artifacts and Wearing Apparel categories, will illustrate this placement. Women’s aprons are worn every day, but certain aprons in the Carnegie Museum of Natural History collection are made specifically for festivals. All aprons were placed into the Wearing Apparel category in order to discuss the “apron” group as a whole. Highly decorated aprons which were intended for festival usage are separately indicated.
It proved impossible to explain fully all manufacturing terminology and methods of the many different materials (basketry, cordage, knots) used in construction of the spec-
66
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imens. Standard references were used-Emery (1980) when discussing the cordage, Ado- vasio (1977) for the basketry, and Shaw (1972) for knot descriptions.
Numbers for the types of cordage and letters for the knots are used throughout the analysis (Couture-Brunette, 1985:tables 1 and 2). All knots are illustrated (Couture- Brunette, 1985:fig. 4). “S” and “Z” twist in cordage is also illustrated (Couture-Brunette, 1985:%. 5).
Some of the cordage consists of a number of spun and plied yams which were combined by twisting. Rather than list these varying numbers of yams as separate constmctions, they were combined in one cordage type (see Couture-Bmnette, 1985:table 1), with the number of plies referred to in parentheses after the cordage type number. For example, cordage type #12(4) consists of four pieces of 2 ply s spin Z twist yams, all combined in an S twist. Similarly, cordage type #13(4) is composed of four pieces of 3 ply s spin Z twist yams, combined in an S twist. Cordage types #5 and #6, as well as types #11 through #15, all consist of varying numbers of plies, and will be used with parenthetical numbers indicative of these.
All Yanomama words in the text are set in italics. Spelling follows SafRrio (1980, 1985). Due to the great number of Yanomama languages and dialects, these words are specific to the Catrimani River villages only.
Wearing Apparel
Men and women do not wear an abundance of clothing. The only item which men wear every day is a string around the waist, to which they tie their penis foreskin. Occasionally men will also wear a “belt” of native cotton. Women wear cotton “aprons” which consist of a thick belt in the back with a short fringe in the front.
Both men and women pierce their ears. Women also pierce the center and comers of their lower lip, and the nasal septum. Everyday “jewelry” consists of straws worn in the lip holes, and straws or bunches of grass in the nose and ears. Feather earrings are worn by both men and women, usually during festivals.
Traditionally, necklaces are made from seeds, shells, feathers, and other animal parts stmng on yamaasik fiber cordage or cotton string. Only men wear necklaces made from animal parts, although women utilize feathers and shells in earrings, necklaces, and aprons.
Many men and women have begun to wear Western shorts and T shirts since the arrival of Westerners on a frequent basis on Brazilian Highway BR 210. Young men no longer pierce their ears, and women near the highway do not pierce their lips or nose (Saffirio, 1980; Saffirio et aL, 1983). Increasing numbers of necklaces and earrings are made from trade beads and scrap metal acquired from Westerners.
Women's Clothing
“Aprons” (Figs. 1-2), which are worn every day, are a sign of a woman’s modesty. They are made by women from beginning to end; the cotton is spun and the cordage is plied and twisted by them. Women also wear armbands made from flowers, grasses, or cotton.
Aprons (pesimak). — For analytical purposes, the elements of the apron
1986
Couture-Brunette— Yanom AM A Material Culture, II
67
Fig, 2.— Beaded woman’s apron, CMNH number 32703-12. This would be worn during a festival.
were divided into three separate constructions —belt, fringe, and cord. The “belt” is the back portion of the apron. It is constructed in the same manner as the cotton hammock (Couture-Brunette 1985: figs. 28, 29). Stakes are placed in the ground the necessary distance apart, and one continuous strand of cotton yam is wrapped back and forth. The ends are then tied with a cotton cord, and the stakes are removed. Belts contain approximately 150 warp strands. They average twice the length of the front portion of the apron: the “fringe.” This is constmcted by folding short pieces of cordage in half over a length of cotton cord. The pieces are then twined tightly with cotton or yamaasik string just below the cord. One end of the fringe cord is tied permanently to the belt. The term “cord” denotes both the cord over which the fringe is folded and twined, and the cord which ties the two ends of the belt together. In most cases these two pieces of cordage are not the same, differing usually in the number of plies contained within the twisted rope.
Decorative seeds, beads, shells, or feathers are occasionally tied to each end of the fringe. The entire apron is sometimes dyed with nara (a reddish dye) in shades of brown, yellow, red, or orange.
Table l.— Yanomama women’s aprons in the collections of Carnegie Museum of Natural History.
68
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VOL. 55
•1.8 « 6
|
T |
in |
o- |
NO |
q |
in |
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no’ |
in |
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d |
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CM |
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m |
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m |
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00
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d •'; |
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cU |
i3 |
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JS |
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c/) |
GO |
N |
c/5 |
c/3 |
c/3 |
c/3 |
c/3 |
N |
oo |
N |
c/3 |
C/3 |
c/3 |
C/O |
C/3 |
|
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C/D |
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|
u |
uK K |
|
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|
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<n NO <n |
|
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<Nfn(N(N(NfN<N<N(N(N(NeN O fo ' cn fn fn m to tn ro fn tn
On On'^OnOnOnOnOnOnOnOnOnOs
(N (N fN fN r-J fn ro (n tn m
5 5t ^ 5 5
On On On On On
^ ^
o <n
N (N
fN
(N
in m"
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^No -CM ^
oo '~'oo oo NO NO ..
.sCn ^ -r-N>-NO^ - ? ^ .-OA^.-V
r-^t^r^'^^.-<en«n s ...in-HCNTi-NO
-d d
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(Nfnmmorommmminfn-g^ ommmmm
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^ I I I I 1 I I 1 I I I I I I I I I I
<n
-44L #13 (7); 1 1 (4) #7 #13 (2) A, C, G(2) none Z slant 28.4 19.4
-44M #1 1 (6); 13 (3) #9 #13 (2) A(6), C S slant S slant 40.4 15.8
-44N* #13(2) #11(3) #14(2) Z slant S slant 35.5 16.5
-440 #13(11); 15(7) #7; 9; 10 #15(2) A, C, E S slant S slant 47.6 25.6
-44P #13(6) #9 #13(2) A(7), C Z slant S slant 51.2 21.6
Table \. — Continued.
1986
Couture-Brunette— Yanom AM A Material Culture, II
69
.2 5
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tN |
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,-4 |
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fN (N CN (N fN
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CO |
CO |
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|
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d |
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CO |
m |
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Os |
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Os |
OS |
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|
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CN |
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CN |
CN |
CN |
CN |
CN |
m |
t Two halves of fringe made by two people (?); cordage and decoration are both dilferent.
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The aprons are fairly standardized in construction, although some cordage variation is apparent in Table 1. The belt cordage is the most consistent element in all aprons. Of the 27 specimens in collection 32703, 24 of them (89%) used cordage type #9 for the belt, either alone or in combination with another type. Cordage types #10, #1 1(3), and #14(4) were each used on one belt, and type #7 (in combination with type #9) was used on two. Fourteen of the 16 aprons from the 32735 collection (87.5%) used cordage type #9 on the belt. Two belts used a combination of cordage types #9 and #7, and one belt used type #8.
Twelve of the collection I belts have occasional rows of twining to hold the warp yams together. Of these belts, 75% of the twining is S slant and 25% is Z slant (Couture-Bmnette, 1985:fig. 15). Ten of the collection II belts have rows of twining, 90% S slant and 10% Z slant. This mixture of S and Z slant twining is interesting in view of the fact that 100% of the basketry twining is S slant. Stitch slant in twining is a recognizable attribute, which is standardized over a cultural area (Adovasio, 1977:30). It is unusual to find this variation in the aprons, which utilize twining only incidentally, and not in the twined basketry. There, the greater number of specimens and the greater amounts of twining would make some variation more understandable.
The cordage used on the fringe is generally the same cordage present on the belt, but is doubled back on itself and re-twisted. For example, 2 ply s spin Z twist cotton cordage (type #9), the most common belt cordage, is doubled and S twisted to produce the 2 ply s spin z plied S twisted cordage of type #13(2), the most common fringe cordage. This cordage is present on 24 of the 27 collection I aprons and 14 of the 16 collection II aprons.
The twining which secured the pieces of fringe to the cord is S slant on 78% of the 32703 aprons and Z slant on 22%. Of the 32735 fringes, 87.5% of the aprons have S slant twining, and one each are Z slant and stitched.
The most interesting fact about the twining on both belts and fringes is the fact that S slant twining on one is often paired with Z slant twining on the other. This can also be seen on Table 1, where eight of the 32703 aprons, representing 67% of the belts which have twining, pair S slant twining on the belt with S slant twining on the fringe. One apron exhibits Z slant twining on both belt and fringe, while two of the aprons, 17% of the belts which have twining, combine Z slant twining on the belt with S slant twining on the fringe. The same is tme of the 32735 aprons; of the 10 belts with twining, 70% of them have S slant twining on both belt and fringe, 10% pair S slant on the belt with Z slant on the fringe, and 1 0% pair Z slant on the belt with S slant on the fringe. One of the aprons utilizes S slant twining on the belt and stitching on the fringe. These facts suggest that belts and fringes are
1986
Couture-Brunette-— Yanom AM A Material Culture, II
71
made by different women, given the standardization of stitch slant in twining which was discussed earlier. This deduction was later confirmed by the collector (Saffirio, personal communication).
The cords, which tie the ends of the belt and serve as the ‘‘backbone” to the fringe, are different from one another on 16 of the collection I aprons (59%) and 1 3 of the collection II aprons (81%). These differences are generally exhibited in the number of plies contained in the cord; if this variance is regarded as a function of cord diameter they are as standardized as the other apron elements. Cordage type #13(variable number of plies) is used on 25 of the 27 collection I aprons in at least one of the two possible places (belt or fringe) and on all of the collection II aprons on either the belt or fringe. The number of s spin Z twist plies used in cordage type #13 was widely variable, with four to eight the most common amount. Of the 32703 aprons, cordage type #13 with six to eight plies was the most popular, whereas on the 32735 aprons four and five plies were more often used. One apron in the 32703 accession used a five strand sennit cord and another exhibited a commercially braided cord. One of the 32735 aprons used cord made from yamaasik fiber.
Knot type “A” was the most popular knot present on the aprons. Nearly all of the cords were finished off with this knot on each string end. Knot type “C,” the second most popular knot, was used to attach the fringe to the belt on 23 of the 27 collection I aprons and 12 of the 16 collection II aprons. The other knots present on the aprons were generally used to attach decorative elements to the fringe. Occasionally the fringe had additional pieces of cordage attached to the main cord with knot type “H,” the lark’s head.
Taking all these facts into consideration, the “standard” apron con- sists of cordage type #9 on the belt, cordage type #13(2) on the fringe, and cordage type #13(variable number of plies) making up the cord. Twenty-two of the 27 aprons in collection I, 81%, are standard. The percentage was even higher for the 32735 aprons, where 14 of the 16 aprons, 87.5%, are standard.
The two accessions differ more in their mode of decoration than in their method of construction. These decorative differences between collections I and II are a reflection of collection bias; the earlier 32703 accession contains more aprons which were worn every day, while many festival aprons were collected for the later 32735 accession.
Only four specimens from collection I have decorative accessories added to the apron. Two aprons (32703-44E and -440) have shells strung on cotton or yamaasik string and tied to each fringe end, whereas one apron (- 1 4C) has patches of curassow skin and feathers tied to each fringe end. The fourth apron, - 1 2, is an entirely different construction (Fig. 2). The belt is standard cordage type #9 and the cord is type
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Fig. 3.— Two traditional women’s necklaces of tirimoku seeds, CMNH numbers 32703-7A and B (left). Right, two aroariki, men’s necklaces worn for protection against evil spirits and spells, CMNH numbers 32703-8A and B.
#12(5). The fringe, however, consists of a 27.4 by 13.2 cm beaded rectangle in red, white, and light and dark blue. There are red (cordage type #10) and black (cordage type #9) tassels at each comer, and a small red fringe along the bottom of the apron (cordage type #10). The beaded design is a geometric motif with opposing triangles within bands of contrasting color. The beadwork on an apron such as this is very intricate and time consuming, and the apron was very expensive to purchase from the maker. It was brought from Themaim, a Maxiko- piutheri woman from the Mucajai River, in December 1979.
Most of the collection II aprons are decorated with beads, tirimoku seeds, shells, and feathers tied to each fringe end. Feathers are the most popular decorative .elements; specimens 32735-291, -296, and -300 use toucan, curassow, and lovely cotinga feathers to decorate each fringe end, and the fringe on apron -293 is constmcted entirely from curassow feathers stitched to the cord. Aprons -297, -298, and -299 used curas- sow and toucan feathers in combination with beads, shells, and seeds. Apron -294 utilizes strings of beads and seeds at each fringe end, while -288 makes use of strings of seeds.
1986 Couture-Brunette— -Yanom AM A Material Culture, II 73
Fig. 4.— Three kotho, belts for boys and men. From top: 32703- 16B, -16C, and -16A.
The fringe on apron -292 appears to have been made by different women. While the cordage, type #13(2), is the same throughout the fringe, the diameter of both the spun and plied yams is quite different. Likewise, the decorations at each end of the fringe are different; one side has shells stmng on cordage types #9 and #1 3(5), whereas the other side decoration is beads and seeds. This suggests that decorative ele- ments on aprons may be specific to individual women, another de- duction later confirmed by the collector (Saffirio, personal communi- cation).
Armbands. -—Most of the decorative arm ornaments worn by women are composed of highly perishable flowers or grasses; therefore, no specimens of these are present in the Carnegie Museum of Natural History collections. The only women’s armbands in the collection are a pair of cotton armbands for a young girl. Although at first glance they appear to be loom woven cloth, the fabric stmcture is actually plain interlinking crossed right over left (Emery, 1980:61). This circular method of construction produces a seamless, stretch fabric.
32735-21 A&B, wao kik, owned by Yaikom. Pink cotton girl’s armband, seamless, cordage type #9, D— 10,1 cm.
Necklaces. Carnegie Museum of Natural History collection
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contains three traditional seed necklaces (Fig. 3). These are rarely made by women today because of the availability of trade beads.
32703-7A, bought from a Hewenahipiutheri woman, December 1979. Tirimoku seeds on cordage type #12(3) tied with three knot “A”s. L— 40.6 cm.
32703-7B, same as 32703-7A except cordage type #3 tied in knots “A” and “G.” L— 55.9 cm.
32735-35, yanuak, woman’s necklace. Fourteen loops of small, round black seeds strung on cordage type #3 tied in knot “F.” Two knot “A”s one string end. L— 43.8 cm.
Men’s clothing
The penis string is the only item of clothing considered essential for everyday men’s wear (Saffirio, personal communication). Occasionally men will also wear a “belt” of native cotton (Fig. 4). Initially this is made by women in the same fashion as the belt portion of the women’s apron. But after the ends have been tied and the stakes removed, the entire length of the belt is very tightly wrapped with yanaaasik or cotton string every 2 to 5 cm. The belt is often dyed red or brown with nara.
Belts (kothoj.
32703- 16A, man’s belt. Cordage type #9 belt; cordage #13(5,6) cord. Loose warps, no wrapping. L— -56.6 cm,
32703-16B, man’s belt, from Mahuku of Hewenahipiutheri village, December 1979. Cordage type #7 belt with cordage type #9 tie every 2 cm. Cordage type #3 cord tied at each end with knots “E,” “G,” and “F.” Yams dyed red with nara; three purple serpentine lines painted down belt length. L~73.2 cm.
32703- 16C, boy’s belt. Cordage type #9 belt; cordage type #13(4) cord. Belt wrapped with green commercial string every 3 cm. Tied each end with knots “A,” “F,” and “H.” L~38.6 cm.
32703-301, boy’s belt. Cordage type #7 belt; cordage type #13(6) cord. Tightly wrapped with cordage type #1 every 2 cm; tied with knots “A”(4) and “E”(2). Dyed red with nara, L— 36.2 cm.
Necklaces.— ThQSQ necklaces (Fig. 5) are worn only by men, because they are made of animal parts from game the men have hunted and killed.
32703-9, thihi nak, jaguar tooth necklace from Paxeko, a 30 year old good hunter in Hewenahipiutheri village. Cordage type #3, knot “E”(?). L— 36.9 cm,
32703-10, opo nak, from Iropitheri village November 1979. Armadillo teeth tied to cordage type #1 with knot “E.” L— 39.4 cm.
32735-43, opo sina. Cordage type #3 necklace, with pendant of two armadillo tails tied to untwisted yamaasik fiber with knots “A” and “E.” L— 37.9 cm.
32735-84, necklace of cordage type #3 with pendant of two cock of the rock bird upper beaks and crests stitched together with commercial pink thread. Tied together with knots “G” and “F.” L— 36.5 cm.
Festival Artifacts
This category includes all items used at feasts and for ritual or spir- itual purposes. Items of clothing (feather armbands, headbands, and
1986
Couture-Brunette— Yanom AM A Material Culture, II
75
Fig. 5. —Two necklaces made from animal parts, and thus worn only by men. CMNH numbers 32703-9 (left) and 32703-10 (right).
earrings) worn only during festivals are included, although highly dec- orated women’s aprons, usually worn during festivals, are grouped into the “Wearing Apparel” category with the rest of the aprons as discussed previously.
Feasts, which can last up to a week, are primarily political events among the Yanomama. Although each village is self-sufficient in terms of daily life and food procurement, alliances with other villages are promoted for marriage alliances and cooperative warfare. The Catri- mani River Yanomama follow a prescriptive bilateral cross-cousin marriage rule and use an ‘Troquois-Dra vidian” kinship system (Chag- non, 1977:56; Saffirio, 1985:1 18). However, one village often does not provide enough cross-cousins to satisfy this rule. A man must fre- quently seek a wife outside of his own village as a result (Smole, 1976: 76, 94). When he finds an eligible female, he is obliged to reciprocate to her father or brothers with his own sister. Alliances between villages are strengthened by the double tie (Chagnon, 1977:55).
Warfare is another situation calling for village alliances. Several vil- lages may band together to fight a common enemy, or one village may flee to an ally when warfare drives them out of their own territory.
76
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They are occasionally forced to live in the allied village for a lengthy period of time —up to several years —until their new gardens begin to produce (Chagnon, 1977:97ff).
Both the Brazilian Indian Agency (FUNAI) and missionary groups have discouraged warfare among the Yanomama in recent years. As a result, ritual chest pounding duels during festivals are currently re- placing fatal raiding parties (Saffirio, personal communication). The duels allow the Yanomama to settle grievances among themselves or with allied villages. These duels are usually performed during the fes- tivals.
Body Decoration
Festival wear consists of feather or animal skin armbands and head- bands for men and feather earrings for men and women. In addition, both sexes paint themselves elaborately and cover their hair with bird down.
Yanomama men and women paint their bodies frequently during the dry season and almost daily during the rainy season with black {nara uxirim) and red {nara wakirim) paint. Body painting is especially colorful and elaborate during feasts, so the paints and storage equipment were placed into the Festival category.
Red body paint is made from seeds {nara moko) of the Bixa orellana tree. These have a greasy red outer covering, which is rubbed off and molded into a ball (Fig. 14) or made into a thick liquid to be stored in gourds.
32735-93, black paint gourd, owned by Carrera. End sealed with wax. String through hole below gourd neck: cordage type #3 tied in two knot “A”s. L— 10.7 cm W-8.1 cm.
32735-342, black paint gourd? sealed with leaf plug. Neck wrapped with scotch tape, no string. L— 11.2 cm W—7.7 cm.
32703-29, lump of red paint. L— 5.9 cm.
32735-310, same as 32703-29. L-=7.0 cm.
32735-311, same as 32703-29. L^7.8 cm.
32703-23, red paint gourd sealed with wax. Cordage type #3 through hole in gourd neck, knot “A” on visible end. L— 18.8 cm W— 1 1.6 cm.
Apparel. —Festival armbands for women are made of leaves and flowers; none of these are present in the Carnegie Museum of Natural History collection. Festival armbands for men are made from bird feathers and body parts.
White bird down feathers (horomaep) are also used by both Yano- mama men and women. They coat the hair with resin, and then cover their heads with the white down. The down on women occupies only a fringe around the head, but men apply it to completely cover the hair. The down is stored in gourds {horokoto), and retrieved by poking a stick into the cut-oflf top.
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Fig. 6.— Feather earrings {kurakaas sina) worn by men and women during festivals. All are accession 32735.
32703-20A, horokoto with horomaep, from Mahuku of Hewenahipiutheri village De- cember 1979. Top cut off; hole for tie string through neck. L—15.4 cm. 32703-20B, same as 32703-20A. L— 16.5 cm.
32703-20C, same as 32703-20A. L-19.3 cm.
32703-2 1C, corked with wad of leaves. Cordage type #1 1(3) tied through hole with knot ‘T”(?). L-12.4 cm.
32703-52A, collected 1979-”80. Cordage type #3 tied in neck hole with knot “A.” L— 15,1 cm.
32703-52B, same as 32703-52A. L-15.4 cm.
32735-106, top cut off; no hole in neck for cordage. L— 12,9 cm.
32735-107, corked with leaf wad; cordage type #4 tied through hole with knots “E” and “A.” L-12.1 cm.
32735-108B, two holes drilled through neck; cordage type #3 threaded through both holes and tied inside gourd with unidentifiable knot. L— 16.6 cm.
32735-108B, no stopper; cordage type #9(?) through hole. L— 25.1 cm.
As has been discussed previously, men wear items made from animal body parts such as wings, skins, teeth, and tails. Women, however, are permitted to wear feather earrings. These earrings, also worn by men, consist primarily of single feathers inserted into pieces of straw (Fig. 6). Sometimes the feathered end of the straw is smeared with resin.
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Table 2. — Yanomama feather earrings in the collections of Carnegie Museum of Natural
History.
|
Type |
Accession number |
Feathers (number) |
Cordage |
Length (cm) |
|
seisi sina |
32735-46 A&B |
lovely cotinga |
#2 |
9.5 |
|
-47 A&B |
lovely cotinga |
yamaasik |
11.2 |
|
|
-53 A&B |
lovely cotinga |
yamaasik |
11.0 |
|
|
-54 A&B |
lovely cotinga |
yamaasik |
17.0 |
|
|
kurakaas sina |
-55 A&B |
parrot (single) |
none |
16.5 |
|
-56 A&B |
parrot (single) |
none |
17.0 |
|
|
-57 A&B |
parrot (single) |
none |
16.6 |
|
|
-58 A&B |
parrot (single) |
resin |
19.4 |
|
|
-59 A&B |
parrot (single) |
resin |
19.7 |
|
|
-60 A&B |
parrot (single) |
yamaasik |
22.5 |
|
|
-61 A&B |
parrot (single) |
yamaasik |
21.4 |
|
|
-62 A&B |
parrot, curassow |
yamaasik |
15.6 |
|
|
-63 A&B |
parrot, curassow |
yamaasik |
17.7 |
|
|
-64 A&B |
parrot, curassow |
yamaasik |
17.3 |
|
|
-65 A&B |
curassow (two) |
none |
13.4 |
|
|
-66 A&B |
curassow (two) |
none |
9.9 |
|
|
-67 |
curassow (two) |
none |
12.8 |
|
|
-68 A&B |
curassow (two) |
none |
11.3 |
|
|
-69 A&B |
curassow (two) |
none |
11.3 |
|
|
-70 A&B |
curassow (two) |
none |
11.5 |
|
|
-71 A&B |
toucan, parrot, curassow |
#2 (?) |
18.7 |
|
|
-72 A&B |
parrot, toucan |
none |
14.3 |
|
|
-73 A&B |
parrot, curassow |
#1 |
15.0 |
|
|
-74 A&B |
guan, curassow |
#1 |
12.2 |
|
|
-75 A&B |
parrot (single) |
none |
11.9 |
|
|
-76 A&B |
parrot (single) |
none |
11.6 |
|
|
-77 A&B |
guan (single) |
none |
7.2 |
|
|
-78 A&B |
guan (single) |
none |
10.0 |
|
|
-79 A&B |
guan (single) |
none |
11.7 |
|
|
-80 A&B |
parrot (single) |
none |
11.3 |
|
|
-81 A&B |
guan (single) |
none |
11.6 |
|
|
-82 A&B |
curassow (two) |
none |
12.4 |
|
|
-83 A&B |
guan, curassow |
yamaasik |
13.0 |
either to protect underlying fiber wrapping or to hold the unwrapped feather in place.
Table 2 presents the data for the two types of feather earrings. The first, seisi sina, consists of lovely cotinga bird skin and feather patches wrapped to pieces of wood, which is used in place of straw because of the weight of the earring. One pair of earrings is wrapped with the rare Z twist yamaasik fiber cordage, whereas the others are secured with unplied yamaasik.
The second type of earring, kurakaas sina, utilizes a variety of feath-
1986
Couture-Brunette— Yanom AM A Material Culture, II
79
ers. Parrot and curassow are the most popular types, whereas toucan and guan are occasionally present. The commonest earring is the single or paired feather inserted into straw, seen in 2 1 of the 29 pairs. Eight pairs have a single earring in the center surrounded by a “bouquet” of curassow or toucan feathers.
Nine pairs use fiber wrapping to secure the feathers, and two pairs have resin coating the inserted end. The unplied yamaasik is again the most popular wrapping, but two pairs use S twist yamaasik, and one pair utilizes the rare Z twist yamaasik fiber.
Men’s clothing. — Men wear both arm- and head-bands during festi- vals. These are made by the men from bird or other animal parts. There are two types of armbands. The first, ara sina, consists of one to four long red tail feathers from the macaw {Ara macao) tied to pieces of
80
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VOL. 55
Table 2>. — Yanomama ara sina armbands in the collections of Carnegie Museum of
Natural History.
|
Accession number |
Macaw feathers |
Bunched feathers |
Cordage type |
Length (cm) |
|
32703-6A |
four |
parrot & macaw |
#1 |
66.1 |
|
-6B |
four |
white-winged guan |
#1 |
65.6 |
|
-6C |
four |
macaw, parrot, and white- winged guan |
#1 |
64.2 |
|
-35A |
three |
parrot |
#1 |
67.5 |
|
-35B |
three |
white-winged guan |
#1 |
59.4 |
|
~35C |
four |
white-winged guan |
#1 |
67.7 |
|
-35D |
three |
white-winged guan |
#1 |
65.6 |
|
-35E |
three |
white-winged guan |
#1 |
63.0 |
|
-35F |
four |
macaw, parrot and white- winged guan |
#1 |
64.0 |
|
-35 G |
three |
macaw |
#1 |
68.7 |
|
32735-121A |
four |
white-winged guan |
#1 |
60.9 |
|
-121B |
four |
white-winged guan |
#1 |
66.7 |
|
-122A |
four |
white-winged guan |
#1 |
51.8 |
|
-122B |
four |
white-winged guan |
#1 |
64.1 |
|
-123 |
one |
white-winged guan |
#1 |
60.4 |
|
-124 |
one |
parrot & white-winged guan |
#9 |
61.6 |
|
-125 |
three |
white-winged guan |
#1 |
69.5 |
|
-126 |
three |
white-winged guan |
#1 |
68.0 |
|
-127 |
four |
white-winged guan |
#1 |
52.1 |
|
-128A |
three |
white-winged guan |
#1 |
62.9 |
|
-128B |
three |
white-winged guan |
#1 |
66.4 |
|
-129 A |
three |
white-winged guan |
#1 |
68.4 |
|
-129B |
three |
white-winged guan |
#1 |
66.1 |
|
-130A |
four |
white-winged guan |
#1 |
66.2 |
|
-130B |
four |
white-winged guan |
#1 |
62.0 |
|
-131A |
three |
white-winged guan |
#1 |
64.7 |
|
-131B |
three |
white-winged guan |
#1 |
64.6 |
|
-132A |
four |
macaw & parrot |
#1 |
54.3 |
|
-132B |
four |
parrot |
#1 |
68.6 |
|
-133A |
four |
macaw |
#1 |
68.4 |
|
-133B |
four |
macaw |
#1 |
69.6 |
|
-134A |
four |
parrot |
#1 |
58.9 |
|
-134B |
four |
parrot |
#1 |
72.2 |
|
-135A |
four |
parrot |
#1 |
76.7 |
|
-135B |
four |
parrot |
#1 |
60.0 |
|
-136A |
four |
macaw & parrot |
#1 |
61.8 |
|
-136B |
four |
macaw & parrot |
#1 |
59.0 |
|
-137A |
four |
parrot |
#1 |
65.2 |
|
-137B |
four |
parrot |
#1 |
53.9 |
|
-138 |
three |
parrot |
#1 |
69.7 |
|
-139 |
three |
macaw |
#1 |
62.0 |
|
-140 |
three |
parrot |
#2 |
63.5 |
1986
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Table 3. — Continued.
|
Accession number |
Macaw feathers |
Bunched feathers |
Cordage type |
Length (cm) |
|
-141 |
three |
parrot |
#1 |
65.6 |
|
-142A |
four |
parrot |
#1 |
63.1 |
|
-142B |
three |
parrot |
#1 |
57.3 |
|
-143 A |
three |
parrot |
#1 |
65.9 |
|
-143B |
three |
parrot |
#1 |
72.0 |
|
-144 A |
three |
parrot |
#1 |
67.4 |
|
-144B |
three |
parrot |
#1 |
63.3 |
|
-MSA |
four |
parrot |
#1 |
65.5 |
|
-MSB |
four |
parrot |
#1 |
61.4 |
|
-146 A |
four |
parrot |
#1 |
57.9 |
|
-M6B |
four |
parrot |
#1 |
51.2 |
|
-147 |
two |
parrot |
#2 |
65.8 |
|
-148 |
four |
parrot |
#1 |
59.2 |
|
-149 A |
four |
parrot |
#1 |
59.2 |
|
-M9B |
four |
parrot |
#1 |
58.5 |
|
-ISOA |
four |
parrot |
#1 and #2 |
51.4 |
|
-ISOB |
four |
parrot |
#1 |
59.2 |
|
-ISIA |
three |
parrot |
#1 |
61.9 |
|
-ISIB |
three |
parrot |
#1 |
58.9 |
|
-152A |
three |
parrot |
#1 |
71.1 |
|
-152B |
three |
parrot |
#1 |
64.2 |
|
-153A |
three |
parrot |
#1 |
69.9 |
|
-153B |
three |
parrot |
#1 |
59.4 |
|
-1S4A |
three |
parrot |
#1 |
66.5 |
|
-1S4B |
three |
parrot |
#1 |
67.4 |
|
-15SA |
three |
parrot |
#1 and #2 |
70.8 |
|
-ISSB |
three |
parrot |
#1 |
67.3 |
bamboo or palm wood (Fig. 7). Bunches of parrot {Amazona sp.), curassow {Crax alector), or white-winged or piping guan {Aburria pip- He) feathers are tied in a “bouquet” at the base of the longer macaw feathers. The armband is then tied around the upper arm with the long red feathers pointing upwards. They are worn in pairs, with usually one on each arm.
Table 3 presents the data for the ara sina armbands. Length, cor- responding to the height of the long red tail feathers when worn, is not a critical factor in matching pairs of armbands; six pairs differ in size between pieces by 10 cm or more. It is more important to match the feathers, both in number of long red macaw feathers and in color and species of the “bouquet” feathers. All but two sets of armbands matched both the number of long red macaw feathers and the “bouquet” feathers
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VOL. 55
Fig. S. — Yaro sina, man’s festival armband made from bird tails.
on both armbands. One pair, 32735- 142A & B, matched four long red macaw feathers on one armband with three on the other. Only 32735- 132A & B did not match the “bouquet” feathers on the two armbands.
An interesting application of these facts can be tested with accession 32703 armbands, which were not delivered in pairs. Utilizing the cri- teria discussed above for matching pairs, it can be suggested that pairs are composed of specimens -6B and -35C, and -6C and -35F. Both of these pairs then have matched macaw and “bouquet” feathers. Both the macaw and the “bouquet” feathers match on three specimens (-35B, -35D, and “35E); any two of these three could form a pair of armbands. The “bouquet” feathers of -6 A and -3 5 A or -35G are somewhat matched. Whatever the pairs of the last two groupings, it is clear that the arm- bands from accession 32703 are not completely paired; the final set of, for instance, -35E and -35G is composed of completely unmatched “bouquet” feathers.
Cordage type #1 is clearly the choice to wrap the feathers to the wood. Cordage type #2, the rare Z twist yamaasik fiber, is present on four armbands, and s spin Z twist cotton cordage, type #9, on one. Interestingly, pairs 32735- 150A & B and -155A & B utilize both S
1986
Couture-Brunette —Yanom AM A Material Culture, II
83
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84
Annals of Carnegie Museum
VOL, 55
Fig. 9. — Yaw sina, man’s festival headband made from bird skins, wings, and tails.
and Z twist yamaasik fiber on one armband of the pair, whereas the other exhibits standard S twist yamaasik fiber. It is possible that these armbands which utilize mixed cordages were made by two individuals, one of whom wrapped the long red macaw feathers to the wood while the other attached the ‘‘bouquet” feathers. Accession 32703 armbands use S twist yamaasik fiber exclusively.
The second type of men’s armband, the yaw sina (Fig. 8), is com- posed of bird wings, tails, and skin. Aracari {Ptewglossus pluricinctus), parrot, toucan (Ramphastos rucanus), and macaw wings, tails, and skins are tied together to lengths of string. These lengths are tied around the upper arm, leaving the wings dangling.
Men usually wear the yaw sina armbands as a pair, one on each arm. Thus the armbands discussed in Table 4 are actually pairs of
1986
Couture-Brunette —Yanom AM A Material Culture, II
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Table S. — Yanomama yaro sina headbands in the collections of Carnegie Museum of
Natural History.
|
Accession number |
Wings and tails |
Headpiece |
Cordage |
Knots |
Length* (cm) |
|
32703-57 |
parrot |
curassow |
#13(3) |
A(3), C, F |
32.4 |
|
-58 |
parrot |
curassow |
#13(3) |
A(3), C, F, J |
24.2 |
|
-59 |
none |
curassow (2) |
#12(6) |
F |
34.8 |
|
-61 |
toucan |
none |
#10 |
H, K |
|
|
-62 |
none |
curassow |
#11 (3) |
25.7 |
|
|
-63 |
none |
curassow |
#11 (3) |
K |
25.1 |
|
-66 |
parrot, curassow |
curassow |
#13(3) |
A(3), C, F |
38.0 |
|
-67 |
toucan, guan |
curassow |
#13(3) |
A(2), C, J |
23.5 |
|
-68 |
toucan, guan, ma- caw, parrot |
curassow |
#3, 5 (3) |
E, F, H, C(2) |
30.4 |
* “Length” refers to the length of the headpiece.
armbands, which have been tied together. Some effort was made to balance the armbands; a pair with a toucan tail on one armband will be partnered with a toucan tail on the other. Only the smaller guan or parrot wings are occasionally present on one member of the pair but not the other.
Table 4 indicates that toucan tails are the parts used most consistently by all who make the yaro sina armbands. Only one specimen uses another bird tail in addition to the toucan tails, and all armbands but 32703-93 have toucan tails on them. Macaw wings are the second most popular bird part, present on four of the seven armbands. When the other parts are considered, primarily skins with feathers from the breast or head, macaw parts are present on five of the seven armbands.
Cordage type #9, the 2 ply s spin Z twist cotton, is used on six of the seven armbands. Cordage type #1 1(3) is present on four specimens, and type #13(2) on three. Only two armbands exhibit yamaasik fiber cordage, and only in minor amounts.
An interesting fact about the yaro sina armbands stems from the variation possible in their mode of construction. There are many types of cordage to choose from, and even more knots which can be used. As with stitch slant in twining, discussed previously, knots tend to reflect individual choice. Taking these facts into consideration, arm- band pairs 32703-54 and -65 were probably made by the same man. Both armbands used knot “H” exclusively to the the string to the bird parts. None of the other specimens uses knot “H” in this fashion; in fact, none of the other specimens uses knot '‘H” in any fashion. A lark’s head is not a “fixed” knot, but rather a suspended knot capable of movement along another piece of string. Although it can be rendered relatively immobile by the tightness of the tie, it is unusual to find it used as a method of permanent attachment. The specimens are not
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Annals of Carnegie Museum
VOL. 55
Fig. 1 0. —Incomplete yaw sina headbands, consisting of just the curassow skin headband.
overly similar in any other attribute; they both make heavy use of toucan parts, as do many of the other yaro sina armbands; they both use cordage types #9 and #13(2), which are the first and third most popular cordages; and one uses yamaasik fiber cordage, found on one other specimen. Nevertheless, the attachment of the parts to the string is unusual enough to merit special attention, and use of the rare lark’s head knot is a distinguishing characteristic which links these two pairs of armbands and separates them from the others.
The second item of men’s festival apparel is the headband made of bird parts or monkey skin. While the monkey headbands are simply made of skinned monkey tails, the feather yaro sina headbands, like the yaro sina armbands, are complex constructions of bird wings and tails tied to cotton or fiber cordage (Fig. 9). The band which passes across the forehead is made of curassow skin and feathers. A string is tied at each end of the band to make the head size adjustable, and the bird wings and tails dangle at each temple.
The yaro sina headbands are not as balanced as the yaro sina arm- bands; little effort was made to place similar wings and tails at each temple. As can be seen on Table 5, many of the headbands (32703-
1986
Couture-Brunette— Yanomam A Material Culture, II
87
Fig. 11.— Wisa sina, man’s festival headband made from a skinned black saki monkey tail. Accession 32735.
58, "59, -62, and -63) consist of just the curassow headband with no dangling parts (Fig. 10). Although bands are occasionally worn this way, it is usual to complete them with other bird parts at the temples. The reverse situation is found on 32703-61, which consists only of the earpieces to a headband; it has not yet been added to, or has been detached from, the curassow band.
Cordage type #13(3) is the most popular cordage used on the yaw sina headbands, present on four of the nine specimens. The rest of the cordage consists of two pieces of cordage type #1 1(3), and one each of types #12(6), #10, #3, and #5(3). In this respect the yaw sina headbands differ from the yaw sina armbands; cordage usage on the armbands was very consistent, with cordage type #9 present on six of the seven armbands, type #11(3) on four, and type #13(2) on three. Yamaasik fiber was used on one headband. Knot “A” was the most popular knot in total number of ties, whereas knots ‘‘C” and “F” were used on the greatest number of headbands.
A second type of headband, also utilizing feathers, is woven of masik vine. Curassow and toucan feathers are inserted into the weave in
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Annals of Carnegie Museum
VOL. 55
Fig. \l.~Horokoto with yakoana, hallucinogenic drug storage gourd. CMNH number 32703-22B.
patches of red, yellow, and black. This headband is not adjustable to different sizes.
32735-20, mayop ahuk, owned by Tixo. Band is tied at four points with cordage types #1 and #3 in knots “F”(2) and “A.” A 12 cm length in the back of the headband has no feathers. D™ 18.0 cm.
Other festival headbands consist of skinned monkey tails (Fig. 1 1). The fur piece is flattened, and an adjustable string is tied at each end.
32735-194, wixa sina. Black saki monkey skin tied with cordage type #3 in knot “H” at one end. L--(tail only) 42.7 cm.
32735-195, same as -194 but string tied with knot “E.” L— 41.2 cm.
32735-196, wixa sina tied with cordage type #19 in knot “C.” L— 36.9 cm.
Drug Containers
Most men use hallucinogenic drugs during festivals, al- though shamans make frequent use of them when performing sha- manistic rituals. The drug ( Virola sp.), also known as ebene, is made from inner tree bark (Chagnon et al., 1971:72-74). The latex from the tree is used as the poison on monkey arrows (Couture-Brunette, 1985: 501).
1986
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Fig. 13. —Chest pounding tools. From top: paruk xayau, CMNH number 32703-25 A; paruk xayau, CMNH number 32703-25B; suhumuk, CMNH number 32703-25C.
The moist scrapings are mixed with ashes and kneaded into a ball, with saliva providing additional moisture if necessary. It is then placed over the fire on a container to dry (Chagnon, 1977:23). The residue is ground into a powder, and stored in gourd containers until needed (Fig.
12).
32703-22A, horokoto with yakoana, from Mahuku of Hewenahipiutheri village, Decem- ber 1979. Corked with ball of wax; cordage type #9 through hole. Knot “A” inside gourd; two knot “A”s other end. L— 14.2 cm.
32703-22B, same as 32703-22A but corked with roll of animal hide. Cordage type #3 through hole in neck; tied with knot “E.” Outside of gourd covered with resin. L— 15.9 cm.
32735-102, sealed with wax. Cordage type #3 through neck hole; two knot “A”s one end and three on the other. L— 1 1 .4 cm.
32735-103, brown wood cork. Hole drilled in neck; no string. L— 15.1 cm.
32735-104, owned by Mahuku. Top sealed with wax, no hole or string. Split down side. L-12.4 cm.
32735-105, top sealed with wax. Cordage type #3, no visible knots. L— 15.3 cm. 32735-341, sealed with wax. L— 12.2 cm.
The drug is blown into the nostrils with a mokamosi, a hollow cane tube. One end of the tube is lined with wax while the other is unmod- ified. The tube is loaded with a dose of yakoana, and the receiver places
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CAre^MftO* CS4 6«
VAOOmA ry^O
Fig. 14.— Haya maro, deer bone flute (top) and nara wakirim, lump of red body paint (bottom). Accession 'ill 35.
the waxed end inside one nostril. Another man expels a powerful gust of air into the cane. A man will take a few doses of the drug at a time in each nostril.
32703-41, collected in 1979--80. Lengthwise cracks and splits along the sides of the tube. L— 8^4 cm.
32735-89, four parrot feathers in the waxed end. L— 69.8 cm.
32735-90, same as 32735-89. L^69.0 cm.
Chest Pounders
Ritual chest pounding duels during festivals have replaced much of the revenge warfare among the Catrimani Yanomama. Blows are de- livered with the aid of chest pounders (paruk xayau), wood branches which are smoothed and polished (Fig. 1 3). No effort is made to trim off protruding stubs, and the ends may be pointed or flat.
32703-25A, from Akasi and Paruk of Uxiutheri village, November 1979. Ends unpoint- ed. L— 35.9 cm.
32703-25B, same as 32703-25A but both ends pointed. L— 40.4 cm.
32735-109, owned by Carlos. Both ends pointed; approximately 5 cm from each end red and yellow toucan feathers are wrapped onto the pounder with cordage type #1. L— 22.1 cm.
1986
Couture-Brunette —Yanom AM A Material Culture, II
91
32735-263, one end pointed and other rounded. Cordage type #9 wrapped 10 times around one end. L— 28.9 cm.
32735-264, ovoid, both ends pointed. Small hole partially drilled into one side. “ME” carved on other side. L— 18.2 cm.
32735-265, forked. Two top ends pointed, other (bottom) end flat. L— 20.2 cm.
A second type of ritual dueling tool, suhumuk, is used in a different fashion. This object, also made of wood, is blade shaped with pointed ends and serrated teeth in the center (Fig. 1 3). Both ends are grasped by one man, who then hugs his opponent and digs the teeth into his back while the opponent does the same.
32735-25C, from Akasi and Paruk of Uxiutheri village, November 1979. Four teeth;
one end pointed, other flat. L— 38.5 cm.
32735-191, four teeth; both ends pointed. L— 77.6 cm.
32735-31, maama suhumuk, stone found by Honi on the path near the Arapari River (KM 135 of Highway BR 210). Honi said these were not longer used in dueling, but were made and used by ancestors. L— 23.4 cm.
Miscellaneous Festival Accessories
The Yanomama use two kinds of musical instruments, both of them flutes. The first is a deer bone flute (Fig. 14). Its use is restricted to festivals, when it is played by the guests as they draw near the host village. The leg bone is hollowed and holes are drilled into one side. The joint forms a natural closure for one end, while the other end of the flute is open. The flute is usually reddened with nara. The second flute, which is a child’s toy imitation of the ritual flute, will be discussed in Part III as a miscellaneous construction.
32703-49A, haya maro, collected 1979-80, 2 holes drilled in top. L---12.4 cm. 32703-49B, same as 32703-49A but three holes drilled in top. L— 15.1 cm.
32703-49C, same as 32703-49B. L-14.6 cm.
32735-50, two drilled holes and one natural hole, faint traces of nara. L— 18.2 cm. 32735-51, same as 32735-50. L-=17.8 cm.
32735-52, three drilled holes, reddened with nara. L— 18.9 cm.
32735-303, three drilled holes alternate with three “X”s scratched onto the surface.
Reddened with nara. L— 19.0 cm.
32735-304, same as 32735-303. L--19.4 cm.
Two necklaces, although occasionally worn outside of festivals, are intimately associated with the spirit world and were thus grouped in the Festival category with the drug equipment. The first, aroariki (Fig. 3), is worn for protection against evil spells and spirits. It is composed of cut pieces of tubers which have been dyed brown. The pieces are threaded on brown dyed string.
32703-8 A, from Puuxim of Wakathautheri village. Cordage type #10, eight knot “A”s. L— 35.4 cm.
32703-8B, same as 32703-8A. Cordage type #3, with knots “A”(5), “E”(2), and “J”(2). L--38.2 cm.
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The other type of necklace, marasik, is worn by men. It acts as an aphrodisiac, attracting partners of the opposite sex. It consists of cut pieces of tubers, which are dyed red and tied to reddened cotton string.
32735-44A, cordage type #11(3) tied around center of each tuber in knots “E,” “G,” and “F.” Ends tied together with two knot “A”s. L--70.0 cm.
32735-44B, same as 32735-44A but string tied around tubers with knot “E.” Ends tied together with knot “E.” L--69.1 cm.
Additional Comments
The two categories of Wearing Apparel and Festival Artifacts are intimately related, inasmuch as most festival artifacts are items to be worn. The division of specimens into selected categories was intended to reflect the function of the artifact. However, in two cases the con- struction of the specimen outweighed the functional aspects. Specifi- cally, the women’s aprons and the body paint groups were discussed together under one category, although in function each group should have been separated. In the case of the aprons, this combined discussion followed a constructional basis. Everyday and festival aprons are made the same way, and differ only in the added decorative elements. As this was the case, the “apron” group was combined into one discussion section under “Wearing Apparel” rather than presenting separate but identical constructional data in two categories. The “body paint” group was similarly combined for the same reasons.
Acknowledgments
I wish to thank Dr. Giovanni Saffirio for his invaluable advice and explanations on particularly puzzling specimens. Drs. James B. Richardson and David R. Watters kindly read through initial drafts and provided helpful comments. Dr. Hugh H. Genoways and the staff of the Section of Mammals, Drs. Kenneth Parkes and D. Scott Woods of the Section of Birds, and Dr, Juan Parodiz and Ms. Jay Tripp of the Section of Malacology assisted in the identification of materials used in manufacture of many of the artifacts. Ms. Nancy Perkins of the Division of Exhibits drafted the knots as well as the cordage/ basketry twist examples. Specimen photographs were taken by Ms. Barbara Gundy; all other photographs are used with the kind permission of Giovanni Saffirio.
Literature Cited
Adovasio, J. M. 1977. Basketry technology— a guide to identification and analysis. Aldine Publ. Co., Chicago, Illinois, 182 pp.
Chagnon, N. a. 1974, Studying the Yanomamo. Holt, Rinehart, and Winston, New York, New York, 270 pp.
— . 1977. Yanomamo: the fierce people. Holt, Rinehart, and Winston, New York,
New York, 2nd ed., 174 pp.
Chagnon, N. A., P. Le Quesne, and J. M. Cook. 1971. Yanomamo hallucinogens: anthropological, botanical, and chemical findings. Current Anthropology, 12: 72-74.
Couture-Brunette, L. 1985. Yanomama material culture in the Carnegie Museum
1986
Couture-Brunette— Yanom AM A Material Culture, II
93
of Natural History. Part I. Food procurement and household articles. Ann. Carnegie Mus. 54:487-532.
Emery, I. 1980. The primary structure of fabrics. The Textile Mus. Press, Washington, D.C., 2nd ed., 339 pp.
Migliazza, E. 1972. Yanomama grammer and intelligibility. Ph.D. dissertation, In- diana Univ., University Microfilms #72-30,432, Ann Arbor, Michigan, 457 pp.
Saffirio, G. 1980. Some social and economic changes among the Yanomama of northern Brazil (Roraima): a comparison of “forest” and “highway” villages. Un- published MS thesis, Univ. Pittsburgh, Pennsylvania, 1 1 9 pp.
— 1985. Ideal and actual kinship terminology among two Yanomama villages of
the Catrimani River basin (Brazil). Unpublished Ph.D. dissertation, Univ. Pitts- burgh, Pittsburgh, Pennsylvania, 244 pp.
Saffirio, G., R. Hames, N. Chagnon, and T. Melancon. 1983. The impact of contact: two Yanomamo case studies. Pp. 1--52, in Working papers on South American Indians, Bennington College Press, Bennington, Vermont, 6.
Shaw, G. R. 1972. Knots, useful and ornamental. First Collier Books ed., Macmillan Publ. Co. Inc., New York, New York, 194 pp.
Smole, W. 1976. The Yanoama Indians: a cultural geography. Univ. Texas Press, Austin, 272 pp.
Back issues of many Annals of Carnegie Museum articles are available, and a few early complete volumes and parts are listed at half price. Orders and inquiries should be addressed to: Publications Secretary, Carnegie Museum, 4400 Forbes Avenue, Pittsburgh, Pa. 15213.
ISSN 0097-4463
ANNALS
ofCAf!jNECIE MUSEUM
CARNEGIE MUSEUM OF NATURAL HISTORY 4400 FORBES AVENUE ® PITTSBURGH, PENNSYLVANIA 15213 VOLUME 55 23 MAY 1986 ARTICLE 5
STANDARD KARYOLOGY OF NINE SPECIES OF VESPERTILIONID BATS (CHIROPTERA: VESPERTILIONIDAE)
FROM THAILAND
Karen McBee
Rea Postdoctoral Fellow, Section of Mammals
John W. Bickham^
SONGSAKDI YeNBUTRA^
Jarujin Nabhitabhata^
Duane A. Schlitter Curator, Section of Mammals
Abstract
Karyotypes of nine species of vespertilionid bats from Thailand are described. Pip- istrellus mimus (2n = 34, FN = 46), Tylonycteris robustula (2n = 32, FN = 50), Murina leucogaster (2n = 44, FN = 50), and Miniopterus schreibersi (2n = 46, FN = 52) have karyotypes essentially identical to ones previously reported from other regions. Pipis- trellus pulveratus (2n = 32, FN = 50) is reported for the first time and differs by six Robertsonian fission/fusion events from the primitive MyotisA.ik.Q karyotype. Karyotypes
95
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for Hesperoptenus tickelli (2n = 32, FN = 50) and H. blanfordi (2n = 34, FN = 60) are reported for the first time and parallel the extreme morphological differences between the two species. Harpiocephalus mordax (2n = 40, FN = 62) is very distinct from other members of the subfamily Murininae but is apparently derived from a Murina-\ik& ancestor. Kerivoula papillosa (2n = 38, FN = 52) though considered little differentiated from primitive vespertilionines has a relatively highly derived karyotype similar to Vespertilio.
Introduction
The family Vespertilionidae is distributed worldwide in temperate and tropical regions. It is the largest family in the order Chiroptera including approximately 33 genera and 3 1 3 Recent species (Koopman, 1984). Thirteen genera and 34 species are known to occur in Thailand (Lekagul and McNeely, 1977).
Previous karyotypic studies led Pathak and Sharma (1969) to suggest that the family has two very different patterns of chromosomal vari- ability. Some genera such as Myotis exhibit remarkable homogeneity with all examined species having 2n = 44, FN = 50 or 52. Others such as Pipistrellus (2n = 26, 28, 30, 32, 34, 36, 38, 42, 44 and FN = 44, 46, 48, 50, 52) are much more heterogeneous. These studies, however, mostly have been restricted to New World (Baker and Patton, 1967; Bickham, 1979<2, 1979Z?) and European (Bovey, 1949; Capanna and Civitelli, 1970; Fedyk and Fedyk, 1970; Zima, 1978) species. Karyo- typic data for African, Australian, and Asian vespertilionids are sparse. For example, karyotypes have been reported for only one species of vespertilionids from Thailand (Harada et al., 1982Z?).
This study presents standard karyotypes of nine species in seven genera and four subfamilies from Thailand. Karyotypes of five of these species have been reported from other regions (Pathak and Sharma, 1969; Manna and Talukdar, 1965; Yong et al., 1971; Bickham and Hafner, 1978; Harada and Kobayashi, 1980; Harada, 1973; Ando et al., 1977). New data are presented for four species and one subfamily.
Materials and Methods
All animals were collected in Thailand using mist nets. Upon capture, all animals were subcutaneously injected with a weak solution of baker’s yeast, sugar, and water (Lee and Elder, 1980) to stimulate bone marrow mitosis. Twenty-four hours later, animals were sacrificed and humeri removed. Karyotypes were prepared in the field from bone marrow cells suspended in a hypotonic solution (0.075 M KCl) for approximately 25 min and then fixed in a 3:1 solution of methanol : glacial acetic acid (Baker et al., 1982). Three to four drops of the fixed cell suspension were dropped onto clean, dry microscope slides and ignited with a match. After the flaming suspension extinguished itself, any remaining liquid was carefully drained away and slides were stained in a 2% solution of Geimsa in 0.01 M phosphate buffer. Diploid (2n) and fundamental (FN) numbers were deter- mined from counts of a minimum of 10 mitotic spreads. Description of chromosome morphology follows the nomenclature of Patton (1967). All specimens were prepared as
1986
McBee et al.™ Thailand Vespertilionid Karyology
97
museum skins and skulls or alcoholics and are housed in the Carnegie Museum of Natural History (CM), the Texas Cooperative Wildlife Collection, Texas A&M University (TCWC), or The Museum, Texas Tech University (TTU).
Specimens Examined
Pipistrellus mimus.—UTnM Thani Prov.; Lan Saka Dist., Huai Kha Khang Wildlife Sanctuary, 15°29'N, 99®18'E (CM 88129 M, 88130 F); Huai Kha Khang Wildlife Sanc- tuary, 2.7 km S Khao Nang Rum Wildlife Research Station, 15®27'N, 99®18'E (CM 88132 M); Huai Kha Khang Wildlife Sanctuary, 1.5 km W Khao Nang Rum Wildlife Research Station, 15°29'N, 99n7'E (CM 88131 M).
Pipistrellus pulveratus. — Uthai Thani Prov,; Lan Saka Dist., Huai Kha Khang Wild- life Sanctuary, 3.7 km S, 1 km E Khao Nang Rum Wildlife Research Station, 15®27'N, 99°18'E (CM 88134 F, CM 88136 F).
Tylonycteris robustula. — SuKAT Thani Prov.; Tha Chang Dist., 15 km N, 23 km W Ban Maruan, 09°18'N, 98®58'E (CM 88149 F, CM 88151 F, CM 88152 F, CM 88140 F, TTU 41257 F, TK 21416 F).
Hesperoptenus blanfordi.—UTHM. Thani Prov.; Lan Saka Dist., Huai Kha Khang Wildlife Sanctuary, 15°19'N, 99°18'E (CM 881 14 M, TTU 41255 F); Huai Kha Khang Wildlife Sanctuary, 1.5 km W Khao Nang Rum Wildlife Research Station (TK 21279 F).
Hesperoptenus tickelli. —Uthai Thani Prov.; Lan Saka Dist., Huai Kha Khang Wild- life Sanctuary, 3.7 km S, 1 km E Khao Nang Rum Wildlife Research Station, 15®27'N, 99®18'E (CM 881 19 M, TK 21 193 M); Huai Kha Khang Wildlife Sanctuary, 2.0 km S Khao Nang Rum Wildlife Research Station, 15°30'N, 99°16'E (CM 88117 M).
Kerivoula papillosa. —Surat Thani Prov.; Tha Chang Dist., 15 km N, 23 km W Ban Maruan, 09°18'N, 90®58'E (CM 88164 F).
Miniopterus schreibersi haradai. —Uthai Thani Prov.; Lan Saka Dist., Huai Kha Khang Wildlife Sanctuary, 2.7 km S Khao Nang Rum Wildlife Research Station, 1 5°30'N, 99®16'E (CM 88156 M); Huai Kha Khang Wildlife Sanctuary, 3.7 km S, 1 km E Khao Nang Rum Wildlife Research Station, 15°27'N, 99“18T (CM 88157 M).
Murina leucogaster.— Uthai Thani Prov.; Lan Saka Dist., Huai Kha Khang Wildlife Sanctuary, Khao Nang Rum Wildlife Research Station, 15®29'N, 99®18'E (CM 88162 F, CM 88163 F).
Harpiocephalus mordax. —Uthai Thani Prov.; Lan Saka Dist., Huai Kha Khang Wildlife Sanctuary, Khao Nang Rum Wildlife Research Station, 15®29'N, 99®18'E (CM 88159 F).
Results
Table 1 is a summary of the known standard karyotypic data for the family Vespertilionidae including those reported here. The standard karyotypes of eight species representing seven genera and four sub- families are presented in Figs. 1--3. A brief description of these karyo- types follows.
Subfamily Vespertilioninae
Pipistrellus mimus (2n 34, FN = 46; Fig. la). —Four animals
examined have a karyotype that consists of six pairs of large metacentric to submetacentric chromosomes and one large subtelocentric pair. There are nine pairs of acrocentric chromosomes ranging in size from medium
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Annals of Carnegie Museum
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Table l.—A summary of known standard karyotypic data for the family Vespertilionidae. SM—submetacentric, M—metacentric, ST—subtelocentric, and A— acrocentric.
Taxon
Subfamily Vespertilioninae Myotis auriculas Myotis austroriparius
Myotis bechsteini Myotis blythi
Myotis brandti Myotis californicus Myotis capaccinii
Myotis dasycneme Myotis daubentonii
Myotis elegans Myotis emarginatus
Myotis evotis
Myotis fortidens Myotis frater
Myotis grisescens
Myotis horsfieldi
Myotis hosonoi
Myotis keaysi
Myotis keenii
Myotis leibii Myotis lucifugus Myotis macrodactylus
|
2n |
FN |
X |
Y |
|
44 |
52 |
SM |
A |
|
44 |
50 |
SM |
SM |
|
44 |
50 |
SM |
A |
|
44 |
52 |
M |
A |
|
44 |
52 |
SM |
A |
|
44 |
50 |
SM |
A |
|
44 |
50 |
SM |
A |
|
44 |
50 |
SM |
SM |
|
44 |
50 |
_ |
_ |
|
44 |
52 |
M |
A |
|
44 |
50-52 |
SM |
A |
|
44 |
54 |
SM |
|
|
44 |
52 |
M |
A |
|
44 |
50 |
SM |
SM |
|
44 |
50 |
SM |
A |
|
44 |
56 |
M |
A |
|
44 |
52 |
M |
A |
|
44 |
50 |
SM |
SM |
|
44 |
52 |
SM |
A |
|
44 |
50 |
SM |
A |
|
44 |
50 |
SM |
A |
|
44 |
50 |
SM |
A |
|
44 |
50 |
SM |
A |
|
44 |
50 |
SM |
A |
|
44 |
52 |
SM |
A |
|
44 |
50 |
SM |
A |
|
44 |
50 |
SM |
A |
|
44 |
50 |
— |
|
|
44 |
50 |
SM |
SM |
|
44 |
50 |
SM |
A |
|
44 |
50 |
SM |
SM |
|
44 |
50 |
SM |
SM |
|
44 |
52 |
SM |
A |
|
44 |
52* |
SM |
A |
|
44 |
50 |
SM |
A |
Authority
Bickham, \919b Baker and Patton, 1967 Bickham, \919b Zima, 1978 Baker, 1970 Baker et aL, 1974 Zima, 1982
Baker and Patton, 1967 Manfreddi Romanini et aL, 1975
Zima, 1978 Bovey, 1949 Fedyk and Fedyk, 1970 Zima, 1984
Baker and Patton, 1967 Bovey, 1949 Radjhabli et al., 1969, 1970
Zima, 1978
Baker and Patton, 1967 Bickham, \919b Osborne, 1965 Harada and Yoshida,
1978
Baker and Patton, 1967 Bickham, 1979^
Harada and Kobayashi, 1980
Harada, 1973 Harada and Yoshida,
1978
Bickham, 1979^
Baker and Bickham, 1980 Baker and Patton, 1967 Bickham, \919b Baker and Patton, 1967 Baker and Patton, 1967 Harada, 1973 Obaraetal., 1976a Harada and Yoshida,
1978
1986
McBee et al.— Thailand Vespertilionid Karyology
99
Taxon
Myotis milieri Myotis myotis
Myotis mystacinus Myotis nattered
Myotis nigricans
Myotis oxygnathus
Myotis pruinosus Myotis sodalis
Myotis thysanodes
Myotis velifer
Myotis {Pizonyx) vivesi Myotis volans Myotis yumanensis
Lasionycteris noctivagans
Pipistrellus abramus
Pipistrellus affinm Pipistrellus babu Pipistrellus endoi
Pipistrellus hesperus Pipistrellus kuhli
Pipistrellus mimus
Table 1. — Continued.
|
2N |
FN |
X |
Y |
|
44 |
52 |
SM |
__ |
|
44 |
50 |
M |
A |
|
44 |
50 |
SM |
A |
|
44 |
50 |
SM |
A |
|
44 |
56 |
M |
A |
|
44 |
50 |
SM |
__ |
|
44 |
50 |
SM |
__ |
|
44 |
52 |
SM |
A |
|
44 |
50 |
SM |
SM |
|
44 |
50 |
SM |
A |
|
44 |
50 |
SM |
A |
|
44 |
56 |
M |
A |
|
44 |
50 |
SM |
A |
|
44 |
52 |
SM |
ST |
|
44 |
50 |
SM |
— |
|
44 |
50 |
SM |
A |
|
44 |
50 |
SM |
SM |
|
44 |
52 |
SM |
A |
|
44 |
50 |
SM |
SM |
|
44 |
50 |
SM |
A |
|
44 |
50 |
SM |
SM |
|
44 |
50 |
SM |
SM |
|
44 |
50 |
SM |
__ |
|
44 |
50 |
SM |
A |
|
20 |
28 |
SM |
A |
|
20 |
28 |
SM |
A |
|
26 |
44 |
ST |
A |
|
26 |
44 |
A |
A |
|
26 |
44 |
A |
A |
|
36 |
50 |
SM |
A |
|
36 |
50 |
M |
A |
|
38 |
50 |
A |
__ |
|
38 |
50 |
A |
|
|
28 |
46 |
SM |
A |
|
44 |
50 |
SM |
A |
|
44 |
50 |
SM |
__ |
|
44 |
50 |
SM |
A |
|
34 |
__ |
__ |
__ |
Authority
Reduker et al., 1983 Bovey, 1949 Bickham and Hafner,
1978
Iliopoulou-Georgudaki and Giagia, 1984 Radjhabli et al., 1969, 1970
Ando et al., 1977 Haradaand Yoshida, 1978 Zima, 1978
Baker and Patton, 1967 Bickham, \919b Baker and Bickham, 1980 Radjhabli et al., 1969, 1970
Bickham and Hafner,
1978
Harada and Uchida, 1982 Baker and Patton, 1967 Bickham, \919b Baker and Patton, 1967 Bickham, \919b Baker and Patton, 1967 Bickham, \919b Baker and Patton, 1967 Baker and Patton, 1967 Baker and Patton, 1967 Bickham, 1979^
Baker and Patton, 1967 Bickham, 1979<2 Takayama, 1959 Harada, 1973 Obara et al., \916b,
1976c
Pathak and Sharma, 1969 Dulic, 1981 Ando et al., 1977 Ando et al., 1980 Baker and Patton, 1967 Capanna, 1968 Baker et al., 1974 Zima, 1982 Manna and Talukdar,
1965
100
Annals of Carnegie Museum
VOL. 55
Table Continued.
Taxon
Pipistrellus mordax Pipistrellus nanus
Pipistrellus nathusii
Pipistrellus pipistrellus
Pipistrellus savii
Pipistrellus pulveratus Pipistrellus subjlavus
Nyctalus furvus
Nyctalus lasiopterus
Nyctalus leisleri Nyctalus noctula
Eptesicus andinus Eptesicus brasiliensis
Eptesicus capensis
Eptesicus circumdatus Eptesicus diminutus Eptesicus furinalis
Eptesicus fuscus
Eptesicus guadeloupensis
|
2N |
FN |
X |
Y |
|
38 |
48 |
M |
A |
|
34 |
46 |
SM |
A |
|
34 |
46- |
M |
A |
|
36 |
50 |
M |
A |
|
44 |
51 |
SM |
A |
|
42 |
50 |
M |
|
|
42 |
50 |
M |
A |
|
42 |
51 |
M |
A |
|
44 |
50 |
M |
— |
|
44 |
50 |
M |
A |
|
44 |
50 |
SM |
A |
|
44 |
52 |
M |
A |
|
44 |
50 |
SM |
A |
|
44 |
50 |
__ |
— |
|
44 |
50 |
SM |
A |
|
32 |
50 |
SM |
__ |
|
30 |
56 |
SM |
A |
|
30 |
50 |
SM |
|
|
44 |
50 |
SM |
A |
|
44 |
50 |
SM |
A |
|
42 |
50 |
SM |
A |
|
42 |
50 |
SM |
A |
|
42 |
50 |
SM |
A |
|
42 |
50 |
SM |
A |
|
46 |
54 |
SM |
|
|
42 |
50 |
SM |
A |
|
42 |
50 |
SM |
A |
|
42 |
50 |
M |
A |
|
42 |
50 |
M |
M |
|
50 |
48 |
SM |
A |
|
50 |
48 |
SM |
A |
|
50 |
48 |
SM |
A |
|
32 |
50 |
SM |
A |
|
50 |
48 |
SM |
__ |
|
50 |
48 |
SM |
A |
|
50 |
48 |
SM |
A |
|
50 |
48 |
SM |
A |
|
50 |
48 |
SM |
A |
|
50 |
48 |
SM |
A |
|
50 |
48 |
SM |
A |
Authority
Pathak and Sharma, 1969 This study
Pathak and Sharma, 1969 Peterson and Nagorsen,
1975
Bovey, 1949 Fedyk and Ruprecht,
1976
Zima, 1978 Bovey, 1949 Fedyk and Ruprecht,
1976
Zima, 1978 Zima, 1982 Zima, 1984 Capanna, 1968 Park and Won, 1978 Zima, 1982 This study
Baker and Patton, 1967 Bickham, \919a Ando et al., 1977 Harada et al., 1982<2 Tsuchiya et al,, 1972 Harada, 1973 Ando et al., 1977 Harada et al., 1982t2 Fedyk and Fedyk, 1970 Dulic et al., 1967 Vorontsov, 1969 Zima, 1978 Zima, 1984
Baker and Patton, 1967 Baker and Patton, 1967 Baker et al., 1982 Peterson and Nagorsen, 1975
Heller and Volleth, 1984 Williams, 1978 Baker and Patton, 1967 Williams, 1978 Baker and Patton, 1967 Bickham, 1979a Genoways and Baker, 1975
1986
McBee et al.— Thailand Vespertilionid Karyology
101
Table \.— Continued.
|
Taxon |
2N |
FN |
X |
Y |
Authority |
|
Eptesicus hottentotus |
50 |
48 |
SM |
— |
Peterson and Nagorsen, 1975 |
|
Eptesicus japonensis |
50 |
48 |
SM |
SM |
Ando et al., 1977 |
|
Eptesicus lynni |
50 |
48 |
SM |
A |
Bickham, 1979^? |
|
Eptesicus nilssoni |
50 |
48 |
__ |
— |
Ando et al., 1977 |
|
50 |
50 |
M |
A |
Zima, 1978 |
|
|
50 |
48 |
M |
__ |
Zima, 1982 |
|
|
Eptesicus serontinus |
50 |
48 |
SM |
A |
Baker and Patton, 1967 |
|
50 |
48 |
SM |
A |
Vorontsov, 1969 |
|
|
50 |
52 |
SM |
SM |
Fedyk and Fedyk, 1970 |
|
|
50 |
48 |
SM |
— |
Baker etal., 1974 |
|
|
50 |
48 |
SM |
A |
Bickham, 1979^2 |
|
|
50 |
48 |
SM |
_ |
Baker and Bickham, 1980 |
|
|
Vespertilio murinus |
38 |
50 |
M |
A |
Vorontsov, 1969 |
|
38 |
54 |
M |
A |
Zima, 1978 |
|
|
38 |
50 |
— |
— |
Obara and Saitoh, 1977 |
|
|
Vespertilio orientalis |
38 |
50 |
M |
A |
Ando et al., 1977 |
|
38 |
50 |
SM |
A |
Obara and Saitoh, 1977 |
|
|
Vespertilio superans |
38 |
50 |
M |
A |
Vorontsov, 1969 |
|
38 |
50 |
M |
A |
Zima, 1978 |
|
|
Histiotus montanus |
50 |
48 |
SM |
A |
Williams and Mares, 1978 |
|
Tylonycteris pachypus |
46 |
52 |
A |
M |
Yonget al., 1971 |
|
Tylonycteris robustula |
32 |
52 |
A |
M |
Yonget al., 1971 |
|
32 |
52 |
A |
M |
This study |
|
|
Hesperoptenus blanfordi |
34 |
60 |
A |
- |
This study |
|
Hesperoptenus tickelli |
32 |
46 |
ST |
M |
This study |
|
Nycticeius humeralis |
46 |
48 |
SM |
A |
Baker and Patton, 1967 |
|
46 |
48 |
SM |
A |
Bickham, 1979^? |
|
|
Scotoecus hindei |
30 |
50 |
ST |
SM |
Nagorsen et al., 1976 |
|
Rhogeessa genowaysi |
42 |
50 |
SM |
SM |
Baker, 1984 |
|
Rhogeessa parvula |
44 |
50 |
SM |
SM |
Baker and Patton, 1967 |
|
44 |
50 |
SM |
Bickham and Baker, 1977 |
||
|
Rhogeessa tumida |
42 |
50 |
SM |
SM |
Baker and Patton, 1967 |
|
30 |
50 |
__ |
_ |
Baker, 1970 |
|
|
42 |
50 |
SM |
SM |
Bickham and Baker, 1977 |
|
|
34 |
50 |
SM |
SM |
Bickham and Baker, 1977 |
|
|
32 |
50 |
SM |
SM |
Bickham and Baker, 1977 |
|
|
30 |
50 |
SM |
ST |
Bickham and Baker, 1977 |
|
|
34 |
50 |
_ |
__ |
Baker and Bickham, 1980 |
|
|
30 |
50 |
__ |
Baker and Bickham, 1980 |
||
|
52 |
52 |
_ |
Honeycutt et al., 1980 |
||
|
34 |
50 |
SM |
__ |
Baker et al., 1985 |
|
|
32N |
50 |
SM |
_ |
Baker etal, 1985 |
|
|
32B |
50 |
SM |
__ |
Baker et al, 1985 |
|
|
30 |
50 |
SM |
A |
Baker et al, 1985 |
102
Annals of Carnegie Museum
VOL. 55
Table \. — Continued.
|
Taxon |
2N |
FN |
X |
Y |
Authority |
|
Scotophilus dinganii |
36 |
52 |
A |
M |
Schlitter et al., 1980 |
|
36 |
62* |
“ |
_ |
Peterson and Nagorsen, 1975 |
|
|
Scotophilus heathi |
36 |
52 |
M |
A |
Sharma et al., 1974 |
|
Scotophilus kuhlii |
36 |
52 |
M |
A |
Pathak and Sharma, 1969 |
|
36 |
48 |
M |
A |
Harada et al., 1982^? |
|
|
Scotophilus temminckii |
36 |
52 |
SM |
A |
Pathak and Sharma, 1969 |
|
36 |
48 |
SM |
A |
Harada and Kobayashi, 1980 |
|
|
Scotophilus viridis |
36 |
54 |
A |
M |
Schlitter et al., 1980 |
|
Lasiurus borealis |
28 |
46 |
SM |
A |
Baker and Patton, 1967 |
|
28 |
48 |
SM |
A |
Baker and Mascarello, 1969 |
|
|
28 |
48 |
SM |
A |
Bickham, 1979<2 |
|
|
Lasiurus cinereus |
28 |
46 |
SM |
A |
Baker and Patton, 1967 |
|
28 |
48 |
SM |
A |
Bickham, \919a |
|
|
Lasiurus ega |
28 |
48 |
SM |
A |
Bickham, \919a |
|
Lasiurus ega panamensis |
28 |
46 |
A |
A |
Baker and Patton, 1967 |
|
Lasiurus ega xanthinus |
28 |
46 |
SM |
A |
Baker and Patton, 1967 |
|
Lasiurus inter medius |
26 |
40 |
SM |
A |
Baker and Patton, 1967 |
|
26 |
42 |
A |
A |
Baker, 1970 |
|
|
Lasiurus seminolus |
28 |
48 |
SM |
A |
Baker and Mascarello, 1969 |
|
28 |
48 |
SM |
A |
Bickham, 1979<2 |
|
|
Barbastella barbastellus |
32 |
52 |
__ |
— |
Matthey and Bovey, 1948 |
|
32 |
50 |
M |
A |
Bovey, 1949 |
|
|
32 |
50 |
SM |
A |
Capanna et al., 1968 |
|
|
32 |
52 |
SM |
A |
Zima, 1978 |
|
|
Barbastella leucomelas |
32 |
50 |
SM |
A |
Ando et al., 1977 |
|
Plecotus auritus |
32 |
52 |
__ |
Matthey and Bovey, 1948 |
|
|
32 |
50 |
M |
A |
Bovey, 1949 |
|
|
32 |
54 |
SM |
A |
Fedyk and Fedyk, 1970 |
|
|
Plecotus auritus auritus |
32 |
54 |
M |
A |
Ando et al., 1977 |
|
32 |
52 |
M |
A |
Zima, 1978 |
|
|
Plecotus auritus sacrimontis |
32 |
50 |
M |
A |
Harada, 1973 |
|
Plecotus austriacus |
32 |
50 |
SM |
A |
Baker, 1970 |
|
32 |
54 |
SM |
A |
Fedyk and Fedyk, 1970 |
|
|
32 |
50 |
SM |
A |
Baker et al., 1974 |
|
|
32 |
52 |
M |
A |
Zima, 1978 |
|
|
Plecotus phyllotis |
30 |
50 |
_ |
__ |
Baker and Patton, 1967 |
|
30 |
50 |
SM |
A |
Baker and Mascarello, 1969 |
|
|
Idionycteris phyllotis |
30 |
50 |
SM |
_ |
Bickham, 1979a |
|
30 |
50 |
SM |
““ |
Stock, 1983 |
1986
McBee et al.— Thailand Vespertilionid Karyology
103
Table \. — Continued.
|
Taxon |
2N |
FN |
X |
Y |
Authority |
|
Plecotus rafinesquii |
32 |
50 |
A |
A |
Baker and Mascarello, 1969 |
|
Plecotus townsendi |
32 |
48 |
__ |
— |
Baker and Patton, 1967 |
|
32 |
50 |
A |
A |
Baker and Mascarello, 1969 |
|
|
32 |
50 |
A |
A |
Bickham, 1979a |
|
|
32 |
50 |
A |
A |
Stock, 1983 |
|
|
Euderma maculatum |
30 |
50 |
SM |
A |
Williams et al., 1970 |
|
30 |
50 |
SM |
- |
Stock, 1983 |
|
|
Subfamily Miniopterinae |
|||||
|
Miniopterus australis |
46 |
50 |
SM |
A |
Harada and Kobayashi, 1980 |
|
Miniopterus magnater |
46 |
50 |
SM |
A |
Harada and Kobayashi, 1980 |
|
Miniopterus schreibersi |
46 |
50 |
_ |
__ |
Matthey and Bovey, 1 948 |
|
46 |
50 |
SM |
A |
Baker et al., 1974 |
|
|
46 |
50 |
SM |
A |
Bickham and Hafner, 1978 |
|
|
46 |
50 |
SM |
A |
Bickham, 1979a |
|
|
46 |
50 |
SM |
A |
Harada and Kobayashi, 1980 |
|
|
Miniopterus schreibersi ha- |
46 |
52 |
SM |
A |
This study |
|
radai |
|||||
|
Miniopterus schreibersi fuli- |
46 |
52 |
SM |
A |
Harada, 1973 |
|
ginosus Subfamily Murininae |
|||||
|
Murina aurata |
44 |
60 |
SM |
A |
Ando et al., 1977 |
|
Murina leucogaster |
44 |
50 |
— |
Harada, 1973 |
|
|
44 |
58 |
SM |
A |
Ando et al., 1977 |
|
|
44 |
50 |
SM |
A |
This study |
|
|
Harpiocephalus mordax |
40 |
62* |
- |
This study |
|
|
Subfamily Kerivoulinae |
|||||
|
Kerivoula papillosa |
38 |
52** |
This study |
||
|
Subfamily Nyctophilinae |
|||||
|
Antrozous pallidus |
56 |
50 |
SM |
A |
Bickham, 1979a |
|
Bauerus dubiaquercus |
44 |
52 |
SM |
A |
Engstrom and Wilson, 1981 |
|
* Obara et al., 1976^, report on inversion polymorphism in chromosome 5. |
** Includes sex chromosomes in FN.
* Examination of the figure in Peterson and Nagorsen (1975) gives a FN = 52. This probably represents a typographical error.
104
Annals of Carnegie Museum
VOL. 55
114 4# 4A 4A #•
la
II!
Fig. 1 . — The standard karyotypes of: a) Pipistrellus mimus F (CM 88 1 35), 2n = 34, FN = 46, inset F. mimus M (CM 88131); b) Pipistrellus pulveratus F (CM 88136), 2n = 32, FN = 50.
to minute. The X is medium-sized and submetacentric and the Y is small and acrocentric.
Pipistrellus pulveratus (2n = 32, FN = 50; Fig. lb). —The autosomal complement includes eight pairs of metacentric or submetacentric chro- mosomes ranging in size from large to medium. There is one large pair and one small pair of subtelocentric chromosomes, and five pairs of acrocentric chromosomes ranging from medium-sized to small. The X is medium-sized and submetacentric.
Tylonycteris robustula (2n = 32, FN = 50; Fig. 2a).— The karyotype shown here is similar to that reported by Yong et al. (1971), but there are slight differences. Both studies report 2n = 32 with nine pairs of metacentric to submetacentric chromosomes. However, our specimens had one pair of medium-sized subtelocentric chromosomes and five pairs of acrocentric chromosomes ranging from medium-sized to mi- nute, whereas Yong et al. (1971) reported two pairs of subacrocentric
1986
McBee et al.— Thailand Vespertilionid Karyology
105
|
ll U M M *kH )<« 0#' A5 1 XX 1 |
2a • <* T XY |
|
HI )) >1 >1 >11 II II 18 M H •• |
2b 1. XY |
|
lir V VI n IX 1 KR w *• Ift IM 0» ( |
2c XY |
Fig. 2.— The standard karyotypes of: a) Tylonycteris robustula F (TK 21416), 2n = 32, FN = 50; inset T. robustula M. (CM 88 1 52); b) Hesperoptenus tickelli M (TCWC 47481), 2n = 32, FN = 46; c) Hesperoptenus blanfordi F (CM 88114), 2n = 34, FN = 60.
and two pairs of acrocentric chromosomes. The X is large and acro- centric and the Y is a small metacentric chromosome.
Hesperoptenus tickelli (2n = 32, FN = 46; Fig. 2b).— The autosomal complement contains eight pairs of metacentric to submetacentric chromosomes ranging from large to medium-sized. There also are seven pairs of medium-sized to minute acrocentric chromosomes. The X is a large subtelocentric and the Y is a small metacentric chromosome.
Hesperoptenus blanfordi (2n ^ 34, FN = 60; Fig. 2c).— The auto- somal complement includes 1 3 pairs of metacentric to submetacentric chromosomes gradually decreasing in size from large to small. There
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Annals of Carnegie Museum
VOL. 55
is one pair of medium-sized subtelocentric chromosomes and two pairs of medium-sized acrocentric chromosomes. The X is a medium-sized subtelocentric chromosome, and the Y is a small subtelocentric chro- mosome.
Subfamily Kerivoulinae
Kerivoula papillosa (2n = 38, FN = 52; Fig. 3a).— Metacentric to submetacentric chromosomes include three large pairs, one medium- sized pair and one small pair. There is one medium-sized pair and one large pair of subtelocentric chromosomes, and 1 2 pairs of acrocentric chromosomes grading from large to small. The sex chromosomes were not identified in this species, but are probably the medium-sized pair of submetacentric chromosomes. The fundamental number includes the presumed sex chromosomes.
Subfamily Miniopterinae
Miniopterus schreibersi haradai (2n = 46, FN = 52; Fig. 3b).— The autosomal complement includes two large and one medium-sized pairs of metacentric chromosomes, and one medium-sized pair of subtelo- centric chromosomes. There are 1 8 acrocentric pairs ranging from large to small. One medium-sized acrocentric pair has a secondary constric- tion near the centromere. The X is a medium-sized submetacentric chromosome. Other reports of karyotypes from Miniopterus schreibersi (Harada, 1973; Ando et al., 1977; Bickham and Hafner, 1978) iden- tified the smallest pair of chromosomes as biarmed. This was not apparent from our preparations; however, the medium-sized subtelo- centric pair of chromosomes we observed is evidently unique to Thai Miniopterus schreibersi and is not seen in European (Bickham and Hafner, 1978) or Japanese (Harada, 1973) populations.
Subfamily Murininae
Murina leucogaster (2n = 44, FN = 50; Fig. 3c).— There are two pairs of large metacentric, and two pairs of medium-sized to small submetacentric autosomes. The autosomal complement is completed by 1 7 acrocentric pairs ranging from large to small. The X is medium- sized and submetacentric. This karyotype is similar to that of Murina leucogaster from Atesu, Japan (Harada, 1973), but the third largest
Fig. 3.— The standard karyotypes of: a) Kerivoula papillosa F (CM 88164), 2n = 38, FN =52 (FN includes sex chromosomes); b) Miniopterus schreibersi haradai F (CM 88157), 2n = 46, FN = 52; c) Murina leucogaster F (CM 88163), 2n = 44, FN = 50.
1986
McBee et al,— Thailand Vespertilionid Karyology
107
108
Annals of Carnegie Museum
VOL. 55
chromosome is considerably smaller in our material than in the Jap- anese bats.
Harpiocephalus mordax (2n ^ 40, FN = 62).— Poor field preparation of Harpiocephalus mordax made chromosome analysis difficult, but counts of metaphase spreads consistently gave a diploid number of 40. The chromosomal complement includes three large pairs, one medium- sized pair and two small pairs of metacentric to submetacentric chro- mosomes and five pairs of subtelocentric chromosomes. There are nine pairs of acrocentric chromosomes gradually decreasing from large to small. The single individual examined was a female, so sex chromo- somes could not be identified but are probably the medium-sized pair of submetacentric chromosomes. The fundamental number includes the presumed sex chromosomes.
Discussion
This karyological study is consistent with earlier studies (Capanna and Civitelli, 1 970; Baker and Patton, 1967; Pathak and Sharma, 1 969; Bickham, 1979Z?), which have indicated extensive chromosomal vari- ability between genera in vespertilionid bats. Within genera for which karyotypic data have been obtained on more than one species, three different patterns of chromosomal variability are apparent. Of the 1 5 genera (Table 1) for which more than one species have been karyotyped, 1 1 , including members of three subfamilies, can be characterized as conservative genera. These are genera in which all species have the same or nearly the same standard karyotype. Myotis (2n = 44, FN = 50, 52) and Eptesicus (2n = 50, FN = 48, with the exception of E. capensis) are typical representatives of this pattern. Scotophilus, Ves~ pertilio, Barbestella, Lasiurus, Plecotus, Miniopterus, and Murina also exhibit this pattern. Genera exhibiting the second pattern of variation are interspecifically variable. Five genera currently fill this group (Table 1). Pipistrellus has nine different diploid numbers among the 1 5 species that have been karyotyped. Nyctalus, with data from four species, shows three different karyotypes and Tylonycteris and Hesperoptenus each has two different karyotypes for two species. The third pattern is one of intraspecific variability and is best documented in the genus Rhogeessa (Table 1). Three species possess at least nine different karyo- types. Rhogeessa parvula has a 2n = 44, FN = 50 karyotype, and R. genowaysi has 2n = 42, FN = 50. R. tumida, however, has 2n = 30, 32a, 32b, 34, 42, 44, 52 and FN = 50, 52).
Pipistrellus exhibits such wide variability that even with karyotypes for approximately one third of the recognized species no real patterns of karyotypic relationships are evident within the genus. Several species share the Myotis-MkQ 2n = 44, FN = 50 karyotype considered primitive for the family (Bickham, \919a, 1 9 79Z?; Baker and Patton, 1967). Many
1986
McBee et al.— Thailand Vespertilionid Karyology
109
of the other karyotypes can be related to each other and to the primitive Myotis-likQ karyotype on the basis of Robertsonian fusions and fissions. Pipistrellus pulveratus has a karyotype of 2n = 32, FN 50 in two individuals. The karyotypes appear to differ from the primitive Myotis- like karyotype by six Robertsonian fission-fusion events.
The karyotype of P. mimus cannot be so directly derived from the Myotis-\ik£ primitive. Manna and Talukdar (1965) reported a karyo- type of 2n ^ 34 from southwestern India and Pathak and Sharma (1969) found 2n = 38, FN = 48 for the same species in northeastern India. We found 2n == 34, FN = 46 for four individuals from north- western Thailand. This karyotype can be derived from the 2n == 38 karyotype through one centric fusion and the loss, possibly through tandem fusion, of one small pair of acrocentric chromosomes. The 2n = 38 karyotype has 6 pairs of large metacentric to submetacentric chromosomes and 12 pairs of acrocentric chromosomes. The Thai karyotypes have an additional biarmed chromosome that is subtelo- centric and three fewer pairs of acrocentrics. Neither karyotype can be derived from the Myotis-likQ primitive karyotype without the loss or tandem fusion of at least one pair of acrocentric chromosomes. These data support the suggestion of Pathak and Sharma (1969) that trans- locations other than Robertsonian fusions may play an important role in chromosomal evolution in some groups of Pipistrellus, These authors also suggest that the two karyotypic forms of P. mimus may represent cryptic species. Cryptic species differentiated by karyotypes have been discovered for the family among the many karyotypic forms of Rho- geessa (Baker, 1984), so this explanation of karyotypic variation in P, mimus is not unreasonable. An alternative explanation of the chro- mosomal variability observed in P. mimus may involve intraspecific variability. This phenomenon is very rare among vespertilionids but is well documented within the genus Rhogeessa. In either case, P. mimus merits comprehensive cytogenetic study throughout its range in southern Asia.
Our karyotype of Tylonycteris robustula is slightly different from that reported by Yong et al. (1971). The extremely small chromosomes they consider biarmed are probably acrocentric. The T. robustula autosomal karyotype readily can be derived from the Myotis-\ikQ primitive con- dition by a series of six centric fusions. The X chromosome, however, has experienced a pericentric inversion to an acrocentric condition and the Y, either a pericentric inversion or the addition of a heterochromatic short arm making it biarmed. An acrocentric or subtelocentric X chro- mosome is rare among the Vespertilionidae occurring in only two species of Pipistrellus, two species of Scotophilus, Scotoecus hindei, two species of Lasiurus, two species of Plecotus, and two species of Hesperoptenus (Table 1). Tylonycteris robustula also has diploid and
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Annals of Carnegie Museum
VOL. 55
fundamental numbers in common with the two Plecotus. The karyo- types are at least superficially the same, except both Plecotus have acrocentric rather than biarmed Y chromosomes. Assumptions of ho- mology on the basis of standard karyotypes must be made with caution, however (Bickham and Baker, 1977; Baker et al., 1985; Haiduk and Baker, 1982). The similarities between the plecotine genus, Plecotus, and the vespertilionine genus Tylonycteris are likely the result of con- vergence. Tate (1942) considered Tylonycteris and Philetor as derived from an ancestor similar to the Pipistrellus jojfrei group. Neither Phile- tor nor any members of the P. joffrei group have been karyotyped for comparison, however.
The genus Hesperoptenus is poorly understood systematically. Four species are currently recognized, two of which are known from only a few specimens. The more common forms, H. tickelli and H. blanfordi, are very different from one another morphologically. Tate (1942) com- mented that if the genus was not polyphyletic it at least contained strongly differentiated species. Hesperoptenus blanfordi and H. tickelli also are karyologically distinct. Whereas the two species have similar diploid numbers, H. blanfordi has one of the highest FNs reported for the family, and H. tickelli has an FN of 52, among the most commonly found in the family. To derive one karyotype from the other would require one fission/fusion event and five pericentric inversions or het- erochromatic additions. The H. tickelli karyotype can be derived from the Myotis-likt primitive karyotype through four Robertsonian fusions and the loss or tandem fusion of two pairs of acrocentrics. There also has been a pericentric inversion changing the primitive biarmed X to a derived subtelocentric configuration. The Y is a derived small meta- centric chromosome, either through pericentric inversion or hetero- chromatic addition. The H. blanfordi karyotype is more difficult to derive from the Myotis-M^iQ primitive requiring at least five Robert- sonian fusions and five pericentric inversions in the autosomal com- plement. The X chromosome also is inverted to an acrocentric con- dition. The more parsimonious scenario might consist of H. tickelli diverging from the Myotis-\i\iQ ancestor with H. blanfordi being a high- ly divergent offshoot of H. tickelli. Chromosomal data support the conclusion that H. tickelli and H. blanfordi, at best, are only distantly related. Ryan (1966) and Koopman (1971) thought Hesperoptenus was closely related to Glauconycteris and Chalinolobus. Hill (1976) con- sidered dental differences between the three genera to be too great and considered Hesperoptenus more closely aligned with the genus Scoto- philus. Hesperoptenus tickelli and some Scotophilus have FN and uni- armed X chromosomes in common. H. tickelli has one fewer pair of biarmed chromosomes and one fewer pair of acrocentric chromosomes
1986
McBee et al.— Thailand Vespertilionid Karyology
1
than Scotophilus, however. No species of Glauconycteris or Chalinolobus have been karyotyped for comparison.
Our karyotype of Miniopterus schreibersi haradai agrees well with previous reports. Miniopterinae is considered the most derived subfamily of the Vespertilionidae, even being accorded familial status by some authors (Mein and Tupinier, 1977) yet the karyotype found throughout this subfamily differs from the primitive Myotis-li^Q karyo- type by a single Robertsonian fission and two pericentric inversions (Bickham and Hafner, 1978). Harada (1973) found in M. s. fuliginosus, and we found in M. s. haradai, a medium-sized subtelocentric chro- mosome apparently unique to Thai members of the species.
Members of the subfamily Murininae have been regarded as a spe- cialized offshoot of an early Myotis-VikQ ancestor (Miller, 1907). The subfamily contains two genera, Murina and Harpiocephalus. All mem- bers of Murina karo typed so far have had a standard karyotype essen- tially identical to the 2n = 44 MyotisAi\.Q primitive, agreeing with the early divergence of Murininae from the vespertilionine line. Tate ( 1 94 1 ) considered the second genus, Harpiocephalus, as a very specialized offshoot of the line leading to Murina, and Miller (1907) termed Har- piocephalus as one of the most aberrant genera of the family. The Harpiocephalus karyotype is derived from the primitive Myotis-\ik.Q karyotype and the Murina karyotype by two possible pericentric in- versions indicating that Harpiocephalus probably evolved from a Mu- rina-likQ ancestor rather than diverging earlier from the line leading to Murina.
The subfamily Kerivoulinae has been considered the least specialized of the vespertilionid subfamilies being closely related to the “least progressive” genera of the subfamily Vespertilioninae (Tate, 1941). The karyotype of Kerivoula papillosa can be derived from the primitive Myotis-\ik& karyotype through two Robertsonian fusions and the loss or tandem fusion of one pair of acrocentric chromosomes.
Within the Vespertilionidae, Kerivoula shares a similar standard karyotype with the Japanese Pipistrellus endoi and members of the genus Vespertilio. In the past, the entire genus Pipistrellus has been considered a part of Vespertilio, but Zima (1978) considers the dis- tinctive karyotype of Vespertilio as justification for separate generic status. Ando et al. (1980) suggest P. endoi may be a link between the genus Vespertilio and its Pipistrellus -likQ ancestor. The subfamily Ker- ivoulinae may have a similar link to its Pipistrellus -likQ ancestor here. There are no other data to link the three, however, and postulation of a common origin is only speculative.
Unquestionably, Robertsonian fusions and fissions have played a major role in chromosomal evolution of the family Vespertilionidae
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(Bickham, 1979^). However, the standard karyotypes reported here indicate a greater importance for non-Robertsonian rearrangements such as inversions and translocations as evolutionary mechanisms than was previously thought (Bickham, 1979a; Bickham and Baker, 1977). Pericentric inversions, tandem fusions, or heterochromatic additions apparently have occurred in Tylonycteris robustula, Miniopterus schrei- bersi haradai, Kerivoula papillosa, and Harpiocephalus mordax in their evolution from the 2n = 44 ancestral karyotype. Hesperoptenus tickelli and H. blanfordi may show an especially high incidence of pericentric inversions, requiring up to five possible inversion events to be derived from the ancestral karyotype. Examination of these standard karyo- types emphasizes how poorly understood are vespertilionid karyolog- ical relationships. Speculations about relationships based on standard karyotypes can be misleading, however. G-band analysis has indicated that constant genera such as Myotis are indeed as constant as was assumed from standard karyotypes (Bickham, 1 919b). It also has point- ed out extreme chromosomal differences where standard karyotypes indicated homology (Baker et al., 1985). Chromosomal banding anal- ysis should allow a more accurate assessment of the mechanisms of chromosomal evolution seen in the family Vespertilionidae. G-banding also will provide a means to test apparent homologies between groups such as Kerivoula and Vespertilio. The extensive variability between, and possibly within, species of Pipistrellus also will be much better characterized by G-banding. Comparison of conserved and derived chromosome sequences revealed by G-banding is imperative to an understanding of systematic relationships among the Vespertilionidae.
Acknowledgments
We especially thank Craig S. Hood and Stephen L. Williams for their unflagging assistance in the field. Keith Studholm, Dorothy Pumo, Bill Frucht, and Carleton J. Phillips also provided assistance in the field. Dr. Niphan Ratanaworabhan acted as our hostess and provided invaluable assistance in Thailand. Tweewat Polpakdee, Somechai Kwanchareon, Supachai Sittilert, Monthida Sitathani, Puangtong Boonsong, and Precha Luecha acted as our guides and translators, enthusiastically helping us in every possible way. Gerard McKieman and Karen Muller aided in our literature search and Brian G. Hanks, Priscilla K. Tucker, Michael S. Smolen and Thomas E. Lee provided aid in the laboratory, and Teresa A. Heiner assisted in preparation of earlier manuscripts. Don E. Wilson and Rodney L. Honeycutt provided constructive reviews that enhanced the final manuscript. McBee received partial funding from a Tom Slick Graduate Research Fel- lowship at Texas A&M University. This study was supported in part by NSF Grant PCM-8202794, NIH Grant AIO 4242, and the Office of University Research Services, Texas A&M University.
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ISSN 0097-4463
ANNALS
0/ CARNEGIE MUSEUM
CARNEGIE MUSEUM OF NATURAL HISTORY 4400 FORBES AVENUE * PITTSBURGH, PENNSYLVANIA 15213 VOLUME 55 23 MAY 1986 ARTICLE 6
RESULTS OF THE CARNEGIE MUSEUM OF NATURAL HISTORY EXPEDITIONS TO BELIZE. L SYSTEMATIC STATUS AND GEOGRAPHIC DISTRIBUTION OF SIBON NEILLI (REPTILIA, SERPENTES)
C. J. McCoy
Curator, Section of Amphibians and Reptiles
Abstract
The nominal species Sibon neilli Henderson, Hoevers and Wilson (type-locality, “vi- cinity of Belize City, Belize”) is shown to be a southern subspecies of Sibon sanniola Cope (type-locality, “Chichen Itza, Yucatan, Mexico”), confirming the arrangement pro- posed by Kofron (1985). Sibon sanniola neilli differs from Sibon s. sanniola in having a banded, rather than spotted, color pattern, and higher ventral and subcaudal scale counts.
Introduction
Sibon neilli Henderson, Hoevers, and Wilson (1977) is an enigmatic member of the snake fauna of Caribbean Central America. The species was described from a unique type-specimen, collected in “the vicinity of Belize City, Belize District, Belize.” Henderson et al. (1977) rec- ognized that Sibon neilli is very similar to 5. sanniola, a species en- demic to the northern part of the Yucatan Peninsula, but diagnosed S. neilli on the basis of higher ventral and subcaudal scale counts, a different pattern of supralabial, postocular, and temporal scale contacts, and a banded color pattern. Kofron (1985),
Submitted 24 September 1985.
117
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McCoy— Status of Sibon neilli
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tional specimens of S. neilli and without comment, considered S. neilli a subspecies of Sibon sanniola.
During field work in Belize in 1984 we collected a specimen of Sibon at Chaa Creek, Cayo District, that has the characteristic banded dorsal pattern of S. neilli. In addition, there is a specimen in Carnegie Museum of Natural History from El Peten, Guatemala, that also fits the de- scription of Sibon neilli. I have compared these specimens with the holotype and previously only known specimen of S. neilli, with all other Sibon of this group available from Belize, and with an extensive series of Sibon sanniola from Yucatan and Quintana Roo, Mexico (see Specimens Examined).
Results
Ventral and subcaudal scale — Data summaries for ventral
and subcaudal scale counts for Sibon sanniola and Sibon neilli are presented in Table 1. Both the ventral and subcaudal counts for the holotype of Sibon neilli exceed the known ranges for these counts in S. sanniola, as pointed out by Henderson et al. (1977). The range of ventral counts for males in the “southern” population overlaps the range for the “northern” population, as do the ranges of ventral counts for females in the two populations. Subcaudal counts in males barely overlap in the two populations, and overlap slightly in females. Al- though the sample size for the southern population is small, it appears that the differences in segmental counts are consistent. A specimen from “British Honduras” (FMNH 4247, male) has 153 ventrals and 67 caudals, both counts within the ranges for the northern population.
SupralabiaVpostocular-temporal contact.— The holotype of Sibon neilli has 9-9 supralabials, with the 4th, 5th, and 6th entering the orbit. The 7th supralabial is in contact with the lower postocular and the primary temporal. Henderson et al. (1977) regarded the 7th supralabial- postocular-temporal contact as a distinctive character of S. neilli, stat- ing that it “occurs occasionally in sanniola, but in no other species of Sibon."" In reality, this is the contact pattern that is most common in Sibon sanniola. In a series of 51 S. sanniola from Yucatan and Quintana Roo the 7th supralabial contacts only the lower postocular and primary temporal 88% of the time (right and left sides of the head scored separately). In the remaining 12% the upper tip of the 7th supralabial, posterior angle of the lower postocular, and anterior tips of the primary and secondary temporals make a single point contact. In specimens with the more common pattern, the anterior tip of the primary temporal makes a broad contact with the postocular, pre- venting contact of the 7th supralabial with the secondary temporal.
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Fig. \.~Sibon sanniola neilli (CM 105981), Chaa Creek, Cayo Dist., Belize; from a Kodachrome photo of the living specimen by Paul S, Freed.
Color pattern. — The holotype of Sibon neilli has a series of 34 dark dorsal crossbands on a lighter ground color, faded almost to white in the specimen. The widest crossbands are IVi scales long at the middorsal line, and taper to about 1 Vi scales at the lowermost dorsal scale rows. The edges of the bands are jagged, not straight. The bands extend ventrally only to the outer ends of the ventral scales. The center of the belly is marked with a series of roughly paired, longitudinal dark “dash- es,” each usually two ventral scales long. There are about 21 dark crossbands on the tail, which become increasingly crowded toward the tail tip. The dark nuchal band extends ventrally to the edges of the ventrals, and anteriorly as a middorsal dark bar that reaches the frontal scale. The holotype is much faded, resulting in enhanced contrast be- tween the dorsal dark bands and the interspaces (Henderson et al., 1977:fig. 1).
The specimen from Chaa Creek, Belize (CM 105981), has 42 dorsal body bands that taper ventrally from IVi to IV2 scales wide, and 22 bands on the tail. The nuchal band has a broad anterior extension that reaches the parietals, and the top of the head is lighter brown. The sides of the head are much lighter. The belly is lightly marked with indistinct brown smudges. The only significant difference between the
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McCoy— Status of Sibon neilli
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pattern of this specimen and that of the holotype is the amount of contrast between the dorsal bands and the interspaces. In life, the dorsal bands of CM 105981 were medium brown, and the interspaces dark tan, providing minimal contrast (Fig. 1).
The specimen from Las Cahas, Guatemala (CM 58282), has 38 barely discernible dark bands on the body, and a banded tail. The bands are only slightly darker than the medium brown ground color. The head and nuchal pattern are typical, and the belly is moderately well-marked with longitudinal dashes.
The typical color pattern of Sibon sanniola is a series of dark, light- edged middorsal spots, frequently fused to form an irregular “zig-zag” line, on a light brown to ash gray background. The lateral and ventro- lateral spot series may either persist, be reduced, or fade completely. The tail is spotted along the middorsal line. The belly typically is marked with pairs of dark longitudinal dashes. The nuchal band, with anterior extension onto the head, is as described for Sibon neilli.
A juvenile specimen from Xunantunich, Belize (MCZ 56994, 180 mm total length), has a pattern of about 47 middorsal spots (many fused), and two series of lateral dark spots on each side. A very similar pattern occurs in juvenile S. sanniola (CM 49739, 140 mm; CM 49754, 156 mm). This suggests that the banded adult pattern of S. neilli may result from ontogenetic fusion of the dorsal, lateral, and ventrolateral spot series, which remain discrete in 5. sanniola.
Conclusions
In size, habitus, and most details of scutellation Sibon neilli and Sibon sanniola are identical. The supposedly diagnostic arrangement of supralabial, postocular, and temporal scales of S. neilli actually is consistent with the pattern usually found in S. sanniola. Only the color pattern and numbers of ventral and subcaudal scales are distinctive characters of the .S', neilli population. Although Kofron (1983) shows the range of S. sanniola (including S. neilli) as being continuous from northern Yucatan and Quintana Roo southward into Belize and El Peten, no specimens are available from the critical areas where inter- gradation would be expected (Lee, 1980 and personal communication). The southernmost precise locality for Sibon sanniola is Felipe Carrillo Puerto, Quintana Roo (Peters, 1953), although FMNH 4247 from “British Honduras” has both scale counts and color pattern typical of S. sanniola. The range of S. neilli extends from coastal central Belize (Belize City), southwestward into El Peten, Guatemala. Despite the apparent hiatus, I assume that the range of the species is continuous, as there is no ecological discontinuity between southern Quintana Roo and central Belize, and I interpret the morphological differences be-
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tween the southern and northern populations as clinal variation within a single species.
Clinal variation is common in colubrid snake species that have ex- tensive latitudinal ranges on the Yucatan Peninsula. Such clinal vari- ation typically involves either the color pattern or segmental counts, or both. Examples are Conophis lineatus (Wellman, 1963), Leptodeira frenata (Duellman, 1958), Ninia sebae (Schmidt and Rand, 1957), and Leptophis mexicanus (Oliver, 1 948), to cite a few of many species that illustrate this variational pattern. I conclude that Kofron (1985) was correct in allocating the name Sibon neilli to the southern population of Sibon sanniola that is characterized by a banded color pattern and higher ventral and subcaudal scale counts.
Specimens Examined
Sibon sanniola neilli. — Belize: Belize Dist., vicinity of Belize City (MPM 8929, ho- lotype). Cayo Dist., Chaa Creek, 5 mi. S San Ignacio (CM 105981); vicinity of Augustine (MPM 8208); Xunantunich (MCZ 56994). Guatemala: El Peten, Las Canas (Munici- pality San Luis) (CM 58282). Total 5 specimens.
Sibon sanniola sanniola.— Belize: “British Honduras” (FMNH 4247). Mexico: Quin- tana Roo, Pueblo Nuevo X-Can (CM 45778-45785, 46844-46845, 46881-46883, 49056, 49062, 49136, 49154, 49159, 49163). Yucatan, Chichen Itza (FMNH 20609, 20613, 26988, 36257-36258, 36268, 36272, 36276, 36285, 36287, 36289, 36296); Kantunil (FMNH 36264, 36286, 36288, 36270); Libre Union (FMNH 36259, 36262, 36266, 36273, 36278, 36280-36283, 36290, 36294-36295); Piste (CM 46955-46958, 47004, 47142-47148, 49734-49741, 49742-49748, 49749-49756); Progreso (FMNH 40734- 40735); Yokdzonot (FMNH 36261, 36263, 36265, 36267, 36269, 36271, 36274-36275, 36277, 36279, 36284, 36290-36293, 36297). Total 101 specimens.
Acknowledgments
Field work in Belize was supported by a grant from the O’Neil Museum Trust, Carnegie Museum of Natural History. For assistance in the field I thank D. Scott Wood, Robert C. Leberman, Paul and Mara Freed, and a column of anonymous army ants that enabled us to capture the Chaa Creek Sibon. I am indebted to the Flemings of Chaa Creek for their hospitality, to Ray E. Ashton, Jr., of International Expeditions, Inc., for logistical aid, and to Dora Weyer for steadfast support of our work in Belize. Collecting permits were granted by Mr. O. Rosado, Department of Forestry, Belize Ministry of Natural Resources. For loans of specimens I thank R. F. Inger and Hymen Marx, Field Museum of Natural History (FMNH), Robert W. Henderson, Milwaukee Public Museum (MPM), and Pere Alberch, Museum of Comparative Zoology, Harvard University (MCZ). I also thank Julian C. Lee for information on localities, and Ellen J. Censky for technical help.
Literature Cited
Duellman, W. E. 1958. A monographic study of the colubrid snake genus Leptodeira. Bull. Amer. Mus. Nat. Hist., 114:1-152.
Henderson, R. W., L. G. Hoevers, and L. D. Wilson. 1977. A new species of Sibon (Reptilia, Serpentes, Colubridae) from Belize, Central America. J. HerpetoL, 11: 77-79.
Kofron, C. P. 1983. Female reproductive cycle of the Neotropical snail-eating snake Sibon sanniola in northern Yucatan, Mexico. Copeia, 1983:963-969.
1986
McCoy— Status of Sibon neilli
123
1985. Systematics of the Neotropical gastropod-eating snake genera, Tropi- dodipsas and Sibon. ]. HerpetoL, 19:84-92.
Lee, J. C. 1980. An ecogeographic analysis of the herpetofauna of the Yucatan Pen- insula. Univ. Kansas Mus. Nat. Hist., Misc. Publ., 67:1-75.
Oliver, J. A. 1948. The relationships and zoogeography of the genus Thalerophis Oliver. Bull. Amer. Mus. Nat. Hist., 92:157-280.
Peters, J. A. 1953. Snakes and lizards from Quintana Roo, Mexico. Lloydia, 16:227- 232.
Schmidt, K, P., and A. S. Rand. 1957. Geographic variation in the Central American colubrine snake, Ninia sebae. Fieldiana: ZooL, 39:73-84.
Wellman, J. 1963. A revision of snakes of the genus Conophis (Family Colubridae, from Middle America). Univ. Kansas Publ., Mus. Nat. Hist., 15:251-295.
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ANNALS
0/ CARNEGIE MUSEUM
CARNEGIE MUSEUM OF NATURAL HISTORY
4400 FORBES AVENUE ® PITTSBURGH, PENNSYLVANIA 15213 VOLUME 55 23 MAY 1986 ARTICLE 7
KARYOTYPES OF ELEVEN SPECIES OF MOLOSSID BATS FROM AFRICA (MAMMALIA: CHIROPTERA)
Steven A. Smith ‘
John W. Bickham^
Duane A. Schlitter
Curator, Section of Mammals
Abstract
Standard karyotypic data are reported for 1 1 species of molossid bats collected from Somalia and Cameroon, Africa. Chromosomal data are reported for the first time for Chaerephon ansorgei, C. aloysiisabaudiae, Mops midas, M. spurrelli, M. thersites, M. brachypterus, M. petersoni, M. demonstrator, and M. nanulus (all were formerly mem- bers of the genus Tadarida). Karyotypes for two of the species we examined have been reported previously. Although our data corroborate the karyotype of C. pumila described by Dulic and Mutere (1973), our karyotypic analysis of M. condylurus differs substantially from that presented by these authors. In addition to these data, we provide a summary of the available karyotypic data for molossid bats studied to date.
Introduction
The Molossidae is a group of insectivorous, swift-flying bats that live in tropical and temperate parts of the world. More than half of the 9 1 or so extant species have been regarded as members of the genus Tad- arida (Corbet and Hill, 1980); the remaining species are spread among
* Address: Department of Wildlife and Fisheries Sciences, Texas A&M University, Col- lege Station, Texas, 77843.
Submitted 10 June 1985.
125
126
Annals of Carnegie Museum
VOL. 55
Table Summary of karyotype morphology for 11 species of African molossid bats. Letter designations are: M—metacentric, SM—submetacentric, ST-subtelocentric, A —
acrocentric.
|
Species |
Large M |
Medium M |
Medium ST |
Small ST |
Medium- small A |
X |
Y |
FN |
|
Chaerephon ansorgei Chaerephon aloysiisa- |
1 |
3 |
4 |
2 |
13 |
ST |
A |
66 |
|
baudiae |
1 |
3 |
4 |
2 |
13 |
SM |
A |
66 |
|
Chaerephon pumila |
1 |
2 |
3 |
0 |
17 |
SM |
A |
58 |
|
Mops midas |
1 |
3 |
4 |
2 |
13 |
SM |
A |
66 |
|
Mops condylurus |
1 |
3 |
4 |
2 |
13 |
SM |
A |
66 |
|
Mops spurrelli |
1 |
3 |
4 |
1 |
14 |
SM |
_ |
64 |
|
Mops thersites |
1 |
3 |
3 |
1 |
15 |
SM |
ST |
62 |
|
Mops brachypterus |
1 |
3 |
0 |
0 |
19 |
SM |
_ |
54 |
|
Mops petersoni |
1 |
3 |
0 |
0 |
19 |
SM |
A |
54 |
|
Mops demonstrator |
1 |
2 |
1 |
0 |
19 |
SM |
A |
54 |
|
Mops nanulus |
1 |
2 |
1 |
0 |
19 |
SM |
A |
54 |
1 1 Other genera. Until recently, taxonomic assignments and systematic relationships among the family members had not been examined world- wide. Freeman (1981), based upon a phenetic study of morphological traits, provided the first major review of the family. She restricted the genus Tadarida to include only nine species and assigned the remainder to Chaerephon, Mops, Mormopterus and Nyctinomops (all former sub- genera of Tadarida).
Karyotypic data for the Molossidae are available for 25 species, only six of which are inhabitants of the Old World. In this paper we analyze the karyotypes of 1 1 African molossid species belonging to the genera Chaerephon and Mops, and summarize the chromosomal data (Tables 1 and 2) now available for 35 species representing 10 of the 12 genera recognized by Freeman (1981).
Methods and Materials
Standard karyotypes were obtained in the field from bone marrow preparations (Patton, 1967) of live caught animals. A minimum of five representative chromosome spreads were examined from each individual to determine diploid (2n) and fundamental numbers (FN). Photomicrographic enlargements of suitable spreads were used in the final analyses.
Chromosomes were divided into large and medium-sized metacentric, medium and small subtelocentric, and medium to small acrocentric morphological classes. Deter- mination of centromere position was difficult because differential contraction of nearly acrocentric chromosomes caused variation in the number of countable arms. We follow Warner et al. (1974) in being conservative in the determination of biarmed versus acrocentric conditions and reiterate their warning that FN values are somewhat arbitrary and subjective.
Taxonomic designations follow Honacki et al., 1982 (see Freeman, 1981).
1986
Smith et al.— Molossid Karyotypes
127
|
Xir XX XK XX M MtK |
Ad |
|
do dd dl dh All |
XY |
|
fkA M |
A |
|
X* tn A XX M lA M M M |
A.-. V V |
|
do ftD ot If« An |
A T |
|
B |
|
|
^ 0 XX M /M M M A* AM |
l/t |
|
X Y |
|
|
c |
|
|
W X« xr> ft firt lifi |
An |
|
OH nn n/% rfikM aa «a |
X Y |
|
D |
Fig. L— Representative karyotypes of A) Chaerephon ansorgei from Cameroun, B) Chae- rephon aloysiisabaudiae from Cameroun, C) Mops midas from Somalia, and D) Mops condylurus from Somalia.
Species Accounts
A summary of the chromosomal morphology for the species ex- amined in this study is presented in Table 1. Representative karyotypes are presented in Figs, 1-3,
All species examined in this study were characterized by a diploid
128
Annals of Carnegie Museum
VOL. 55
number of 48. Fundamental numbers ranged from 54 to 66. All of these species had, minimally, four biarmed autosomal elements in- cluding one large pair of metacentric and at least one medium-sized pair of metacentric |Chromosomes. In all cases the large metacentric pair was twice the size of the next largest chromosome pair. The X chromosomes were medium-sized and submetacentric or subtelocen- tric in all species; the Y chromosome was medium-sized and acrocentric in all but one species.
A brief description of the karyotypes for each species reported herein follows.
Chaerephon ansorgei (Thomas, 1913)
Fig. lA, 2n = 48; FN = 66; U
The autosomal complement includes one pair of large metacentric, three pairs of medium metacentric, four pairs of medium subtelocen- tric, and 1 3 medium to small acrocentric chromosomes. The X chro- mosome is medium-sized and subtelocentric, and the Y is medium- sized and acrocentric.
Chaerephon aloysiisabaudiae (Festa, 1907)
Fig. IB, 2n = 48; FN = 66; IS
The karyotype of this species is identical to C ansorgei except the X chromosome in C aloysiisabaudiae appears submetacentric rather than subtelocentric.
Chaerephon pumila (Cretzschmar, 1826)
2n = 48; FN = 58; SSS, 6$9
The karyotype of our specimens is identical to that reported for this species by Dulic and Mutere (1973).
Mops midas (Sundevall, 1843)
Fig. 1C, 2n = 48; FN = 66; 3SS, 399
This species is karyotypically identical to the above-mentioned Chaerephon species and shares the submetacentric condition of the X chromosome observed in C aloysiisabaudiae.
Mops condylurus (A. Smith, 1833)
Fig. ID, 2n = 48; FN = 66; 4SS, 599
The karyotype of M. condylurus is identical to both M. midas and C aloysiisabaudiae.
1986
Smith et al.— Molossid Karyotypes
129
Mops spurrelU (Dollman, 1911)
Fig. 2A, 2n - 48; FN - 64; 355
The chromosomal complement from female specimens of M. spur- relU differ from M. condyiurus in the absence of one less small sub- telocentric pair and the presence of an extra acrocentric pair. The X chromosome is submetacentric.
Mops thersites (Thomas, 1903)
Fig. 2B, 2n = 48; FN ^ 62; 355, 499
The autosomes are nearly identical to M. spurrelU but there is one less medium-sized subtelocentric and one additional acrocentric pair present. The X chromosome is submetacentric but the Y appears to be subtelocentric instead of the more commonly observed acrocentric condition.
Mops bmchypterus (Peters, 1852)
Fig. 3 A, 2n - 48; FN - 54; 19
The autosomes of the female specimen examined consist of one large pair of metacentric, three pairs of medium-sized metacentric and 1 9 medium to small acrocentric pairs. Although morphologically similar to M. thersites, it differs chromosomally by lacking subtelocentric pairs and having additional acrocentric pairs. The X chromosome presum- ably is submetacentric.
Mops petersoni (El Rayah, 1981)
Fig. 3B, 2n - 48; FN - 54; 15, 19
In addition to being morphologically similar, this species is karyo- typically identical to M. brachypterus.
Mops nanuius J. A. Allen, 1917
Fig. 3C, 2n - 48; FN - 54; 255, 299
M. nanuius differs from M. petersoni and M. brachypterus by having one less medium-sized metacentric pair and the presence of a medium- sized subtelocentric pair. The sex pair is identical to M. petersoni.
Mops demonstrator 1903)
Fig. 3D, 2n = 48; FN - 54; 15
The karyotype of this species is identical to M. nanuius.
Discussion
Until Freeman’s (1981) recent revision of the Molossidae, phylo- genetic relationships and taxonomic assignments within the family
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Annals of Carnegie Museum
VOL. 55
|
M SK AA Ad lA Ad AA |
AK |
|
M tt 40 M «• A« It* |
X X |
|
4A §• • «« |
A |
|
XS Kt n M AA A . |
Y V |
|
f/A (^6 Ofl A0 AO 04 |
A T |
|
B |
Fig. 2.— Representative karyotypes of A) Mops spurrelli from Cameroun, B) Mops ther- sites from Cameroun.
were largely unexplored. Warner et al. (1974) suggested that chromo- somal studies might be beneficial in evaluating these relationships. Chromosomal data now available for 36 molossid species representing 10 of 12 genera recognized by Freeman (1981) are summarized in Table 2.
We detected no intraspecific chromosomal variation within any of the species examined in this study. This is noteworthy for two reasons. First, our karyotypes of M. condylurus (FN = 66) from Afgoi, Somalia, differ substantially from the karyotype of this species (FN = 56) re- ported from Kisubi, Uganda, by Dulic and Mutere (1973). These lo- calities are several hundred kilometers apart and this suggests either that considerable geographic variation in the karyotype occurs within this species or there are two species currently recognized as M. con- dylurus. Secondly, our data support the specific distinctiveness of M. spurrelli and M. nanulus. Freeman (1981) recognized the morpholog- ical similarity between these two taxa and noted Koopman’s (1975) suggestion that they might be conspecific. Our data indicate that M. nanulus (FN = 54) and M. spurrelli (FN = 66) differ by five pairs of biarmed chromosomes, and considering the scarcity of intraspecific karyotypic variation within this family, it would seem likely that the two taxa are specifically distinct.
Variation in FN for the species we examined ranged from 54 to 66 (Table 1). These karyotypes can be conveniently divided into three
1986
Smith et al.— Molossid Karyotypes
131
CC n IK
lA M fn OA Ifi lA M
AR RR (ft (# (ft %( ••
^A^XX' ««
00 AA AA A« M NO Ml M ••
#11 ^41 HA AA A# A# lO* *>« *•>
ITtlUil
il AOONNAI'^ MM M
ii tH »• »l» W ti oo
n/lf Mn
AA tfl HA Ail at «# at Ai ft
U
A
(#
B
X Y
c
h
XY
D
Fig. 3.— Representative karyotypes of A) Mops brachyptems from Cameroon, B) Mops petersoni from Cameroun, C) Mops nanulus from Cameroun, and D) Mops demonstrator from Cameroun.
groups. The high FN group (FN 62-66) includes both species of Chaerephon and four of eight Mops species. Within this group, differ- ences between the FN = 62-66 karyotypes apparently involve the absence of medium and small subtelocentric autosomes. Our exami- nation of Chaerephon pumila (FN ^58) agrees with the karyotype of this species reported by Dulic and Mutere (1973), and forms an inter- mediate FN group. Again, differences between the intermediate and
132
Annals of Carnegie Museum
VOL. 55
Table 2.— Summary of molossid karyotype data.
|
Species |
In |
FN |
Reference |
|
Chaerephon aloysiisabaudiae'^ |
48 |
66 |
This study |
|
Chaerephon ansorgeP |
48 |
66 |
This study |
|
Chaerephon bivittata^ |
48 |
54 |
Peterson and Nagorsen, 1975 |
|
Chaerephon plicata^ |
48 |
54 |
Harada and Kobayashi, 1980; Harada et al., 1982 |
|
Chaerephon pumila^ |
48 |
58 |
Dulic and Mutere, 1973; this study |
|
Eumops auripendulus |
42 |
62 |
Warner et al., 1974 |
|
Eumops glaucinus |
38 |
64 |
Warner et al., 1974 |
|
40 |
64 |
Warner et al., 1974 |
|
|
Eumops perotis |
48 |
56 |
Baker, 1970; Warner et al, 1974 |
|
48 |
58 |
Wainberg et al., 1974 |
|
|
Eumops underwoodi |
48 |
56 |
Warner et al., 1974 |
|
Molossops abrasus |
34 |
60 |
Warner et al., 1974; Gardner, 1977 |
|
Molossops greenhalli |
34 |
_ |
Linares and Kiblisky, 1969 |
|
34 |
60 |
Baker, 1970; Warner et al., 1974 |
|
|
Molossops temminckii |
42 |
56 |
Gardner, 1977 |
|
Molossus ater |
48 |
58 |
Warner et al., 1974 |
|
Molossus molossus |
48 |
56 |
Baker and Lopez, 1970 |
|
48 |
58 |
Warner et al., 1974 |
|
|
Molossus rufus |
48 |
58 |
Wainberg et al., 1974 |
|
Molossus Sinaloa |
48 |
58 |
Warner et al., 1974 |
|
Mops brachypterus' |
48 |
54 |
This study |
|
Mops condylurus^ |
48 |
66 |
This study |
|
48 |
56 |
Dulic and Mutere, 1973 |
|
|
Mops demonstrator^ |
48 |
54 |
This study |
|
Mops midas^ |
48 |
66 |
This study |
|
Mops nanulus^ |
48 |
54 |
This study |
|
Mops petersonP |
48 |
54 |
This study |
|
Mops spurrellP |
48 |
64 |
This study |
|
Mops thersites^ |
48 |
62 |
This study |
|
Mormopterus kalinowskii’^ |
48 |
56 |
Warner et al., 1974 |
|
Mormopterus setiger'^ |
48 |
54 |
Warner et al., 1974 |
|
Nyctinomops aurispinosus^ |
48 |
58 |
Warner et al., 1974 |
|
Nyctinomops femorosacus^ |
48 |
58 |
Warner et al., 1974 |
|
Nyctinomops laticaudatus^ |
48 |
58 |
Warner et al., 1974 |
|
Nyctinomops macrotis^ |
48 |
58 |
Warner et al., 1974 |
|
48 |
56 |
Baker, 1970 |
|
|
Otomops martiensseni |
48 |
56 |
Dulic and Mutere, 1973 |
|
Promops centralis |
48 |
58 |
Warner et al., 1974 |
|
Promops nasutus |
40 |
54 |
Wainberg, 1966 |
|
Tadarida brasiliensis |
48 |
__ |
Painter, 1925 |
|
48 |
54 |
KniazefF et al., 1967 |
|
|
48 |
56 |
Warner et al., 1974; Baker et al, 1982 |
|
|
Tadarida fulminans |
48 |
54 |
Peterson and Nagorsen, 1975 |
* Indicates species formerly recognized as Tadarida, see Freeman (1981).
1986
Smith et al.— Molossid Karyotypes
133
high FN forms appear to be in the absence of small subtelocentric pairs plus the absence of one pair of medium-sized metacentrics. The low FN group (FN =54) includes M. brachypterus, M. peter soni, M. dem- onstrator, and M. nanuius. Karyotypic morphologies of the latter two are identical and differ from the former pair in having one fewer me- dium-sized metacentric pair and an additional pair of medium-sized subtelocentric chromosomes.
Whether or not these karyotype associations reflect phylogenetic re- lationships within genera is difficult to assess from standard karyotypic data. Phylogenetic interpretations based on chromosomal data nec- essarily require identification of homologous pairs using differential staining techniques (see Haiduk et al., 1981). There is, however, little concordance between Freeman’s (1981) phenetic classification and the patterns of karyotypic morphology for these species. Within Chaere- phon, Freeman’s (1981) analysis clusters C. ansorgei (FN = 66) with C. bivittata (FN = 54, Peterson and Nagorsen, 1975). C aloysiisabau- diae (FN = 66) then joins the cluster followed by C. pumila (FN = 58, Dulic and Mutere, 1973; this study) several junctures later, and further still, by C plicata (FN = 54, Harada and Kobayashi, 1980; Harada et al., 1982). Similarly, within the genus Mops, M. demonstrator (FN = 54) clusters phenetically with M. condylurus (FN = 66); M. brachyp- terus (FN = 54) and M. ther sites (FN = 62) pair together. These dis- parities suggest the possibility that either morphological and chro- mosomal characters are evolving at different rates, or that the taxonomic relationships of these taxa need to be reexamined.
Karyotypic stability for bats, in general, has been recognized by several authors (Peterson and Nagorsen, 1975; Gardner, 1977; Baker, 1978; Baker et al., 1982; Baker and Bickham, 1980; Bickham, \919a, \919b\ Bickham and Baker, 1979) and has been suggested for the Molossidae, specifically, by Warner et al. (1974) and Dulic and Mutere (1973). Of the 36 molossid species for which chromosomal data are available only seven species have diploid numbers other than 2n = 48 (Table 1). The modal occurrence of 2n = 48 chromosomes in both Old and New World genera plus the similarity between this number and the proposed primitive diploid number for the Vespertilionidae (Baker, 1970) led Warner et al. (1974) to propose 2n = 48 as primitive for the Molossidae. Our documentation of the 2n = 48 karyotype in 1 1 Old World molossid species further supports this diploid value as primitive for the family.
Specimens Examined
Chaerephon aloysiisabaudiae. — Cameroun: 1 6 km S, 2 km E Yaounde (3®43'N, 1 1°32'E), (16 CM 58678).
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Annals of Carnegie Museum
VOL. 55
Chaerephon ansorgei. —Cameroun: 25 km S, 13 km E Garoua, (9°05'N, 13®30'E), (13 CM 58679).
Chaerephon Cameroon: 24 km S, 13 km E Garoua (9®05'N, 13®30'E), (13
CM 58724); Somalia: Libsoma Farm 6 km S, 17 km W Afgoi (2®05'N, 44®58'E), (333 CM 85438, 85439-85440; 299 CM 85441-85442); Somalia: Bulo Burti (3®51'N, 45®34'E), (433 CM 85455-85456, CM 85459-85460; 499 CM 85457-85458, 85461-85462).
Mops brachypterus.—CAMEROvn: 25 km S, 3 km E Yaounde (3®38'N, ir33'E), (19 CM 58687).
Mops condylurus.—^OMAEw: Libsoma Farm, 6 km S, 17 km W Afgoi (2®05'N, 44®58'E), (433 CM 85423, 85408-85409, 85424; 599 CM 85410-85411, 85412, 85425-85426).
Mops demonstrator.— CAMEROvn: 2 km W Ngaoundere (7®20'N, 13®34'E), (13 CM 58681).
Mops midas. —Somaeia: Libsoma Farm, 6 km S, 17 km W Afgoi (2°05'N, 44®58'E), (333 CM 85429, 85436, 85428; 399 CM 85427, 85430).
Mops nanulus.—CAMEROuw. 25 km S, 13 km E Garoua (9®05'N, 13®30'E), (13 CM 58694); Cameroon: 24 km S, 13 km E Garoua (9®05'N, 13®30'E), (13 CM 58692; 299 CM 58693, CM 58695).
Mops petersoni. —Cameroun: 25 km S, 3 km E Yaounde (3®38'N, 11°33'E), (13 CM 58688; 19 CM 58691).
Mops spurrel/i.—CAMEROUN: 30 km N, 40 km E Obala (4®22'N, 11°58'E), (13 CM 58730); Cameroon: 25 km S, 3 km E Yaounde (3®38'N, 1 1®33'E), (233 CM 58731, CM 58786).
Mops thersites.—CAMEROUN: 30 km N, 40 km E Obala (4°22'N, 1 1®58'E), (13 CM 58737); Cameroon: 25 km S, 3 km E Yaounde (3°38'N, 1 1°33'E), (299 CM 58743, CM 58745); Cameroon: 7 km S, 8 km W Yaounde (3°48'N, 1 r27'E), (233 CM 58739, CM 58741; 299 CM 58740, CM 58742).
Acknowledgments
Field work in Somalia was supported by a grant from the M. Graham Netting Research Fund, Cordelia Scaife May Charitable Trust and the Hays Fund of the American Philo- sophical Society. Assistance in the field was provided by M. J. Smolen, R. Ruiz, Omar Hagi, Abdulwahab Josuf, and Mohamed Ali. Facilities and support in Somalia was graciously supplied by Mohamet Abdi Nur, Minister of Agriculture; Mohamed Abikar, Director, General Ministry of Agriculture; and Abdulcadir Nur, Director, Department of Plant Protection and Locust Control in the Ministry of Agriculture. Abdullahi Ahmed Karani, General Manager, National Range Agency, issued the necessary permits to work in Somalia. John and Jonquil Ash; Bill and Sally Smythe; Tony and Lynette Johnston, United Nations Development Program in Somalia, graciously assisted in numerous ways. Our special thanks to all these individuals.
Field work in Cameroun was supported by grants from the M. Graham Netting Re- search Fund, Cordelia Scaife May Charitable Trust; the Loyalhanna Foundation; and the National Geographic Society. Assistance in the field was provided by L. W. Robbins, R. L. Robbins, and S. L. Williams. Permission to conduct field work and collecting permits were received from the Ministry of Agriculture. We are indebted to Mr. V. Belinga, Director of Forestry Services in Cameroun and Mr. C. Njiti for assistance in obtaining the necessary permits. Laboratory work was supported by NSF grant No. PCM- 8202794 to J. W. Bickham. Partial financial support for field work was received from NIH Grant AIO 4242 to R. Traub.
Literature Cited
Baker, R. J. 1970. Karyotypic trends in bats. Pp. 65-96, in Biology of bats (W. A.
Wimsatt, ed.). Academic Press, New York, 406 pp.
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■ . 1978. Karyology. Pp. 107-155, in Biology of bats of the New World family Phyllostomatidae, Part III (R. J. Baker, J. K. Jones, Jr., and D. C. Carter, eds.). Spec. Publ. Mus., Texas Tech Univ., 16:1-441.
Baker, R. J., and J. W. Bickham. 1980. Karyotypic evolution in bats: evidence of extensive and conservative chromosomal evolution in closely related taxa. Syst. ZooL, 29:239-253.
Baker, R. J., M. W. Haiduk, L. W. Robbins, A. Cadena, and B. F. Koop. 1982. Chromosomal studies of South American bats and their systematic implications. Spec. Publ. Ser. Pymatuning Lab. EcoL, 3:303-327.
Baker, R. J., and G. Lopez. 1970. Karyotypic studies of the insular populations of bats of Puerto Rico. Caryologia, 23:465-472.
Bickham, J. W. 1979a. Chromosomal variation and evolutionary relationships of vespertilionid bats. J. Mamm., 60:350-363.
Bickham, J. W. 1979^. Banded karyotypes of 11 species of American bats (genus Myotis). Cytologia, 44:789-797.
Bickham, J. W., and R. J. Baker. 1 979. Canalization model of chromosomal evolution. Bull. Camagie Mus. Nat. Hist., 13:70-84.
Corbet, G. B., and J. E. Hill. 1980. A world list of mammalian species. British Mus. (Nat. Hist.), London, viii + 226 pp.
Dulic, B., and F. a. Mutere. 1973. Comparative study of the chromosomes of some molossid bats from eastern Africa. Period. Biol., 75:61-65.
Freeman, P. W. 1981. A multivariate study of the family Molossidae (Mammalia, Chiroptera): morphology, ecology, evolution. Fieldiana ZooL, Chicago Mus. Nat. Hist. 1316 n.s. 7.
Gardner, A. L. 1977. Taxonomic implications of the karyotypes of Molossups and Cynomops (Mammalia:Chiroptera). Proc. Biol. Soc. Washington, 89:545-550. Haiduk, M. W., R. J. Baker, L. W. Robbins, and D. A. Schlitter. 1981. Chromo- somal evolution in African Megachiroptera: G- and C-band assessment of the mag- nitude of change in similar standard karyotypes. Cytogenet. Cell Genet., 29:221- 232.
Harada, M., and T. Kobayashi. 1980. Studies on the small mammal fauna of Sabah, East Malaysia II. Karyological analysis of some Sabahan mammals (Primates, Ro- dentia, Chiroptera). Contrib. Biol. Lab. Kyoto. Univ., 26:83-95.
Harada, M., M. Minezawa, S. Takada, S. Yenbutra, P. Nunpakdee, and S. Ohtani. 1982. Karyological analysis of 12 species of bats from Thailand. Caryologia, 35: 269-278.
Honacki, j. H., K. E. Kinman, and J. W. Koeppl (eds.). 1982. Mammal species of the world. Allen Press, Inc., and the Association of Systematics Collections, Law- rence, Kansas, 694 pp.
Kniazeff, a. j., D. Constantine, W. A. Nelson-Rees, D. Schmidt, and R. Owens. 1967. Studies in chiropteran cell lines. 41st. Tech. Progr. Rept. Naval Biol. Lab. Suppl. Rept, CC-8:97-105.
Koopman, K. F. 1975. Bats of the Sudan. Bull. Amer. Mus. Nat. Hist., 154:354-444. Linares, O. J., and P. Kibilisky. 1969. The karyotype of a new record of Molossups greenhalli from Venezuela. J. Mamm., 50:831-832.
Painter, T. S. 1925. A comparative study of the chromosomes of mammals. Amer. Nat, 59:385-408.
Patton, J. L. 1967. Chromosome studies of certain pocket mice, genus Perognathus (Rodentia: Heteromyidae). J. Mamm., 48:27-37.
Peterson, R. L., and D. W. Nagorsen. 1975. Chromosomes of fifteen species of bats (Chiroptera) from Kenya and Rhodesia. Occas. Papers, Royal Ontario Mus. Life ScL, 27:1-14.
Wainberg, R. L. 1966. Cytotaxonomy of South American Chiroptera. Arch. Biol. (Liege), 77:411-423.
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Wainberg, R. L., L. H. Delupi de Bianchini, J. J. Bianchini, and G. E. Pollero de Actis Dato. 1974. Uniformidad cariotipica y radiacion adaptativa en “Eumops” y “Molossus” (Chiroptera, Molossidae). Physis Secc. C Cont. Org. Terr., 33:249-- 254.
Warner, J. W., J. L. Patton, A. L. Gardner, and R. J, Baker. 1974. Karyotypic analyses of twenty-one species of molossid bats (MolossidaeiChiroptera). Canadian J. Genet. Cytol., 16:165-176.
^ ISSN 0097-4463
ANNALS
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VOLUME 55 23 MAY 1986 ARTICLE 8
TRILOBITES FROM THE KEOKUK LIMESTONE (MISSISSIPPIAN) OF MISSOURI
David K. Brezinski^
Research Associate, Section of Invertebrate Fossils
Abstract
A new species of the trilobite genus Griffithides Portlock, G. salinensis, new species, is described. G. salinensis is a rare component of the fauna of the Keokuk Limestone (Mississippian) of St Genevieve County, Missouri. It can be distinguished from other described species assigned to this genus by the posterior termination of the pygidial axis, which is upturned into a large node-like projection. The type specimens of this species were collected from just below the Marginarugus magnus bed of the Keokuk Limestone. In addition to G. salinensis, two specimens also recovered from the same bed are de- scribed and tentatively assigned to the genus Waribole Richter and Richter.
Introduction
The trilobite genus Griffithides Portlock is relatively poorly known from Carboniferous rocks of North America as compared to the many species recognized from correlative rocks of Europe. Of the three North American species recognized, only one, G. bufo Meek and Worthen, is known from more than just the type material. In contrast, specific diversity exhibited by this genus in Europe has prompted some authors (Hahn and Hahn, 1 970, 1971; Hahn et al., 1983) to subdivide the genus
* Present address: The Maryland Geological Survey, The Rotunda —Suite 440, 71 1 W. 40th Street, Baltimore, MD 21211.
Submitted 21 August 1985.
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into a number of subgenera. The recent recovery from the Keokuk Limestone of Missouri of a small number of specimens of Griffithides, which are notably dissimilar to previously recognized species, warrants the erection of a new species. The specimens of Griffithides, and two pygidia tentatively assigned to the genus Waribole were recovered from a light "gray lime grainstone in the Keokuk Limestone along Interstate 55 at Little Saline Creek, St. Genevieve County, Missouri. The spec- imens were recovered as accessories during field collections made for brachiopods by J. L. Carter and A. D. Kollar (Carnegie Museum of Natural History) from a bed just below the Marginarugus magnus bed or about in the middle of the Keokuk. The brachiopod fauna associated with these trilobites is dominated by Productus crawfordsvillensis Well- er, Imbrexia montonana Miller, Rhynchopora beecheri Greger, and Torynifer pseudolineatus (Hall). The lithologic character of the strata, which yielded the trilobites, is similar to that in which other North American species of Griffithides have been found. Moreover, the spec- imens of Waribole? recovered from the Keokuk are similar to pygidia I have recovered from the lime grainstones of the Salem Limestone of southern Indiana. These specimens from the Salem are also found in association with a species of Griffithides. The consistent occurrence of these two trilobite genera together in particular lithologies may suggests a strong ecologic control on their distribution.
Terminology employed in this study follows that utilized by Har- rington (1959).
Systematic Paleontology
Family Proetidae Salter Subfamily Griffithidinae Hupe Genus Griffithides Portlock
Distribution of North American Present in the Keokuk
Limestone (Osagean) of Missouri and Illinois, the Salem Limestone (Meramecian) of Indiana, and the Pitkin Limestone (Chesterian) of Oklahoma.
Diagnosis of North American representatives. —Ctphdilon parabolic in outline, moderately vaulted. Glabella pyriform with frontal lobe moderately to greatly expanded laterally and reaching the anterior mar- gin of the cranidium. Lateral preoccipital lobes well-defined, subtrian-
Fig. \. — Griffithides salinensis, new species, A, C, E, holotype pygidium in dorsal, lateral, and posterior views, CMNH 34553, x2; B, D, paratype pygidium in dorsal and lateral views, CMNH 34498, x2; F, paratype pygidium, dorsal view, CMNH 34499, x2.5; G, H, I, paratype cranidium in dorsal, lateral and anterior views, CMNH 34500, x2.5.
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Fig. 2.— Bivariate plot of maximum pygidial width (W) vs maximum pygidial length (Z) for specimens of Griffithides salinensis, new species, and G. bufo Meek and Worthen.
gular, Ip furrow narrow, deeply incised, widened anterolaterally. Pal- pebral lobes crescentic to semicircular in outline. Eyes of medium size, hemispherical in shape. Lateral border furrow well-defined and narrow, margin sharply rounded. Genal spines short, generally reaching to the third thoracic segment.
Thorax of nine segments, axial rings semicircular in transverse pro- file, ornamented by a row of small granules along the posterior margin. Pleurae sharply rounded at fulcrum, subangular at tip.
Pygidium semicircular to parabolic in outline, strongly vaulted, con- sisting of an axis composed of 1 1-15 rings and pleural fields of 9-14 ribs. Axis tapers posteriorly, does not reach posterior margin, strongly convex and steeply downsloping posteriorly. Pleural ribs extend nearly to the margin, with no well-defined border.
Discussion. — Although Hahn et al. (1983) were able to subdivide the Eurasian representatives of the genus Griffithides into three subgenera, no such division of North American species is possible. The main distinguishing features among North American species of Griffithides lie mainly in the shape (suboval versus parabolic) and structure (num- ber of ribs and rings) of the pygidium. Griffithides can be distinguished from the contemporary trilobite genus Paladin by the presence of a well-developed pygidial border on the latter.
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Table \. — Univariate measurements of select morphological characters of Griffithides salinensis, new species. For discussion and illustration of character definition see Shaw
(1957).
|
Character |
N |
Mean* |
Range* |
|
(W) pygidial width |
7 |
16.2 |
12.7^19.6 |
|
(Z) pygidial length |
7 |
13.4 |
10.1-16.8 |
|
(X) axial width (max.) |
7 |
6.3 |
4.9-7.4 |
|
(Y) axial length |
6 |
12.2 |
8.4-14.7 |
|
number of axial rings |
6 |
15 |
14-15 |
|
number of axial ribs |
5 |
14 |
13-14 |
* Measurements in millimeters.
Griffithides salinensis, new species Figs. 1A~1I
Holotype. -CMNn 34553.
Paratypes.~CMN¥L 34498-34500.
Material ~\0 incomplete pygidia and 1 partial cranidium.
Description.— pyriform with moderate lateral expansion to the frontal lobe. Frontal lobe strongly convex in transverse profile and extends to the anterior margin. In longitudinal profile glabella is nearly flat to mildly convex at the posterior terminus, becoming increasingly convex anteriorly, meeting the anterior margin vertically. Glabella covered by fine granules. Lateral preoccipital lobes subtriangular with granular ornament; Ip furrow well-defined and of medium width, becoming broader toward the dorsal furrow. Palpebral lobes of medium size, semicircular in outline, inclined into the dorsal furrow at about 45® Facial sutures mildly divergent from a to 0, rounded at wider at co than a. Occipital lobe not preserved.
Thorax is unknown.
Pygidium parabolic in outline, moderately vaulted, .83 times as long as wide. Axis tapers posteriorly, .91 the total pygidial length, ,39 the total pygidial (anterior) width, composed of 14 to 15 rings which are semicircular in transverse profile. Posteriormost axial ring is enlarged into a large node or nub that overhangs a slightly concave axial terminus. Each ring is slightly sinuous, being posteriorly bent across axis. A row of 1 2 fine granules ornament the posterior edge of each ring. Pleural fields strongly convex, made up of 13--14 posteriorly recurved ribs that extend nearly to the margin. The anteriormost three or four ribs exhibit a well-defined pleural furrow. A row of fine granules ornament each rib.
Discussion. — G. salinensis, new species, can readily be distinguished from other North American species of the genus by the upturned node- like termination of the pygidial axis, by the greater length to width ratio to the pygidium, and by the greater number of axial rings and pleural ribs. Only G. bufo Meek and Worthen has been recovered in sufficient number to allow any close comparison. Fig. 2 is a bivariate plot comparing the maximum pygidial widths with the maximum py- gidial lengths for specimens of G. salinensis and G. bufo. There is a noticeable difference in the rectilinear trends exhibited by each species.
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Fig. 3.— Waribolel sp. A, B, dorsal and posterior views of complete pygidium, CMNH 34501, x.2.5.
The plot illustrates that G. salinensis possesses a greater length to width ratio than does G. bufo and is larger on the average. Unfortunately, insufficient numbers of G. salinensis are available at present to produce a reliable regression equation. G. salinensis differs from the poorly known G. meramecensis Shumard in that the latter exhibits fewer pygidial ribs and rings (12 and 13, respectively) and lacks the node- like termination of the axis. Both G. meramecensis and G. salinensis were recovered from the Keokuk Limestone of Missouri. The com- parison presented above is based upon the drawing and description of G. meramecensis presented by Shumard (1855). Inasmuch as the lo- cation of the holotype is unknown, comparison must be based upon Shumard’s description and drawing alone. G. salinensis differs from G. pustulosus Snider in that the pygidium of the former exhibits a parabolic outline and the prominent terminal node on the pygidial axis. All Eurasian species of Griffit hides can be distinguished from G. sali- nensis by the terminal axial node on the pygidium.
Subfamily Cyrtosymbolinae Hupe Genus Waribole Richter and Richter Waribolel sp.
Fig. 3A, 3B
Material. — 1 complete and 1 fragmented pygidium from the Keokuk Limestone. IL lustrated specimen CMNH 34501.
Description.— with low vaulting and relief, semicircular in outline with a length/ width ratio of .65. Axis tapers posteriorly, is .85 the total pygidial length and .37 the maximum (anterior) pygidium width, composed of 1 1 rings and terminates, poste- riorly, at inside margin of border. Pleural areas composed of six or perhaps seven ribs which become increasingly obsolete posteriorly. Each rib composed of two bands of approximately equal width. Border well-developed, smooth, and slightly concave to the margins, of nearly equal width all along pygidium.
Discussion. — The genus Waribole is most common in Late Devonian rocks and has a documented range into the earliest Carboniferous. The genus, to the best of my knowledge, has not been definitely documented
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from the stratigraphic interval of the Keokuk Limestone (that is, Vi- sean). Moreover, Waribole has not previously been reported from North America. If these specimens are in fact not specimens of Waribole, then they belong to some very similar genus. At present insufficient specimens are available to make any further inference. The only other North American trilobite genus with pygidial characteristics somewhat similar to these two pygidia is Richterelia; Richter ella, however, ex- hibits a much greater vaulting to the pygidium and the pleural ribs lack the subdivision into anterior and posterior bands.
Acknowledgments
All specimens utilized in this study are reposited in collections in the Section of Invertebrate Fossils, Carnegie Museum of Natural History (CMNH). Thanks are owed to Dr. John L, Carter and Albert Kollar who collected the specimens and made them available to me. Drafts of the manuscript were read by J. L. Carter and C. A. Kertis, and critically reviewed by J. H. Stitt and J. F. Taylor.
Literature Cited
Hahn, G., and R. Hahn. 1970. Trilobitae carbonici et permici 11. (Proetidae: Griffi- thidinae). Pp. 162-331, in Fossilium Catalogus 1. Animalia (F, Westphal, ed.), ’s Gravenhage (Dr. W. Junk N. V.), 1 19.
1971. Revision von Griffithides {Bollandia) (TriL: UnterKarbon). Palaeonto-
graph., 137:109-154.
Hahn, G. R. Hahn, and C. Brauckmann. 1983. Die Trilobitien des belgischen Koh- lenkalkes (Unter-Karbon) 5. Griffithides and Cyphinoides. Geol. et Paleont., 1 7: 109- 135.
Harrington, H. J. 1959. General description of trilobita. Pp. 38-1 17, Treatise on Inverterbrate Paleontology, Part O, Arthropoda 1 (R. C. Moore, ed.), Univ. Kansas Press, Lawrence, Kansas, 506 pp.
Shaw, A. B. 1957. Quantitative trilobite studies II. Measurement of the dorsal shell of non-agnostidean trilobites. J. Paleont., 31:193-207.
Shumard, B. F. 1855. Description of a geological section on the Mississippi River from St. Louis to Commerce. 1st and 2nd Annual Rept., Geol Surv. Missouri, 2:
85-208.
Back issues of many Annals of Carnegie Museum articles are available, and a few early complete volumes and parts are listed at half price. Orders and inquiries should be addressed to: Publications Secretary, Carnegie Museum, 4400 Forbes Avenue, Pittsburgh, Pa. 15213.
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VOLUME 55 15 SEPTEMBER 1986 ARTICLE 10
RESULTS OF THE ALCOA FOUNDATION SURINAME EXPEDITIONS. X. PATTERNS OF CELLULAR DIVERGENCE AND EVOLUTION IN THE GASTRIC MUCOSA OF TWO GENERA OF PHYLLOSTOMID BATS, TRACHOPS AND CHIRODERMA
Keith M. Studholme*’^
Carleton J. Phillips^
Research Associate, Section of Mammals
G, Lawrence Forman^
Abstract
The fundic mucosa in Trachops cirrhosus, Chiroderma trinitatum, and C villosum
(Suborder Microchiroptera) was studied comparatively by transmission electron mi- croscopy. Trachops is an animalivorous species that mainly feeds on Neotropical frogs, whereas both species of Chiroderma are frugivorous. In Trachops, two types of entero- endocrine cells (A and D cells) that possibly produce glucagon and somatostatin, re- spectively, were identified ultrastructurally. In Chiroderma examples of possible A, EC^, Di(H), and G cells were identified. The product in possible G-cells in Chiroderma ultra- structurally matched that found in the pylorus of another stenodermatine bat, Ariteus flavescens, which has been shown to exhibit gastrin-like immunoreactivity. In Trachops
' Address: Department of Biology, Hofstra University, Hempstead, New York 11550.
^ Present address: Department of Neurobiology and Behavior, SUNY Stony Brook, Stony Brook, New York 11794.
^ Address: Department of Biology, Rockford College, Rockford, Illinois 61101. Submitted 7 October 1985.
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the pepsin-producing chief cells are restricted to the basal-most portion of the gastric gland and produce an electron-dense product, whereas in Chiroderma the chief cells occupy up to 75% of the gland and produce a pale product. In Trachops the parietal cells are less abundant and less active than in Chiroderma but mucous neck cells are far more abundant. The gastric pits are shallow in Chiroderma, whereas Trachops has deep gastric pits. The mucous secretory granules in the surface mucous cells in Chiroderma differ ultrastructurally from those in Trachops. Overall, the ultrastructure of gastric mucosa in Trachops resembles that found in Phyllostomus (animalivorous/omnivorous) and Pteron- otus (insectivorous), whereas these features in Chiroderma resemble those of Artibeus (frugivorous). The general histology of the stomach of Chiroderma is similar to that of a megachiropteran, Eidolon helvum, suggesting that histological convergence has occurred in the evolution of the stomach of microchiropteran and megachiropteran fruit bats.
Introduction
Patterns of morphological evolution in bats are complex and un- doubtedly constrained by many factors (Hill and Smith, 1984). Among these, dietary specialization appears to explain many chiropteran mor- phological features. Dental morphology, salivary gland structure, kid- neys, gastric anatomy, relative brain size, jaws, and even length of forearm can be correlated with diet (Phillips, 1971; Phillips et al., 1977; Studier et al., 1983<2; Eisenberg and Wilson, 1978; Findley and Wilson, 1982; Freeman, 1979, 1981). Consequently, bats represent an outstand- ing mammalian model against which hypotheses about the relation- ships among structure, function, and biological role of various ana- tomical features can be tested.
The digestive tract, and the stomach in particular, is especially in- teresting in bats because diet can be correlated with gross anatomy, musculature, histochemistry, and histology (for example, Park and Hall, 1951; Kolb, 1954; Schultz, 1965, 1970; Rouk and Glass, 1970; Hart, 1971; Forman, 1971, 1972; Forman et al., 1979; Kamiya and Pirlot, 1975; Ogunbiyi and Okon, 1976; Okon, 1977; Bhide, 1980; Yamada et al., 1984; Ishikawa et al., 1985). These correlations have been ex- tended through recent studies in which we have demonstrated differ- ences in 1) aspects of the ultrastructure of cell types, 2) possible presence or absence of certain entero-endocrine (endocrine-paracrine) cells, and 3) relative numbers of particular cell types (Phillips and Studholme, 1982; Phillips et al., 1984; Mennone et al., 1 986). A variety of questions has been left unanswered by previous investigations. For example, we do not yet know exactly to what extent cellular and subcellular mor- phology differs in conjunction with diet or to what extent species with similar diets have similar cellular patterns, regardless of systematic associations.
For the present investigation we compared two genera of phyllos- tomid bats, Trachops (a phyllostomine) and Chiroderma (a stenoder- matine), that represent probable extremes in feeding specialization. We
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chose Trachops cirrhosus because it has been characterized as an ani-
malivorous species that feeds on small vertebrates, especially Neo- tropical frogs (Gardner, 1977; Tuttle and Ryan, 1981). As further evi- dence of feeding specialization, it has been shown that Trachops responds differentially to the vocal advertisements of edible and presumably non-edible (poisonous) anurans (Tuttle and Ryan, 1981) and has his- tologically unique accessory submandibular salivary glands that also might relate to its dietary habits (Phillips and Tandler, 1985; Phillips et al., in press). Chiroderma trinitatum and C viliosum are frugivores (Gardner, 1977), and Forman (1973) and Forman et al. (1979) have reported that the stomach of Chiroderma viliosum has an unusually large fundic caecum making it . perhaps the most extreme spe- cialization for plant feeding ...” among the phyllostomids. Insofar as previous cellular comparisons are concerned, Phillips and Studholme (1982) demonstrated a significant ultrastructural difference in chief cells in the fundic mucosa of representative frugivorous and carnivorous (insectivorous and animalivorous) bats, including Trachops and Chi- roderma. This ultrastructural difference was interpreted to mean that pepsinogen secretion is greatest in the frugivores.
The present investigation addressed the following questions. 1) How are dietary and structural specializations expressed at the cellular level in the gastric mucosa of Trachops and Chiroderma? 2) How do spe- cializations in Trachops and Chiroderma compare with other micro- chiropterans for which data are available? 3) How do ultrastructural features of the gastric mucosa in Chiroderma and other frugivorous microchiropterans compare with those found in Old World frugivores of the Suborder Megachiroptera?
Methods and Materials
Specimens of Chiroderma trinitatum (2 females; CM 77599, 77600), Chiroderma viliosum (3 females; CM 76796, 76798, 77601), and Trachops cirrhosus (1 male, CM 63688; 1 female, CM 64048) were collected in the vicinity of Rudi Kappelvliegveld, Brokopondo, Suriname. All specimens have been deposited in the collections of the Section of Mammals, Carnegie Museum of Natural History (CM). The bats were captured in nets at night (1700--2400 h) while feeding. Subsequently, they were kept overnight without food until between 0900 and 1 200 h when they were anesthetized with an intra- peritoneal injection of 0.25 cc of sodium pentabarbitol (50 mg/ml). A polyethylene tube next was inserted into the stomach via the mouth and esophagus and approximately 1 cc of trialdehyde fixative (at ambient temperature) was injected into the digestive tract. After approximately eight minutes, an incision was made into the abdomen and the stomach removed and cut into 2 by 2 mm samples that included portions of the fundus. Only samples from the fundus (Fig. 1) were used for the present report.
The fixation protocol was developed specifically for field projects involving transmis- sion electron microscopy (Forman and Phillips, in press; Phillips, 1985). The primary fixative (based on Kalt and Tandler, 1971) consisted of 3% glutaraldehyde, 1% parafor- maldehyde, 0.5% acrolein, 2,5% dimethyl sulfoxide (DMSO), and 1 mM CaCf in 0.05 M cacodylate buffer at pH 7.2 with 0. 1 M sucrose. All tissues were stored in this primary
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fixative for approximately 20 h at ambient temperature (30--40°C), The fixative then was replaced with fresh buffer (0.05 M cacodylate buffer, pH 7.2, with 0.1 M sucrose); tissues were left in this solution, unrefrigerated, for ten days. When refrigeration was available, the tissues were placed in fresh buffer with 3% glutaraldehyde and stored at 4®C. For processing, the tissues were washed for one hour in 0.05 M cacodylate buffer (pH 7.2) with 0.1 M sucrose and post-fixed for one hour in 1% OSO4 with cacodylate buffer and sucrose. Tissues then were dehydrated in an alcohol series and embedded in Epon 812. Thin sections were post-stained with uranyl acetate (saturated solution in 50% EtOH) and lead citrate (Reynolds, 1963) and examined and micrographed with a Philips 201 transmission electron microscope (TEM) operated at 60 Kv. Semithin (0.5 nm) sections were stained with toluidine blue for light microscopy.
Entero-endocrine cells in both genera were identified solely on an ultrastructural basis using Grube and Forssmann (1979) and Solcia et al. (1981) as guides. The ultrastructure of the acid-producing cells in our specimens was analyzed comparatively by using avail- able experimental data as a guide to recognition of secretory state (Ito and Schofield, 1978; Schofield et al., 1979). Our descriptions thus are based on cells judged to be in the same state of activity in both species.
Results
At the light microscopic (LM) level the fundic mucosa of Trachops and Chiroderma differed greatly (Fig. 1). In Trachops both surface mucous cells and chief cells were conspicuous because they contain toluidine blue stained secretory granules. In Chiroderma gastric pits either were very shallow or lacking and the basal half of each gastric gland was composed mostly of chief cells containing unstained granules. Transmission electron microscopic analysis revealed details of the var- ious cell types, as described in the following paragraphs. At both the TEM and LM levels of comparison no noteworthy individual variation, other than the usual localized differences in fixation, was found among specimens of the same species.
Entero-endocrine (endocrine-paracrine) cells.— In Trachops cirrho- sus only two types of entero-endocrine cells were distinguished. The most common was identified as an A-cell, present in nearly all gastric glands, positioned among the chief and parietal cells in the lower one- half of the gland (Fig. 1). These cells contained abundant spherical, electron-dense secretory granules (averaging 285 nm in diameter) with a narrow “halo” caused by an apparent space between the electron- dense material and the granule membrane (Fig. 2a). The cytoplasm contained scattered lamellar granular endoplasmic reticulum (GER), relatively few mitochondrial profiles, and lipid-like droplets (Fig. 2a). The A-cells frequently were juxtaposed to chief cells. The second cell type, identified as a D-cell, also was found within the most basal portion of the gastric gland, apparently often juxtaposed to A-cells. The D-cells were characterized by spherical secretory product (330 nm in diameter) with a finely granular appearance, exiguous GER, and few mitochon- drial profiles (Fig. 2b)
In Chiroderma, four distinctive types of entero-endocrine cells were
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Fig. L— Diagrams of the gross anatomy of the stomachs and histology of the mucosa in Trachops cirrhosus and a representative of Chiroderma (C. villosum). Tissue samples used for electron microscopy were taken from areas enclosed by circles in the stomach diagrams. The mucosa diagrams illustrate the histology as well as the relative height of the glands and relative numbers of particular cell types. Examples of the ultrastructure of particular cell types can be found by using the “F” labels as a guide to illustrations. Abbreviations are: py, pylorus; duo, duodenum; eso, esophagus.
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found in the lowermost portion of the gastric gland amongst the chief cells. One type contained highly irregular, somewhat granular secretory product (Fig. 3b) and corresponded morphologically to an EQ cell. These cells tended to have a pale, leached appearance with abundant tubular smooth endoplasmic reticulum (SER) and little or no GER. The second cell type most closely resembled the Di(H) cell (Grube and Forssmann, 1979); its cytoplasm contained small, spherical, electron- dense granules (approximately 1 60 nm in diameter), strands of GER, lipid-like droplets, and prominent Golgi complexes. The third type, which was by far the least common, corresponded to an A-cell in having abundant spherical electron-dense granules (Fig, 3a). The fourth type, which was common (possibly associated with each gastric gland), con- tained a mixture of granules most of which were small and contained a coarsely granular substance equivalent to that characteristic of gastrin- secreting G-cells (Fig. 4a, b). These cells were distinctive in that the cell body had elongate processes that extended among the chief cells so that cross-sectioned profiles were seen commonly (Fig. 4b). These elongate processes and the cell body itself were generally wedged in among the chief cells to the extent that chief cell secretory granules often intruded into the cytoplasm of the G-cell (Fig. 4a, b). This image was common and probably not an artifact because the granule mem- brane and cell membranes typically were intact (Fig. 4b). In the pro-
Fig. 2.— a: An A-cell positioned at the base of the fundic gland in Trachops. Note the spherical, electron-dense product (arrow) and lipid droplets (L) typical of these cells. T. cirrhosus, CM 63688. b: An example of a D-cell positioned among chief cells in Trachops. Note the typical secretory product (arrow) of these cells, which are thought to produce somatostatin. T. cirrhosus, CM 63688.
Fig. 3.— a: An example of an A-cell in Chiroderma. The mature secretory product (arrow) can be compared to an immature, electron-dense granule (img) associated with the Golgi complex. C villosum, CM 77601. b: An ECn-cell in Chiroderma; the granules (arrows) typically are elongate rather than spherical and possibly contain serotonin (5-HT). The cytoplasm of these cells often is pale and contains abundant tubular smooth endoplasmic reticulum (SER). A chief cell secretory granule (SG) also can be seen on the left. C villosum, CM 76798.
Fig. 4.— a: A possible G-cell at the base of a fundic gland in Chiroderma. Note how the body of the cell is wedged among adjacent chief cells, whose secretory product (SG) intrudes into the G-cell cytoplasm. The cell and granule membranes (arrow) are intact, suggesting that this close relationship is not an artifact, bl, basal lamina. C. villosum, CM 76798. b: A cross-section through a G-cell process that extends between chief cells filled with product (SG). An arrow denotes an image suggestive of exocytosis; also note the tubular smooth endoplasmic reticulum (SER) and intact chief cell and granule mem- brane (MB). C villosum, CM 76798.
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cesses we found tubular smooth endoplasmic reticulum and images suggestive of exocytosis, suggesting that the G-cell granules might be released here (Fig. 4b).
Chief cells. — The gastric chief cells in Trachops were clustered at the base of the gastric gland (Fig. 1 ). The cells were pyramidal and contained spherical, electron-dense secretory product; the nucleus was irregular or ovoid and basally positioned; the GER profiles usually were swollen and spherical in appearance and the cistemae were filled with pale granular material (Fig. 5 a). The Golgi complex was small and incon- spicuous.
The chief cells in Chiroderma were extremely abundant, occupying at least half of each gland (Fig. 1). These cells differed dramatically from those in Trachops in that the abundant product did not stain with toluidine blue. Additionally, with the TEM the product was swollen, pale granules. Although many of the granules were in the form of individual spheres, others often had a coalesced appearance (Fig. 5b). The nucleus was basally positioned and usually irregular; lamellar GER and large Golgi complexes filled most of the remaining cytoplasm.
Parietal (oxyntic) cells. — The cytoplasm of parietal cells in Trachops was characterized by a modest number of profiles of intracellular can- aliculi. The microvilli of these canaliculi were closely appressed and the intercellular spaces between them were filled with electron-dense product matching that found in the gland lumen (Fig. 6a, b). The cytoplasm of most parietal cells also contained large numbers of mi- tochondrial profiles and tubular and spherical vesicles (Fig. 6b). The parietal cells in our specimens of Trachops nearly always corresponded to an intermediate activity state (following Ito and Schofield, 1978).
In Chiroderma, the parietal cells could be placed into two categories: 1) an intermediate secretory stage, which was most common; and 2) an actively secreting stage, which was far less common (Fig. 7a, b). The ultrastructure of parietal cells in the intermediate stage differed from those in Trachops in the same stage in that these cells invariably had extensive, swollen-appearing intracellular canaliculi with thick, elongate microvilli (Fig. 7b). The cytoplasm of all parietal cells had abundant mitochondrial profiles and numerous vesicles, scattered GER profiles, and concentrations of lipofuscin granules (Fig. 7a, b). In active parietal cells, the intracellular canaliculi were greatly expanded and the
Fig. 5.— a; Chief cell in Trachops; note the spherical electron-dense product (SG) and example of exocytosis (arrow) into the lumen. NU, nucleus. T. cirrhosus, CM 63688. b: Chief cells in Chiroderma; note the pale product (SG) and example of coalescing among secretory granules (CO-SG). NU, nucleus. C. villosum, CM 76798.
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microvilli lining their borders were less compact (Fig. 7a). The cyto- plasm of these cells contained few vesicles; instead, the Golgi complexes were the most prominent feature.
Mucous cells.— In Trachops, the mucous neck cells were scattered among the chief and parietal cells in the lower one-half of the gastric gland. The apex of each mucous neck cell bordered on the gland lumen. The nuclei were basally positioned and the apical cytoplasm contained secretory granules with pale, finely granular material and a distinctive electron-dense droplet occupying about one-fourth of the granule diameter (Fig. 8a).
The surface mucous cells and cells lining the gastric pits differed considerably from the mucous neck cells. The secretory product in these surface cells was more clearly defined (coalescing granules were not observed) and consisted of an electron-dense droplet set against a reticulated, dense substrate (Fig. 9). Some granule polymorphism was apparent; some of these cells also contained spherical or irregular, completely electron-dense granules located in the vicinity of the Golgi complex (Fig. 9).
The mucous neck cells in Chiroderma were relatively small and sparsely distributed and they primarily were positioned among parietal cells (Fig. 8b). The secretory product differed considerably from that in homologous cells in Trachops. The granules in Chiroderma consisted of a mixture of pale, flocculent material along with a coarser, fibrillar material and a small, peripheral electron-dense component (Fig. 8b). Gastric pits were nearly lacking in Chiroderma. The gastric surface consisted of mucous cells and exfoliating parietal cells with dark, dense
Fig. 6, —a: Parietal cell in Trachops; note the typical image of intracellular canaliculi in these cells in this species (arrow). NU, nucleus; Mi, mitochondria; M, mucous neck cell. T. cirrhosus, CM 63688. b: Parietal cell in Trachops; note the intracellular canaliculi (arrow) and tubular and spherical membrane profiles (V) in the cytoplasm. Abbreviations
as above. T. cirrhosus, CM 63688.
Fig. 7.— a: Actively secreting parietal cell in Chiroderma; note the expanded intracellular canaliculus and cluster of lipofuscin granules (arrows). NU, nucleus; Mi, mitochondria; L, lumen. C. villosum, CM 76796. b: Inactive parietal cell in Chiroderma; note the lipofuscin granules (arrows) and compare the intracellular canaliculi (C) to those seen in Trachops parietal cells (Fig. 6). Abbreviations as above. C. villosum, CM 76796.
Fig. 8. —a: Mucous neck cells in Trachops. An immature granule (IMG) in association with the Golgi complex can be compared to mature mucous granules (MSG), which have an electron-dense droplet (arrow). T. cirrhosus, CM 63688. b: Mucous neck cell in Chiroderma. This cell is positioned near parietal cells (P) and is packed with mucous granules (MSG), which have an electron-dense droplet (arrows) but otherwise differ from those seen in Trachops. C. villosum, CM 76798.
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cytoplasm. The surface mucous cells differed considerably from those in Trachops because the secretory product was pale and contained fibrillar material as well as a small, denser sphere at the margin of each granule (Fig. 10). Unlike the situation in Trachops, in Chiroderma the surface mucous cells and mucous neck cells appeared to produce only a single type of granule (Figs. 8b, 10).
Discussion
Comparative transmission electron microscopy has been shown to be a useful addition to histological and histochemical techniques in the study of systematics and evolutionary biology of mammals (for ex- ample, Phillips et al., 1984; Feldman and Phillips, 1984; Tandler et al., 1986; Phillips et al., in press; Phillips and Tandler, in press). An analysis by TEM allows for interspecific comparisons of structure among cells that are thought to be homologous (following criteria of Rieger and Tyler, 1979) and in the same stage of differentiation. In making our comparisons of ultrastructure we recognized the fact that, in effect, we were comparing cells “fixed” at a moment in time and, thus, to some extent the molecular events and organization that we were viewing and comparing were aspects of functional state rather than genic differences that directly determine cell structure (Phillips et al., 1984). On the other hand, in making comparisons between regu- lated, polarized secretory cells in the two genera of bats, we were able to compare directly the ultrastructure of mature secretory granules containing gene products such as mucus, pepsinogen, and a variety of peptide hormones (Phillips and Tandler, in press). In our present in- vestigation we found noteworthy ultrastructural differences among all cell types and many of their products as well as in the relative numbers of particular cell types in the fundic glands of two ecologically divergent species of bats (Table 1).
Entero-endocrine cells are important because they synthesize and secrete a wide variety of peptides as well as 5-hydroxytryptamine (sero- tonin) and these products can have a complex, and as yet not fully understood, controlling or modulating influence on the digestive tract (Pearse, 1969; Grube and Forssmann, 1979; Solciaetal., 1981). Indeed, although we use the terms “entero-endocrine” or “endocrine,” at least some of these cells actually might be regarded as “endocrine-paracrine” cells. Our identification of entero-endocrine cells was based solely on
Fig. 9. ““Surface mucous cells that line the surface of the stomach and the gastric pits in Trachops. Note the variety of product images in the cytoplasm (an ows). Those nearest to Golgi complexes (G) often are the most electron dense. NU, nucleus. T. cirrhosus, CM 63688.
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Fig. 10.— Surface mucous cells in Chiroderma. These cells are packed with secretory granules (MSG) that have an electron-dense droplet (arrow) and resemble those in mucous neck cells (Fig. 8b). Note the sparse microvilli (MV) on the apical surface of these mucous cells. C. villosum, CM 76798.
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Table L—A comparative summary of features of the fundic glands in Trachops cirrhosus and Chiroderma trinitatum and C. villosum.
|
Feature |
Trachops |
Chiroderma |
Comments |
|
Entero-endo- |
A, glucagon |
A, glucagon |
See Figs. 2, 3, 4 |
|
crine (=en- |
D, somatostatin |
EC„, serotonin |
|
|
docrine™ |
D„? |
||
|
paracrine) cells |
G, gastrin |
||
|
Chief cells |
Moderate num- |
Extremely abundant |
Ultrastructural differ- |
|
bers |
ences in secretory product; Figs. 1, 5 |
||
|
Parietal cells |
Small in size, |
Extremely abundant. |
Ultrastructural differ- |
|
low average |
large in size; very |
ences allow for |
|
|
activity level |
high average ac- |
comparisons of ac- |
|
|
in fasted ani- |
tivity level in fast- |
tivity level; Figs. |
|
|
mal |
ed animal |
6, 7 |
|
|
Mucous neck |
Common |
Scarce |
Ultrastructural differ- |
|
cells |
ences in secretory product; Fig. 8 |
||
|
Gastric pit |
Abundant, with |
Very sparse, gastric |
Ultrastructural differ- |
|
and surface |
deep gastric |
pits nearly non-ex- |
ences in secretory |
|
mucous cells |
pits |
istent |
product; Figs. 1, 9, 10 |
ultrastmcture; thus, we could only infer products from experimental literature. The following discussion of entero-endocrine cell products is therefore largely reliant on the accuracy of ultrastructural identifi- cation of particular cells; immunohistochemistry will be necessary for more precise identification.
In Trachops we identified A-cells (Fig. 2), which are thought to secrete glucagon (Moody et al., 1978; Unger et al., 1978) and D-cells, which secrete somatostatin (Hokfelt et al., 1975; Grube and Forssmann, 1979). Cells with essentially the same ultrastructure have been found in other species; A-cells have been described in Pteronotus, Phyllostomus, Car- ollia, and Artibeus, whereas D-cells have been found previously only in Pteronotus and Phyllostomus (Phillips et al., 1984). Yamada et al. (1984) have demonstrated both glucagon- and somatostatin-like im- munoreactivity in endocrine cells in the fundic region of the stomach of the common vampire bat, Desmodus rotundus, so although ultra- structural data are lacking, this species also has both A- and D-cells. Chiroderma differed from Trachops in that A-cells were rare and in having ECn-cells, G-cells, and Di(H) cells, none of which was found in the Trachops material examined by us. However, it must be remem- bered that cell types that are scarce, or common but localized, could easily be overlooked in a TEM survey. Light microscopy of semithin
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(0.5 fjLm) sections does not solve this problem because many endocrine cells cannot be detected and identified with light microscopy and rou- tine staining.
The apparent presence of abundant glucagon- and somatostatin-pro- ducing entero-endocrine cells in Trachops is significant because these cells 1) also are found in other insectivorous, animalivorous, and san- givorous species (Phillips et aL, 1984; Yamada et aL, 1984), 2) are abundant, 3) are positioned among chief and parietal cells, and 4) because each of these products is known to inhibit gastric acid secretion (Konturek et aL, 1975; Kusomoto et aL, 1979; Stachura et aL, 1981). Additionally, the physical proximity of A- and D-cells in Trachops resembles the situation in dogs, in which somatostatin apparently also inhibits, or limits, glucagon production (Kusomoto et aL, 1979). In any case, the abundant presence of A- and D-cells in Trachops might cor- relate with the moderate number of generally inactive parietal cells, just as we found previously in both Pteronotus and Phyliostomus (Phil- lips et aL, 1984). Likewise, the relative rarity of glucagon-producing cells and the possible absence or scarcity of somatostatin-producing cells in Chiroderma correlates with the extreme activity of the large and abundant parietal cells (Figs. 1 , 7).
The presence of and Di-cells in Chiroderma (Fig. 3) cannot readily be related to any particular feature of the fundic gland. The EQ-cells are thought to have endogenous serotonin ( 5 -hydroxy tryp- tamine), which also is found in enteric neurons (Grube and Forssmann, 1979; Gershon, 1981). The physiological roles of entero-endocrine cell serotonin possibly include inhibition of acid release along with stim- ulation of mucus. Some investigators also regard serotonin as an in- trinsic neurotransmitter that modulates smooth muscle and affects blood flow (Gershon, 1981; Ormsbee and Fondacaro, 1985), Examples of an EQ-type of cell also have been found in Phyliostomus hastatus so although we failed to find examples of EQ-cells in our specimens of Trachops, their occurrence in Chiroderma suggests that they might be widespread in phyllostomid bats. This is further supported by Yamada et aL (1984) who reported that they found moderate numbers of sero- tonin-immunoreactive cells in the fundus of vampire bats.
The Di-cells in Chiroderma have an ultrastructure that is indistin- guishable from that found in D, -cells in Artibeus (Phillips et aL, 1984). Some authors have linked D, -cells to production of vasoactive intes- tinal polypeptide (VIP) in the mammalian gut (for example, Grube and Forssmann, 1979), but others (for example, Larsson et aL, 1979) have argued that the peptide produced by D, cells is not VIP but a similar molecule. Some workers now think that VIP in the digestive tract is found only in nerve fibers and nerve cell bodies (Baecker et aL, 1983). Another possibility is that the D, cells in Chiroderma produce motilin
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or bombesin, or both, because these peptides also have been associated with cells that ultrastructurally fall into the Di cell group (Capella et aL, 1978; Solcia et al., 1981). The product and physiological role of Di cells in Chiroderma and Artibeus thus is unknown at the present time.
The possible G-cells found by us at the base of fundic glands in Chiroderma are noteworthy for several reasons. Heretofore, gastrin- producing cells have not been identified in the fundic region of the mammalian stomach (Solcia et al., 1981) and Yamada et al. (1984) did not find any gastrin-like immunoreactivity in the fundus of vampire bats. Secondly, the gastrin peptide is known to stimulate gastric acid secretion (for example. Chew and Hersey, 1982), which is considerable in Chiroderma, if abundance, large size, and ultrastructure of parietal cells are reliable indicators of the level of acid secretion in the absence of quantitative data on gastric acid. Thirdly, the ultrastructure of the G-cells and their cytoplasmic granules is consistent with published descriptions of known G-cells in other mammals (Grube and Forss- mann, 1979; Solcia et al., 1981) and with G-cells in the pylorus of Ariteus flavescens, which were identified by demonstration of gastrin- like immunoreactivity (Mennone et al., 1986). The apparent physical relationship between G-cells and chief cells in Chiroderma is another matter. It is not known with certainty that gastrin stimulates secretion of pepsinogen; instead, chief cells in mammals have been shown to have receptors to the peptides secretin and cholecystokinin (CCK) (Her- sey et al., 1983, 1984). However, the ultrastructure of possible G-cells in Chiroderma differs substantially from the ultrastructure of either secretin or CCK-producing endocrine cells (Solcia et al., 1981) and thus would not be easily confused with either of these. Gastrin and CCK have been shown to share a common C-terminal portion of the mol- ecule (Larsson and Rehfeld, 1977; Solcia et al., 1981) but in mammals they are produced by separate cells and appear to have separate func- tions. Indeed, it seems to be typical of entero-endocrine cells that their various products are stored in membrane-bound granules that have specific and consistent ultrastructural morphology and gastrin and CCK, regardless of their molecular similarities, are found in very different- looking granules (Phillips and Tandler, in press).
Available data on various types of entero-endocrine cells in Tra- chops, Chiroderma, Phyllostomus, Carollia, Glossophaga, Artibeus, Ar- iteus, Erophylla, Desmodus, and Pteronotus (Phillips et al., 1984; Ya- mada et al., 1984; Mennone et al., 1986) raise far more questions than answers. Some peptides (for example, glucagon, somatostatin, and gas- trin) have been investigated to the extent that we can begin to relate physiological data from laboratory studies of other species to our find- ings in bats. However, most other peptides possibly produced in species-
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Specific patterns of differential abundance in bats are far more difficult to integrate with other data because less is known about possible func- tions. The problem is compounded by the fact that some of these molecules are found in both enteric nerves and endocrine cells and may be functionally different in each (Hokfelt et al., 1975; Grossman, 1976; Schultzberg et af, 1980; Gershon, 1981; Miller, 1984). Never- theless, these regulatory molecules probably have played a key role in the evolution of dietary diversity as seen in bats, and differential pro- duction of peptides or differential location of cells that produce some peptides might even contribute to or account for interspecific differ- ences in histology of the chiropteran digestive tract (Mennone et aL, 1986). Various peptides, and 5-HT as well, ultimately might provide a key to our understanding of the evolution of histological and func- tional diversity.
Parietal cell ultrastructure in Chiroderma closely resembles that of Artibeus and Ametrida (Phillips et al., 1984; Phillips, unpublished data) and, judging from experimental studies, is indicative of a high rate of HCl secretion, even in fasted animals (Ito and Schofield, 1978; Scho- field et al., 1979; Black et al., 1980). This high rate of secretory activity may be further substantiated by the abundance of lipofuscin granules (Fig, 7) in Chiroderma parietal cells observed in all of our specimens. This type of lysosome has been correlated with degradation of by- products of cellular metabolism and its presence in parietal cells sug- gests that cellular metabolism is occurring at a high rate (Toth, 1968; Fawcett, 1981).
Chief cells (which produce pepsinogen and rely on gastric acid for conversion to pepsin) also are extremely active in phyllostomid fruit bats, judging from the unusual ultrastructural appearance of the product (Fig. 5) in Chiroderma, Artibeus, and Ametrida (Phillips and Stud- holme, 1982; Phillips et al., 1984). This pattern may be a widespread occurrence in fruit bats because in Eidolon helvum, an African mega- chiropteran, an analysis of digestive enzymes demonstrated high pepsin content in both the stomach and lower esophagus (Ogunbiyi and Okon, 1976) and histological study has revealed abundant chief and parietal cells (Okon, 1977).
Differences among the mucus-producing cells in Trachops and Chi- roderma were significant, but in keeping with previously reported pat- terns (Forman, 1972; Phillips et al., 1984). Trachops produces abun- dant mucus, whereas Chiroderma produces little mucus due to the limited number of mucous neck cells, the virtual absence of gastric pits lined with mucous cells, and the relative scarcity of typical mucous surface epithelial cells. In Trachops, the mucous cells ultrastructurally resembled those found in Pteronotus and Phyllostomus (particularly in
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the latter species) and differed from those in Carollia and Artibeus (Phillips et al., 1984) and Chiroderma. However, Trachops differed from its nearest studied relative {Phyllostomus, cf. Smith, 1976; Hood and Smith, 1 982) in having relatively shallower gastric pits (hence fewer surface mucous cells) and in having a chemically different mucus in the surface cells. In Phyllostomus, the surface mucous cells exhibit toluidine blue metachromasia (Forman, 1972), suggesting the presence of a sulfonated compound (Spicer, 1963), whereas in Trachops the product is negative to toluidine blue staining. The significance of this chemical difference is unknown. It might represent a real interspecific difference but also could be the consequence of physiological state of the animals at time of sacrifice because Ohara et al. (1984) have dem- onstrated that histochemical changes in mucous glycoproteins can oc- cur within hours in fasted laboratory rats. Although our bats all were handled the same way after capture, we have no way of knowing for certain whether they fed before being collected. On the other hand, intraspecific consistency among our specimens does seem indicative of a real interspecific difference.
The greatly reduced mucus-production in Chiroderma seems to be characteristic of Neotropical fruit bats (Forman, 1972, 1973). Although mucus often has been regarded as a major factor in the protection of the gastric epithelium, its relative scarcity in fruit bats with highly active parietal cells suggests otherwise. A recent study (Robert et al., 1984) that demonstrated a lack of correlation between the thickness of the mucus coat and protection of the stomach lining in laboratory rats helps to explain the fruit bat data. But how is the stomach protected in bats such as Chiroderma and Artibeusl One possibility is that the lining of the stomach is protected by salivary gland secretions (Studier et aL, 1983Z?; Phillips et al., 1984). Other possibilities include surface- active phospholipids (Lichtenberger et al., 1983) and H+ disposal by the surface c6lls through a Na+/H+ exchange system (Olender et al., 1984).
In summary, it seems reasonably clear that mucous production (amount and chemistry) in the fundus of the stomach in stenodermatine fruit bats differs substantially from that in phyllostomine animalivo- rous species. The mucus produced in fruit bats is less complex in the sense that mucous neck and surface cells are ultrastructurally the same, whereas in animalivorous species such as Trachops notable differences can be found when one compares the secretory granules in these cells. The biological significance of this intrafamilial divergence in mucous cells and in ultrastructure of mature product is unknown but the con- sistency of the pattern in the genera examined to date (Forman, 1972; Phillips et al., 1984) is noteworthy because when differences in the
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ultrastructure of an exocrine cell product can be correlated with sys- tematics, a foundation is set for studying evolution at the cellular level (Tandler et aL, 1986; Phillips and Tandler, in press).
To what extent do our ultrastructural data provide answers for our original question about the relationship among dietary and structural specialization and cellular features of the gastric mucosa? Although Trachops cirrhosus clearly represents an instance of behavioral evo- lution that includes specialized feeding on Neotropical frogs (Tuttle and Ryan, 1981), the gastric fundic mucosa at the cellular level is similar to that of Phyllostomus hastatus and P. elongatus (Phillips et aL, 1 984), which generally are regarded as “animalivorous” or “insectivorous” (Gardner, 1977). The only differences of note are the possible absence of ECn-cells, the slightly shallower gastric pits, and the absence of toluidine blue positive components in the mucus in Trachops. The fundic mucosa of Trachops thus seems to be somewhat generalized even though these bats have adopted a specialized behavior and feeding strategy.
Chiroderma is very similar to Artibeus at the cellular level (Phillips et aL, 1984). It differs significantly, however, in having a relatively greater abundance of chief cells (45“-75% of each fundic gastric gland as compared to about 25%). Cellular differences in the gastric mucosae of different Neotropical fruit bats are interesting because they suggest the possibility of subtle differences in diet or in the assimilation of nutrients from a shared diet.
Lastly, how do Chiroderma and other previously studied phyllos- tomid fruit bats compare to the megachiropteran fruit bats? This ques- tion is significant because evolutionary convergence is virtually un- studied at the histological, histochemical, and ultrastructural levels. Available data support the theory that frugivory evolved independently in these bats (Kamiya and Pirlot, 1975; Smith, 1976) and some phys- iological studies suggest that the two groups might be different in the ways that they actually regulate their diets or dietary intake (Thomas, 1984). Furthermore, some data can be interpreted to show that mega- chiropteran and microchiropteran bats might have had separate origins altogether (Smith and Madkour, 1980). Given this, the general histo- logical similarities in gastric mucosa of microchiropteran fruit bats of the genera Chiroderma, Artibeus, and Ametrida on the one hand (pres- ent study; Phillips and Studholme, 1982; Phillips et aL, 1984) and the African megachiropteran. Eidolon helvum, on the other, are indeed remarkable. Judging from the published data of Ogunbiyi and Okon (1976) and Okon (1977), the microchiropteran fruit bats and E. helvum have the following features in common: 1) very shallow, almost non- existent, gastric pits; 2) a scarcity of mucous neck cells; and 3) abundant.
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extremely active parietal and chief cells. The extent of histological convergence between the stenodermatines and megachiropterans as a group is less certain because the latter seem to be quite variable and published descriptions are not always adequate for comparisons. How- ever, the stenodermatines do appear to share a variety of histological features with Rousettus, Pteropus, Eonycteris, and Penthetor (Kamiya and Pirlot, 1975; Bhide, 1980) and future ultrastructural comparisons will be of interest.
Acknowledgments
Our research and field work were supported by the Research Corporation (grants C-1251 and C-1855 to Phillips), Hofstra University (HCLAS Executive Committee grants to Phillips), the Alcoa Foundation, and the M. Graham Netting Research Fund (to Dr. H. H. Genoways, Carnegie Museum of Natural History). We are particularly indebted to Hugh H. Genoways for his support and assistance to our research. Stephen L. Williams, Jane Groen, Nadine M. Sposito, Rodney L. Honeycutt, Ben Koop, and Mike Arnold all assisted with the field work. In Suriname, Drs. J. P. Schultz and H. A, Reichart of STINASU were especially helpful and they are gratefully recognized. Lastly, we greatly appreciate the work of our patient typist, Linda Cossen, of the Hofstra Uni- versity Special Secretarial Services.
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Chew, C. S., and S. J. Hersey. 1982. Gastrin stimulation of isolated gastric glands. Amer. J. Physiol., 5:G504-G512.
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Back issues of many Annals of Carnegie Museum articles are available, and a few early complete volumes and parts are listed at half price. Orders and inquiries should be addressed to: Publications Secretary, Carnegie Museum, 4400 Forbes Avenue, Pittsburgh, Pa. 15213.
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VOLUME 55 15 SEPTEMBER 1986 ARTICLE 9
AN INTERNATIONAL SURVEY OF THE POPULAR AND TECHNICAL LITERATURE OF MAMMALOGY
Michael A. Mares ^
Research Associate, Section of Mammals
Janet K. Braun*
Abstract
A questionnaire designed to assess the diversity and abundance of the technical, semi- technical, and popular literature of mammalogy was sent to curators in 1 6 1 mammal collections and museums in 74 countries; 104 responses were received from 55 nations. Results show a great disparity in the availability of the different types of literature among countries. Some nations have a very strong literature at all levels, while many are in great need of all types of literature. Countries that have a rich literature at the semi- technical and popular levels also have a strong foundation in the basic technical literature of mammalogy. A statistical analysis of various socioeconomic factors shows that per capita income (PCI) is strongly positively related to the availability of literature at all levels: increasing the PCI by $ 1 000 per year results in a doubling of the available literature on mammalogy. The analysis suggests that only through international cooperative re- search on basic mammalogy can the PCI block be bypassed and the stage set for an increase in the popular literature of mammalogy.
An overview of the mammalogical literature is presented for each country queried. Responses of the curators are also given. Almost all respondents indicated a willingness to work in a cooperative manner with foreign scientists in producing semitechnical and popular literature in mammalogy.
' Address: Stovall Museum and Department of Zoology, University of Oklahoma, Nor- man, Oklahoma 73019.
Submitted 26 July 1984.
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Introduction
Mammalogy is a diverse discipline, encompassing such areas as sys- tematics, ecology, behavior, physiology, anatomy, and other fields of specialization. The trend in recent years has been away from basic systematic and survey research and toward more scientifically elegant and popular experimental and laboratory-oriented work. We believe that this is a very healthy and exciting pattern, particularly in the developed nations of the world. As Mares (1982, 1985, in press) has pointed out, one very important reason that modem theoretical/em- pirical research on mammals can be performed in developed countries is because the foundational research on mammal species was done decades ago by scores of mammalogists who constructed a framework upon which current research can be supported. It is no accident that most of the important research in state-of-the-art mammalogy is being performed by investigators living in countries that have a long and noble history of basic faunal research. In many cases, the organisms being examined by the current crop of new mammalogists have been studied in one form or another for more than a century.
An unfortunate correlate of the trend toward experimental research is the tendency by some investigators to view foundational systematic and survey research as being somehow less scientific and less inherently valuable than investigations that are dernier cri. This view obtains because in developed countries there is little work left to be done at the foundational level. Moreover, younger biologists, trained in the United States for example, are often quite parochial in their views of nature. This is not said in a condemnatory fashion, but is merely observational. Most young mammalogists in developed countries have only had dealings with very well-studied aspects of their natural en- vironment. They have seldom been at a real loss for either taxonomic or ecological information on any vertebrate species with which they are familiar, and their view of nature reflects this familiarity. They tend to think of nature as a well-studied entity, believing that most of the foundational work was completed early in this century and viewing the research problems that remain to be done as exciting tests of the- oretical questions. We believe that these views are consistent with an educational background in a country that is rich in the foundational literature of mammalogy. We suggest that these sanguine views of basic field biology are in error.
The literature of mammalogy is extremely diverse. Almost all coun- tries of the world support one or more mammal collections or natural history museums that often help to contribute to the basic literature of the science of mammalogy (for example, Genoways and Schlitter, 1981; Hickman, 1981). Hickman (1981) surveyed the field guides that
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had been published to the mammal faunas of most countries of the world. However, in many cases the guides or papers cited by Hickman are quite old (that is, published in the mid-1800’s or early 1900’s), and/or are quite difficult to obtain. More important, as Mares (1985, in press) has argued, field guides and other popular literature owe their existence to the foundational literature of mammalogy, the taxonomic and geographic surveys that first clarify the faunal makeup of a country. It was clear from Hickman (1981), and from our own work in various countries, that there are pronounced differences in the availability of the mammal literature of the world. Some countries seem to have a particularly abundant literature, whereas others have, at best, a very scant literature on mammals. Moreover, we had not seen any reports examining the availability of both the popular literature on mammals and the technical literature that provides its foundational material. If, indeed, these two types of literature bear some relationship to each other, we might expect that few countries lacking a strong basic liter- ature will show a pronounced level of development of popular literature on mammals.
Just exactly how extensive the foundational literature of mammalogy might be is not clear. No one, to our knowledge, has attempted a comprehensive assessment of the actual status of the world’s mam- malogical literature. We feel that information on the availability of such literature on a global scale will help point out potential areas requiring increased research efforts. Such a survey might also indicate that the world we inhabit is less biologically explored than many sur- mise.
In this paper, we present results of a survey of the literature of mammalogy that were obtained through the use of a questionnaire that was sent to most of the mammal collections of the world. A preliminary report on these data was given in Mares (1985); however, we herein present a much more complete literature survey based on many more responses than were included in the earlier report. These data present an overview of the popular and technical literature of mammalogy for many countries of the world. They include data from most countries that have a mammal collection of greater than 50 specimens, and offer some information on the availability of mammal literature for almost all countries of the world that have a mammal collection of even modest size. Taken together, these data give some indication of the current status of the world’s mammal literature, including which countries have reached a high level of literature availability. They also show which countries need a cohesive plan of work to provide either the technical or popular literature, or both, that are required if the country’s literature is to be brought up to acceptable levels.
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Materials and Methods
A questionnaire designed to assess the availability of various types of natural history literature (primarily that dealing with mammals) in different countries of the world was sent out in 1981. Respondents were asked to list available publications and indicate if these publications were available in the common language of the country. The following questions were used: 1) Is there a publication (in the style of a field guide) available that deals with the identification of the mammals of your region? 2) Is there a publication (technical survey) available to the museum specialist which is concerned with the mam- mals of your region? 3) Are there taxonomic keys available to the mammals of your region? 4) Are there any works available that deal with the collecting or preparation techniques of mammals? 5) Are there any publications available that deal with the operation or importance of natural history museums? 6) Would you judge the number of available children’s books dealing with natural history topics to be many, few, or rare? 7) Please list three natural history or museum science subjects that you feel need attention in your particular geographic area. 8) Please list any projects underway that deal with any of the topics listed in the first seven questions. 9) Would you be interested in cooperative research with other museum specialists in preparing publications such as field guides, textbooks, etc., that deal with the fauna of your area? Names of museums, research organizations, collections, and personnel (curators and directors) were obtained from Genoways and Schlitter (1981). Museums or research organizations were queried if: (1) the collections were of a substantial size, or (2) they were the only major museum or collection in a particular country or region.
The countries queried are listed in alphabetical order and questionnaire responses are indicated where applicable. Countries which were not surveyed or which did not respond to the questionnaire are indicated by ‘Wo response received"" before the list of literature citations. Responses given by the respondents are italicized. Additional literature cita- tions, not given by respondents, were added to present a more complete overview of the literature availability for the countries listed and are not italicized. Topics were eliminated if the respondent(s) did not answer a question. Responses to research needed or projects underway which did not directly pertain to mammals or museum science were omitted. Under “Comments” we have either summarized as succinctly as possible, or reproduced verbatim, informal comments of respondents. In some cases we have added additional observations that we feel are particularly important to an understanding of the literature situation in any given country. Willingness of the respondent(s) to work in cooperation with specialists from other museums and institutions are denoted by WTC at the end of the comments if more than one-half of the respondents of a particular country answered positively. The absence of this designation indicates that a negative response was received or that this question was not answered. A concerted effort was made to verify all references given by the respondents. Those references which could not be verified and/or are partial references are denoted by an asterisk following the reference number.
Hickman’s (1981) broad overview of field guides to national mammal faunas includes many old (pre-1940) natural history publications. We have tried not to duplicate works cited in Hickman (1981). Moreover, we have attempted to list only the more readily available recent literature.
Countries responding to the questionnaire were placed into one of six groups based upon the major geographic regions: Latin American, European, Middle Eastern, African, Asian, and Australian. Summaries of responses to each question are presented separately in tabular form (Tables 1-3).
Data on gross national product (GNP), per capita income estimates, population growth rates, population density, literacy, and the percent of the population living in urban areas were obtained from The Global 2000 Report to the President (Barney, 1982) and The World Almanac (Lane, 1983).
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Results
Survey of Responses
Afghanistan: No response received; but see 26 347, 409, 429, 430.
Algeria: No response received; but see 344, 401.
Angola: No response received; but see 279, 401.
Argentina: Museo Municipal de Ciencias Naturales “Lorenzo Scag- lia.” Guides: None; but see 447. Surveys: 496; see also 113, 114, 115, 189, 273, 372, 387, 391, 475. Keys: None; but see 272, Technical manuals: 130*; see also 102. Museum science literature: None. Chil- dren’s literature: Rare. Research needed: Paleontology. Projects un- derway: None; also, Mares et aL, Mammals of Tucuman, Los Mami- feros de Salta. Comments: The recent field guide by Olrog and Lucero is already out of print, and was difficult to obtain even when it was still in print. Greenhall has written a key to the bats of Argentina and Pearson has written a key to the small mammals of Nahuel Huapi and Lanin National Parks. Both of these are published or in press. Most of the surveys published in Spanish are difficult to obtain, while those published in English are either poorly disseminated among the scien- tists in Argentina or are themselves difficult to obtain. Argentina il- lustrates a problem that will be encountered time and again during this survey, and that is the difficulty in obtaining publications that are published in obscure journals or materials that are unpublished and circulated among a few specialists. The literature of mammalogy in Argentina is quite poor, particularly for native students of the Argentine fauna.
Australia: Arthur Rylah Institute for Environmental Research; Australian National Wildlife Collection; Central Australian Museum; Monash University; National Museum of Victoria; Queensland Mu- seum; Queen Victoria Museum and Art Gallery; South Australian Mu- seum; Taronga Zoo; Tasmanian Museum and Art Gallery; The Aus- tralian Museum. Guides: 29*, 218, 234, 300, 312, 313, 314, 389, 500, 663; see also 85, 662. Surveys: 13, 40, 83, 84*, 234, 300, 389, 459, 500, 662, 663; see also 41, 188, 215, 219, 238, 375, 576, 654, 676. Keys: 12, 39, 144, 300, 306, 337, 663. TecMical manuals: There is no publication which is specifically oriented in this direction. The Tas- manian Museum and Art Gallery has a leaflet on collecting animals for the museum. Works published in North America and Europe are used. Museum science literature: 483* gives a historical account of the beginnings of the National Museum of Victoria but does not cover the last 25 years. Several journals are used including: Kalori, of the Mu- seums Association of Australia, Museum, Museum News, Curator, and Museum Journal; see also 533. Children’s literature: Few to many.
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Research needed: Studies on marine mammals; ecology, ethological and conservation studies on native vertebrates; animal population dy- namics; handbooks and field keys; collection and preparation technique manuals; publications dealing with ecological and behavioral tech- niques for naturalists; museum science publications; field guide to ce- taceans recorded in Australian waters; detailed surveys of the mammals of particular geographic regions. Projects underway: The zoology de- partment of the University of Tasmania is publishing a Fauna of Tas- mania Series; two field guides to Australian mammals are in prepa- ration, one by J. W. Calaby and one by M. Archer; B. Marlow is preparing an identification manual (at a technical level) to all the ter- restrial mammals of Australia. Comments: Several respondents indi- cated that a large number of works are currently in preparation. Many indicated that no comprehensive keys are available to Australian mam- mals. Many referred to preparation/collection techniques manuals of the British Museum (Natural History). No publications are available which deal specifically with museum science and literature; some are included as part of a treatment of other topics, however. WTC.
Austria: Naturhistorishes Museum Wien; Oberdsterreichisches Landesmuseum. Guides: JO*, 75*, 90. Surveys: 65*, 411, 432, 670. Keys: JO*, 75*, 432. Technical manuals: 471 and later editions, 161. Museum science literature: None. Children’s literature: Few. Research needed: A modern handbook on European mammals; a survey of Aus- trian mammals; a semipopular identification manual. Projects under- way: All of the above are in preparation. Comments: WTC.
Belgium: Institut Royal des Sciences Naturelles de Beligique; La- boratorium voor Algemende Dierkunde. Guides: 94. Surveys: 192, this publication is not up to date, and is without notes concerning distri- butions’, see also 285. Keys: 575* only for Rodentia, Lagomorpha, and Insectivora. Technical manuals: 285. Museum science literature: None. Children’s literature: Many, but no original work— most are transla- tions. Research needed: Mammal distributions; ecology of the Carni- vora; studies on mammal protection. Projects underway: Mammal dis- tributions. Comments: WTC.
Belize: No response received’, but see 231, 233, 273, 309, 336.
Bolivia: No response received’, but see 27, 28, 113, 114, 115, 273, 666, 695. Comments: Research is currently underway by Sydney An- derson and various collaborators on the mammals of Bolivia. Literature on the mammals of Bolivia is particularly depauperate.
Botswana: No response received’, but see 401, 557, 558.
Brazil: Museu de Ciencias Naturais; Museu Nacional; Museu Par- aense “Emho Goeldi”; Universidade Estadual Paulista. Guides: 65i. Surveys: 113, 114, 420, 641, 642’, see also 45, 46, 47, 115, 126, 273,
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388, 419, 466, 476, 550, 584, 640, 643. Keys: 420, 641, 642, 651; see also 472. Technical manuals: A collection and preparation manual written by the Departamento de Zoologia was published by the Mus. Zool, Univ. Sdo Paulo in 1967 (152). Museum science literature: None. Children’s literature: None. Research needed: Keys to the mammals of southeastern Brazil. Projects underway: Chiroptera of southeastern Brazil; Chiroptera of Mato Grosso, Brasil; chave para determinaqao de quiropteros brasileiros (reformulation and actualization); guia para identificaqdo de quiropteros do sul do Brasil (ndo existe verbas para impressdo) (F. Silva). Comments: Research is currently underway on mammals of the Cerrado by C. Alho and M. Mares. Brazil has an extremely complex fauna of mammals and has only been cursorily examined (474). One respondent, in referring to the availability of field guides for Brazil, noted that a field guide was published long ago but is not available today. This same respondent noted that many of the publications on Brazilian mammals are no longer available, even to specialists. WTC.
Burundi: No response received; but see 401, 639.
Cameroon: No response received; but see 164, 165, 401.
Central African Republic: No response received; but see 40 1 , 528.
Chile: Coleccion Particular de Fabian Jaksic y Jose Yafiez; Instituto de Ecologia y Evolucion; Laboratorio de Citogenetica; Museo de Zoo- logia de la Universidad de Concepcidn. Guides: 412, 454. Surveys: 383, 454; see also 113, 114, 115, 154, 220, 273, 475, 477, 588. Keys: 382, 497; see also 383, 454. Technical manuals: None, although 230 has been used in the past. Museum science literature: None; one re- spondent reported There is something published by Museo Nacional de Historia Natural, Casilla 787, Santiago, Chile. Another respondent suggested 1 54 as a source of information on museum science literature; see also 694. Children’s literature: None or rare. Research needed: Ecology; behavior; distribution and zoogeography; taxonomic studies; keys to Chilean mammals; field guide to Chilean mammals for country and particular regions; literature for the general public; preparation/ collection technique manuals for the specialist and the general public. Projects underway: Miscellaneous projects dealing with particular species. Comments: A recent bibliography of references on terrestrial Chilean mammals is given in 464. Additional information is given in 495. The research on Chilean mammals has increased significantly in recent years, although it has not been reflected in available publications on the topics of interest in this report. Mann’s guide to mammals of Chile is difficult to obtain. WTC.
Colombia: Instituto de Ciencias Naturales; Museo del Instituto de la Salle. Guides: None. Surveys: 62, 63, 268, 269, 270, 282*; see also
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78, 113, 114, 115, 267, 273, 363. Keys: 62, 63. Technical manuals: 202*. Museum science literature: 159. Children’s literature: Few or rare. Research needed: Natural history studies of mammals; systematic studies on any of the various mammal groups (Rodentia, Ursidae, Cer- copithecidae). Projects underway: None. Comments: Cabrera and Yepes, as well as Cabrera’s systematic manual, are either extremely difficult to obtain and very expensive (the former) or out of print entirely (the latter). Many of the works cited for Colombia are old or are published in English. WTC.
Congo: No response received; but see 401, 489.
Costa Rica: No response received; but see 21 1, 231, 233, 309, 310, 323, 626. Comments: The basic information to form a field guide to the mammals of Costa Rica and, indeed, to produce many other types of basic mammalogical materials is available in diverse publications in English. Costa Rica’s fauna is well-studied when compared with the fauna of most of Latin America. A key to the mammals of Costa Rica was written by Hooper but is generally unavailable. A key to the bats of Costa Rica (572) is also generally unavailable.
Czechoslovakia: Insititute of Systematic Zoology; Institute of Ver- tebrate Zoology; National Museum; Zapadoceske Muzeum v Plzni. Guides: 24*, 185, 196*, 245, 465. Surveys: 185, 239, 244, 245, 465. Keys: 24*, 185, 245, 465. Technical manuals: 583*. Museum science literature: None. Children’s literature: None to many. Research needed: A handbook and key to Czechoslovakian mammals. Projects underway: State research programs on nature conservation and on the fauna of Czechoslovakia; handbook of Czechoslovakian mammals; key to Czechoslovakian mammals. Comments: One response indicated that a specialized museum science journal was available, ‘‘Musejni Prace.” WTC.
Denmark: Natural History Museum; Zoological Museum of the University. Guides: 373, 548. Surveys: 139, 432; see also 140. Keys: Keys from a number of different sources and in a variety of languages are available to the specialist. Non- specialists can use handbooks and field guides for identification. Technical manuals: No special publica- tion; techniques are given in a number of field guides for the general public (youth, teachers, sportsmen). Museum science literature: None. Children’s literature: Many. Projects underway: None. Comments: You always want an up-to-date, comprehensive, and confident treatment of the mammal region, but I think we have a fair picture of the Danish mammal fauna and new information is added every year. The same can be said for most countries in N. W. Europe. In referring to the question on the availability of publications dealing with the operation and importance of natural history museums the following comments
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were made: I can think of no special publications. Museums —including natural history museums— are part of the cultural tradition and de- pending on the actual financial situation in the country it may go up and down for the museum. In one period you may pay special attention to research, in the next to exhibition and other public relation (^sicj e.g., in relation to nature conservation and management. In reference to the question dealing with collecting or preparation techniques manuals, the following comments were made: No special publication but tech- niques are mentioned in a number of field guides for youth, guides for teachers in biology, guides for sportsmen, ‘do it yourself books, ' etc. Generally, collecting and preparing a mammal should not be encour- aged in countries in Western Europe and in many countries here mam- mals not considered game or pest species are totally protected.
Ecuador: No response received', but see 14, 51, 113, 114, 115, 273, 667.
Egypt: Wassif s Collection. Guides: 453. Surveys: None', see also 229, 287, 288, 289, 290, 453, 657, 658, 660. Keys: 525, 536, 537, 538, 539, 540, 541, 542, 659', see also 401. Technical manuals: None. Mu- seum science literature: None. Children’s literature: Few. Research needed: [Mammals of] the eastern and western deserts of Egypt; the Sinai. Projects underway: A study of the Western Desert of Egypt is currently underway and is scheduled to be published by the Desert Re- search Institute, Mataria, Cairo (in Arabic). Comments: WTC.
El Salvador: Museo de Historia Natural de El Salvador. Guides: None. Surveys: 106 (currently unavailable), 181', see also 177, 178, 179, 180, 182, 231, 233, 309. Keys: None', see also 231, 233. Technical manuals: 25. Museum science literature: None. Children’s literature: None. Projects underway: Mammals of the Monte Cristo National Park by Jim Hartman (in preparation); New Bats for El Salvador by V. Hellebuyck (in preparation). Comments: There was also a comment indicating that publications dealing with preparation techniques of mammals are available at the museum library for these specialists. It was noted that these were not available for the general public. WTC.
England: Booth Museum of Natural History; British Museum (Nat- ural History). Guides: 93, 136, 137, 139, 141, 142, 559; see also 392. Surveys: 136, 137, 139, 140, 141, 142, 359', see also 42. Keys: 136, 137, 139, 359; a specialist key is available from Mrs. J. Coy, Depart- ment of Environment, Faunal Remains Project, 63 University Road, Southampton S09 5NH England. Technical manuals: 31, 379, 618*, 652’, see also 138. Museum science literature: Various British Museum (Natural History) publications. Children’s literature: Many. Research needed: A list of named collections, their content and location; cura- torial codes and practices; a national inventory of collections. Projects
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underway: A Manual of Curatorship and a Code of Ethics are being developed by Museums Association, a museums professionals group; one or two keys to skeletal material are available from specialist mu- seums groups in England.
Equatorial Guinea: No response received', but see 380, 40 L
Ethiopia: No response received', but see 143, 280, 355, 401, 690, 691, 692, 693.
Federal Republic of Germany: Coll. Pieper/Kiel; Landessamm- lungen fur Naturkunde; Staatliches Museum fur Naturkunde Stuttgart; Zoologische Staatssammlung Munchen; Zoologisches Forschungsinsti- tut und Museum Alexander Koenig. Guides: iO*, 75*, 92, 195, 325*, 341. Surveys: 325* (for Bavaria), 432. Keys: 30*, 75*, 432. Technical manuals: 148, 149, 471 (and later editions), some information given in field guides. Museum science literature: 527 and several papers in Natur und Museum. Children’s literature: Many. Research needed: Detailed distribution maps; studies of post-cranial osteology; survey of the mam- mals of Germany; a history of museum collections; type catalogues. Projects underway: None. Comments: One respondent noted that there are several small papers dealing with the collection and preparation techniques of mammals that were issued for the public but are of questionable value. WTC.
Finland: Zoological Museum, University of Helsinki; Zoological Museum, University of Oulu. Guides: 547, 549. Surveys: 547, 549, 552. Keys: 549. Technical manuals: 549, 552. Museum science liter- ature: None. Children’s literature: Few to many. Research needed: Zoo- geography of the northern parts of Norway, Sweden, Finland, and So- viet Union; history of the distribution of mammals after the Ice Age; microtaxonomy of mammals today. Projects underway: None. Com- ments: One respondent noted that most children’s literature is trans- lated from other languages. WTC.
France: Laboratoire de Zoologie, Mammiferes et Oiseaux; Musee Zoologique. Guides: 89, 519. Surveys: 519', see also 362, 506. Keys: Response affirmative, but none were listed. Technical manuals: 153, 356*. Museum science literature: 80*, 214. Children’s literature: Many. Research needed: Importance of natural history museums; an atlas of the fauna of France; inventory of the fauna of parks and reserves; an altitudinal distribution of the micromammals in the Vosges Mountain area; a precise list of distributions of the Chiroptera; status of a number of “pest” (hamster) and introduced species (raccoon, raccoon dog, nu- tria). Projects underway: Atlas of the distribution of mammals in France (by the Societe Francaise pour IFtude et la Protection des Mammiferes, S.F.E.P.M.). Comments: WTC.
French Guiana: No response received', but see 99, 113, 114, 115, 273, 668.
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German Democratic Republic: Staatliches Museum fiir Tier- kunde Dresden; Zoologie der Sektion Biowissenschaften der Martin- Luther-Universitat. Guides: JO*, 75*, 194, 195, 707. Surveys: 92, 664, 665; see also 432. Keys: 30*, 75*, 92, 194, 195, 707, Technical man- uals: 471 (and later editions). Museum science literature: None. Chil- dren’s literature: Many. Research needed: Operation of natural history museums. Projects underway: None.
Ghana: University of Ghana. Guides: 77, 155, 247. Surveys: Nu- merous publications but no single one; see also 401. Keys: 401. Tech- nical manuals: Not generally available. Museum science literature: None. Children’s literature: Few. Research needed: Guide to Ghanaian mam- mals; books on ecology and scientific method. Projects underway: None. Comments: There is no lingua franca in Ghana except perhaps English which is the language of literate people; few books are published in Ghanaian languages. Books are not generally available in Ghana an- yway, especially under the present foreign exchange crisis. The respon- dent noted further that some books that could be used as children’s books become available periodically. However, the supply of these ma- terials is extremely erratic. The respondent noted that there is only a single copy of Meester and Setzer (197 1-1977) available in the entire country and this new copy is incomplete. WTC.
Greece: No response received; but see 431, 449, 450, 706.
Guatemala: No response received; but see 210, 231, 233, 273, 309, 317, 585.
Guyana: No response received; but see 113, 114, 115, 243, 273.
Honduras: No response received; but see 68, 197, 273.
Hong Kong: No response received; but see 509.
Iceland: Nattwrufraedistofun islands. Guides: None. Surveys: 162*, 223*, 517, 518, 612, 627*. Keys: None. Technical manuals: None. Museum science literature: None. Children’s literature: Rare. Research needed: Natural history of whales and seals; natural history museum publications. Projects underway: None.
India: Central Arid Zone Research Institute; National Zoological Collection of India. Guides: None. Surveys: 60, 72, 167, 187, 381, 478, 479; see also 96, 97, 98, 169, 328, 351, 482, 573, 574. Keys: 72, 166, 168, 478, 479, 681, 682, 683, 684, 685, 686, 687; see also 163. Tech- nical manuals: 71, 187. Museum science literature: None. Children’s literature: None to rare. Research needed: A checklist of mammals; a field guide to the mammals of India; faunal works. Projects underway: Faunal work on the groups not covered by Pocock and Ellerman; an up-to-date checklist of Indian mammals is being prepared by the Zoo- logical Survey of India. Comments: 128 gives a review of research literature. First International Workshop on Management of Zoological Collections: Recent mammal collections in tropical environments
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sponsored by the Zoologcal Survey of India, Calcutta, and the Carnegie Museum of Natural History, Pittsburgh, USA, was held in Calcutta from 19-25 January 1984. The results of this workshop, which dealt almost entirely with museum topics, will be published in the near future. Two recent publications (423, 6 1 3) are available, which concern endangered and threatened animals of India. WTC.
Indonesia: Museum Zoologicum Bogoriense. Guides: 399 (for Bor- neo). Surveys: 131, 357. Keys: 399. Technical manuals: Use primarily publications of the British Museum (Natural History), American Mu- seum of Natural History, and the Museum of Berlin. Museum science literature: None. Children’s literature: Few. Research needed: Taxon- omy and other information on the biology of mammals; research on methods of small pest control; research on zoonosis. Projects underway: Inventory studies of mammals in Indonesia; studies on the biology of murid rodents as pests on agricultural crops. Comments: The respon- dent noted that there is no formal publication on mammal surveys for the Indonesian Archipelago other than for the island of Borneo. A bibliography of the mammals of Southeast Asia is given in 3 1 5. WTC.
Iran: No response received', but see 100, 147, 360, 414.
Iraq: Natural History Research Centre and Museum. Field guides: None. Surveys: 139, 169, 253, 254, 255, 262', see also 19, 140, 251, 426, 522. Keys: 253, 254, 255. Technical manuals: 255. Museum sci- ence literature: None. Children’s literature: Few. Research needed: Field guide to Iraq's mammals; wildlife of Iraq including modern techniques. Projects underway: None.
Ireland: National Museum of Ireland, Dublin; Trinity College, Dublin. Guides: 88, 136, 435*. Surveys: 175, 176, 418; see also 139, 140. Keys: 136, 422, 568. Technical manuals: 31, 618*, 652. Museum science literature: None, but a publication by the Irish Museums Trust was due to be released in 1982. Children’s literature: Few to many. Research needed: Museum maintenance; special techniques manuals; design and presentation of museum material; simple illustrated litera- ture dealing with the common mammals; guides to specific areas of interest in the Natural History Museum. Projects underway: A book on extant Irish animals has recently been published by the Natural History Museum; a book on the history of the Natural History Museum is expected in the near future. Comments: A book on Irish wild mammals was published as part of the Folens Irish Environmental Library Series. WTC.
Israel: No response received; but see 10, 11, 43, 44, 74.
Italy: Collezione Microteriologica di Longino Cantoli; Museo Civi- co di Storia Naturale; Museo Zoologico de “La Specola.” Guides: 90, 699. Surveys: 619, 620; see also 117, 120. Keys: 619, 620, 699. Tech- nical manuals: 619, 620, 698*. Museum science literature: None; al-
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though some visitors guides are available for a number of museums; see also 118, 119. Children’s literature: None to many. Research need- ed: Literature dealing with training people for collecting, preparing, and maintenance of natural history collections; publications concerned with the importance of natural history museums. Projects underway: Soric- Mae deir ambiente bioclimatico mediterraneo (in press, Contoli); Glir- Mae, Arvicolidae, Muridae delL ambiente bioclimatico mediterraneo (in press, Amori, Contoli, and Cristaldi), Comments: One respondent not- ed that some Italian specialists have field guides currently in press to Italy’s most important mammals. Comments from another respondent indicated that survey books having beautiful illustrations that can be used to explain mammalogy to local and national authorities as well as to common people are vitally needed in Italy, as are books that relate nature to the cultural and moral formation of children and to the general public. Moreover, the respondent felt that books dealing with collecting and preparation techniques are especially vital, as are materials that deal with the exhibition of mammalogical material.
Jamaica: Institute of Jamaica, Guides: None. Surveys: None; see also 231, 233, 27 L Keys: 473; see also 231, 233. Technical manuals: None. Children’s literature: Few. Research needed: Study on the Ja- maican cony, Geocapromys brownii; study of the manatee; natural history study of the mongoose. Projects underway: Studies on the man- atee and cony (Natural Resources and Conservation Division, Dr. Pat- rick Fairbairn).
Japan: Department of Oral Anatomy; Laboratory of Wildlife Re- source Ecology; Natural History Museum; National Science Museum. Guides: 303, 304. Surveys: 1, 303, 304, 349, 616*; see also 2, 448, 656, 696. Keys: 1, 303, 350*, 616*. Technical manuals: 4*, 303. Mu- seum science literature: None. Children’s literature: Few to many. Re- search needed: Complete collections of representative animal faunas; zoogeographic studies; field guides and taxonomic keys to the mammal fauna of the Japanese islands. Projects underway: None. Comments: WTC.
Kenya: National Museum of Kenya. Guides: 151, 155; see also 135. Surveys: 17, 329, 330, 331, 332, 333; see also 8, 252, 334, 335. Keys: 148, 401. Technical manuals: Field manual for museums (UNESCO). Museum science literature: None. Children’s literature: Few. Research needed: Conservation of flora and fauna; ecology and population dy- namics studies of mammals; general natural history studies of native species. Projects underway: None. Comments: WTC.
Korea: Natural History Museum. Guides: 680. Surveys: 679*; see also 318, 319, 425. Keys: 679*. Technical manuals; None. Museum science literature: None. Children’s literature: Rare. Research needed: Field guide geared toward the general public and for education; a
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monograph on the mammals of Korea; guide books in English, Projects underway: None.
Lebanon: No response received; but see 43, 44, 366, 367.
Liberia: No response received; but see 346, 401.
Libya: No response received; but see 240, 293*, 401, 415, 486, 494, 535.
Madagascar: No response received; but see 236, 469, 570*, 607.
Malaysia: Sarawak Museum. Guides: 54, 397; see also 399. Sur- veys: 54, 397; see also 55, 56, 146, 260, 396*, 398, 399, 400, 521, 624. Keys: 396*. Technical manuals: None. Museum science literature: None. Children’s literature: Few. Research needed: Field guide to the mam- mals of Malaysia; collection/ preparation technique manuals; museum science literature. Projects underway: None. Comments: An annotated bibliography of the mammals of Malaysia (116) and a bibliography of the land mammals of Southeast Asia (315) are available. WTC.
Mexico: Escuela Nacional de Ciencias Biologicas; Universidad Na- cional Autdnoma de Mexico. Guides: 365, 645, 647. Surveys: 21, 26, 145, 212, 231, 232, 233, 292, 644; see also 22, 23, 49, 52, 105, 125, 127, 201, 203, 273, 320, 321, 322, 323, 326, 364, 438, 491, 492, 529, 625, 661, 678. Keys: 231, 233, 648. Technical manuals: 202*, 646, Folleto de Divulgacidn, Instituto de Biologia, varias ediciones en Es- panol. Museum science literature; None; but see 50, 493. Children’s literature: Few to many. Research needed: Publications explaining the importance of scientific systematic collections; a brief history of the “Coleccion de mastozoologia del Instituto de Biologia, U.N.A.M. ”; lo- cations of natural history specimens from Mexico that are housed out- side of the country. Projects underway: Mamiferos de Mexico, in prep- aration by Villa- R. Comments: WTC.
Mozambique: No response received; but see 191, 563.
Morocco: Museum de I’lnstitut Scientifique. Guides: 456, 457, 520. Surveys: None; but see 95, 401. Keys: None. Technical manuals: None. Museum science literature: None. Children’s literature: Rare. Projects underway: None.
Namibia: State Museum. Guides: None. Surveys: 401, 544; see also 133, 286, 324, 507, 581. Keys: 170, 401, 503; see also 134. Technical manuals: None. Museum science literature: None. Children’s literature: None to few. Research needed: Semi- popular literature on natural his- tory and conservation of mammals; comprehensive study on the mam- mals of Namibia; field guides to small mammals in national parks; semi-popular literature in non-English languages. Projects underway: The mammals of Namibia (by C. G. Coetzee); check-list of mammals of Etosha National Park (by J. E. W. Dixon); taxonomic study of small mammals of southwest Africa/ Namibia (by the State Museum). Com- ments: One respondent pointed out the need for children’s books and
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Other popular material in the Ovambo language and other Bantu lan- guages that are commonly used in Namibia. WTC.
Nepal: No response received', but see 3, 217, 311, 401, 416, 417, 545, 577, 669.
New Zealand: Auckland Institute and Museum; Canterbury Mu- seum; National Museum of New Zealand. Guides: 48, 199, 200, 480, 504*, 677. Surveys: 48, 199, 200, 208, 677. Keys: 200 (key to marine mammals). Technical manuals: None; only general works on taxider- my; nothing specializing in mammals. Museum science literature: 610*; we subscribe to English and American museum journals such as Cu- rator and Natural History. Children’s literature: Many. Research need- ed: Cetaceans; a guide to the fauna and flora of marine reserves.
Nicaragua: No response received', but see 53, 231, 233, 273, 309.
Niger: No response received', but see 340, 401.
Nigeria: D.C.D. Happold Collection of Mammals. Guides: None. Surveys: 510, 511, 512, 513', see also 69, 246, 248, 249, 250, 299, 401. Keys: 511, 512, 513 (for selected species). Technical manuals: None. Museum science literature: None. Children’s literature: None to rare. Research needed: Field guides at all levels for the vertebrates of Nigeria; conservation literature; general ecological principles in the tropics. Proj- ects underway: Mammals of Nigeria by D.C.D. Happold (in prepara- tion, about 400 pp., Oxford University Press).
Northern Ireland: Ulster Museum and Botanic Gardens. Guides: 88, 142. Surveys: 139', see also 140. Keys: 136, 137. Technical manuals: 379, 480*, 652. Museum science literature: None specifically for North- ern Ireland, although three were listed for England. Children’s litera- ture: Many. Research needed: Archaeozoology and the history of the Irish fauna; osteology in natural history museums; a catalogue of the mammal collections in Britain and Ireland. Projects underway: None. Comments: WTC.
Norway: No response received', but see 283, 284,
Oman: No response received; but see 257,
Pakistan: No response received; but see 505, 546.
Panama: Museo de Ciencias Naturales. Guides: 171, 209, 233, 402. Surveys: None; but see 231, 233, 241, 273, 309. Keys: 402; see also 231, 233. Technical manuals: None. Museum science literature: None. Children’s literature: None. Research needed: Guides to preparation techniques; children’s natural history books; keys to the mammals of Panama. Projects underway: None. Comments: WTC.
Papua New Guinea: Papua New Guinea National Museum and Art Gallery; University of Papua New Guinea. Guides: 103, 104, 275, 357, 406, 500, 590, 598, 601, 634, 655, 701, 703; see also 702. Surveys: 121, 275, 276, 277, 294, 357, 368, 369, 378, 393, 394, 395, 403, 404*, 405, 407, 499, 521, 590, 591, 592, 593, 594, 595, 596, 597, 598, 599,
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600, 601, 602, 603, 604, 605, 606, 628, 629, 630, 632, 633, 634, 703, 704; see also 160, 207, 631, 653. Keys: 295, 370, 404*, 702, 705. Technical manuals: 652. Museum science literature: None. Children’s literature: Few to none. Research needed: Handbook to New Guinea marsupials and monotremes; a guide to the collection and preservation of vertebrates: conservation of natural history specimens in the tropics; the role of museums in community education; the role of museum col- lections in scientific research. Projects underway: None. Comments: WTC.
Paraguay: No response received; but see 70, 113, 114, 115, 273, 424, 671.
People’s Republic of China: Department of Vertebrate Taxonomy and Faunology. Field guides: Surveys: 132; see also 16, 18, 183, 498, 689, 700, 708. Keys: None. Technical manuals: None. Museum science literature: None. Children’s literature: Few. Research needed: An up- dated survey of the fauna of China. Projects underway: A synopsis of Chinese mammals with distributional maps. Comments: WTC.
Peru: No response received; but see 113, 114, 115, 273, 343, 460, 461, 462, 463, 565, 566, 567, 609, 621, 623.
Philippines: No response received; but see 15, 59, 263, 315, 358, 487, 488, 523, 530, 608, 614.
Poland: Mammals Research Institute. Guides: 484. Surveys: 348*, 434*, 55i*. Keys: 348*, 434*, 553*. Technical manuals: 433. Museum science literature: Przeglad Zoologiczny, the journal of the Polish Zoo- logical Society contains articles on this subject. Children’s literature: Many. Research needed: None. Projects underway: None. Comments: WTC.
Portugal: Museu Bocage; Museu e Laboratorio Zoologico. Guides: 20, 150, 455. Surveys: 139; see also 140, 172, 377. Keys: 112, 150. Technical manuals: 652. Museum science literature: Two publications by Almaca and one by Sacarrdo are available; the journal Museum published by UNESCO is available. Children’s literature: Rare to few. Research needed: Evolution; mammalogy; zoogeography. Projects un- derway: None. Comments: One respondent noted that the Museum of Natural History was completely destroyed by fire in March 1978 with no specimens surviving the conflagration, WTC.
Republica Dominicana: Museo Nacional de Historia Natural. Guides: None. Surveys: 233, 636; see also 231. Keys: 231, 473, 551, 577; see also 233. Technical manuals: None. Museum science literature: None. Children’s literature: Few. Research needed: Ecology of the ver- tebrate fauna of the West Indies and the Caribbean; natural history, status and evolution of West Indian mammals. Projects underway: MurciNagos de la Republica Dominicana (J. A. Ottenwalder); status y explotacion del manati en la Republica Dominicana; habitat preference
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of the hutia (Plagiodontia aedium); natural history and systematics of Solenodon; marine mammals of the coast of the Dominican Republic; evolution and natural history of Capromyidae and fossil mammals (by C. A. Woods).
.Republic of South Africa: Albany Museum; Department of Zo- ology ^ University of Cape Town; Jonkershoek Nature Conservation Station; Kaffrarian Museum; Transvaal Museum. Guides: 35, 155, 216, 503, 558. Surveys: 170, 401, 503, 544, 562; see also 307, 514, 515. Keys: 170, 401, 503, 562. Technical manuals: Minor publications are available. Museum science literature: Publications available, es- pecially those published in SAMAB; see also 675. Children’s literature: Rare to many. Research needed: Systematic and zoogeographic studies; multi-lingual childrens literature on conservation and general natural history; techniques books on collection and preservation; a field guide to mammals of South Africa; general natural history publications aimed at the general public. Projects underway: Mammals of the Cape Prov- ince by Swanepoel; Mammals of Transvaal by Rautenbach was pub- lished in 1982; The Wildlife Society of Southern Africa is preparing a field guide to the eastern Cape Coast; Mammals of the Orange Free State by Lynch was published in 1983; Mammals of Natal by Pringle. Comments: WTC.
Romania: “Grigore Antipa” Museum of Natural History. Guides: 88. Surveys: 122, 481. Keys: 237*, 305. Technical manuals: 428*, 458*. Museum science literature: 157, 467*; three pre- 193 5 publications were written by Antipa on the importance of museums. Children’s literature: Few; for a summary of available literature see 157. Research needed: Guide to the small mammals of Europe; establishment of general reg- ulations for organizing and maintaining collections of small mammals; a manual which centralizes many of the methods (morphological, cy- togenetic, etc.) used in the systematics of small mammals. Projects underway: An illustrated guide to the fauna of Romania. Comments: WTC.
Saudi Arabia: No response received; but see 111, 253, 254, 255, 256, 427.
Scotland: Aberdeen University; The Royal Scottish Museum. Guides: 108, 138, 141, 142, 359. Surveys: 142, 501. Keys: 108, 138, 139, 141, 142, 359. Technical manuals: 31, 64, 568, 652. Children’s literature: Many. Research needed: A guide to the Wildlife Protection Act of 1981; guides to the vertebrates of the eastern Mediterranean region. Projects underway: Conference is being held to discuss the im- plications of the Act of Parliament; the lack of field guides is known. Comments: WTC.
Senegal: No response received; but see 76, 158, 291, 401.
Singapore: Zoological Reference Collection. Guides: 258, 400, 624.
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Surveys: iii; see also 398. Keys: 258, 259*, 400. Technical manuals: None. Museum science literature: None. Children’s literature: None. Projects underway: None. Comments: WTC.
Somali Republic: No response received', but see 193, 401.
Spain: Museu de Zoologia de Barcelona. Guides: 91*, 109*, 519. Surveys: 432', see also 638. Keys: 129, 436*. Technical manuals: 9. Museum science literature: None. Children’s literature. Few. Projects underway: None. Comments: WTC.
Sri Lanka: No response received', but see 163, 166, 167, 278, 470.
Sudan: No response received', but see 339, 342, 376, 401, 534.
Suriname: No response received', but see 113, 114, 115, 205, 206, 273, 296, 297, 298, 672, 673.
Switzerland: Naturhistorisches Museum Basel; Zoologisches Mu- seum des Universitat Zurich. Guides: 66, 92. Surveys: 66, 432, 490. Keys: 30*, 66, 75*, 92. Technical manuals: None printed in Switzer- land, but several are available from Germany. Museum science liter- ature: Birkhdusar- Verlag published the “Raritdten and curiositdten der natur, sammlungen der naturhistorischen museum Basel” in 1980. Children’s literature: Few to many. Research needed: Natural history of Swiss mammals; importance of natural history museums; survey on distribution of mammals in Switzerland; a museum guide for children. Projects underway: A children's guide is in preparation by the Natur- historisches Museum. Comments: WTC.
Taiwan: No response received', but see 308*, 316.
Tanzania: College of African Wildlife Management; The Serengeti Research Institute. Guides: 155, 401. Surveys: None', but see 235, 582. Keys: None; keys are available to skulls of some selected mammal groups; some materials available on identification on mammalian teeth. Technical manuals: None. Museum science literature: None. Children’s literature: None to few. Research needed: Keys to the mammals of the region; preparation manuals; guides to the importance of natural his- tory museum. Projects underway: Studies on the ecology and behavior of the lion, cheetah, elephant, and mongoose. Comments: WTC.
Thailand: Division of Environmental Biology. Guides: 361. Sur- veys: None', but see 101, 281. Keys: 361. Technical manuals: A prep- aration manual in Thai was published by the TISTR. Museum science literature: None', see also 611. Children’s literature: Rare. Research needed: Mammals of southeast Asia; revisions of many southeast Asian bats and rodents. Projects underway: None. Comments: A bibliography of the land mammals of Southeast Asia (3 1 5) is available. WTC.
The Netherlands: Rijkamuseum van Natuurlijke Histoire; Zoo- logical Museum. Guides: 87, 107*, 302*, 374*. Surveys: 301, 432, 519', see also 86, 285, 485. Keys: 137, 139, 301, 432, 519. Technical manuals: 285. Museum science literature: None. Children’s literature:
1986
Mares and Braun —Mammalogy Literature
163
Many. Research needed: Key to the identification of mammals in owl pellets; a distribution atlas of mammals; a modern and popular hand- book on mammals. Projects underway: An owl pellet key is in prepa- ration at the Rijksmuseum van Natuurlijke Historic, Leiden; an atlas project has just begun and will result from the collaboration of a large number of institutes, scientists, and amateurs. Comments: WTC.
Trinidad and Tobago: No response received; but see 124, 213, 231, 233, 502.
Turkey: No response received; but see 173*, 174*, 421, 451, 452.
Uruguay: Museo Nacional de Historia Natural de Montevideo. Guides: None; see also 186, 586. Surveys: 353, 688; see also 113, 114, 115, 273. Keys: None; but see 354. Technical manuals: None. Museum science literature: A few articles have been published in the Boletin del Museo Nacional de Historia Natural since 1973. Children’s literature: None. Research needed: Ecology; ethology; biogeography. Projects un- derway: Catdlogo sistemdtico de los vertebrados fdsiles sudamericanos by A. Mones. Comments: WTC.
USSR: Zoologica Museum of Moscow University. Guides: Publi- cations available, but none listed. Surveys: Publications available, but none listed; see 5, 6, 7, 58, 190, 221, 222, 224, 225, 226, 227, 228, 264, 265, 266, 338, 410, 437, 440, 441, 442, 443, 444, 445, 446, 554, 555, 579, 580, 589, 615, 617, 637, 650. Keys: Publications available, but none listed; see 57, 73, 543, 578, 649, 697. Technical manuals: None. Museum science literature: None. Children’s literature: Many.
Venezuela: No response received; but see 110, 113, 114, 115, 242, 273, 371, 439, 508, 526, 587. Comments: Although we received no response to the questionnaire, there are several types of ecological and systematic projects in mammalogy that are currently underway in the country. A key to the mammals of Venezuela was written by Handley but is generally unavailable.
Vietnam: Laboratory of Zoology. Guides: 635. Surveys: 81, 82; see also 123, 516. Keys: 635. Technical manuals: 153. Museum science literature: None. Children’s literature: Few. Projects underway: The rodents of North Vietnam (in French) is in preparation; Key to the mammals of Vietnam by Tien is in preparation and will be published in Vietnamese. Comments: A bibliography of the land mammals of Southeast Asia (315) is available. WTC.
Yemen: No response given; but see 524.
Yugoslavia: Zoology Department Collection and Dr. Dulic’s Col- lection. Guides: 198*. Surveys: 156, 413*; see also 345, 468. Keys: 390, 413*. Technical manuals: Only booklets and papers dealing with preparation of animals in general including also the mammals. One very old one published in 1948 is a small introduction in preparation techniques in general. Author of this booklet is P. Alinger. Museum
164
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Table I. — The availability of field guides (fg), surveys (s), keys (k), and museum science literature (msl) within each geographic region. Percentages are given for each geographic region and are based on the total number of responses received from each region.
|
Response |
Latin Ameri- can |
Euro- pean |
African |
Middle East |
Asian |
Aus- tralian |
|
|
Available in the common |
fg |
17% |
90% |
50% |
62% |
87% |
|
|
language |
s |
50% |
86% |
78% |
62% |
69% |
|
|
k |
39% |
83% |
64% |
69% |
81% |
||
|
msl |
17% |
31% |
7% |
8% |
25% |
||
|
Available, but not in the |
fg |
11% |
5% |
21% |
50% |
8% |
|
|
common language |
s |
28% |
7% |
15% |
50% |
23% |
|
|
k |
11% |
2% |
7% |
100% |
23% |
||
|
msl |
5% |
3% |
7% |
8% |
|||
|
Unavailable |
fg |
72% |
5% |
29% |
50% |
30% |
13% |
|
s |
22% |
7% |
7% |
50% |
15% |
25% |
|
|
k |
59% |
14% |
29% |
8% |
19% |
||
|
msl |
78% |
64% |
86% |
100% |
84% |
75% |
|
|
No response |
fg |
||||||
|
s k |
6% |
||||||
|
msl |
2% |
science literature: None. Children’s literature: Few. Research needed: A good and comprehensive book on the mammalian fauna; compiled lists with distributional data; small monographs for different species or groups of species. Projects underway: Investigation of specific Yugo- slavian mammal species. Comments: The need for a manual on mam- mal preparation techniques was emphasized. WTC.
Zaire: Koninklijk Museum voor Midden- Afrika, Musee Royal de I’Afrique Centrale, Belgium. Guides: 155, 236. Surveys: 531, 532; see also 67, 79*. Keys: 401. Technical manuals: None. Museum science literature: 204; a list of West European museums with important col- lections of African small mammals was given in the supplement to the African Small Mammal Newsletter in July 1981. Children’s literature: Rare. Research needed: Importance of conservation of tropical forests; field guides and general natural history information of small mammals; conservation of mammals. Projects underway: None.
Zambia; No response received; but see 37, 38, 401.
Zimbabwe: Museum of Zoology. Guides: 327 , 556, 559, 674. Sur- veys: None; but see 35, 36, 401, 561*, 563, 564, 674. Keys: 556, 559; see also 184. Technical manuals: None; but see 33, 34, 560. Museum science literature: None; but see 32. Children’s literature: Few. Research needed: Translation of available literature into non-English languages; educational materials. Projects underway: None.
1986
Mares and Braun— Mammalogy Literature
165
Table 2. — The availability of mammal collecting and preparation techniques manuals within each major geographic region. Percentages are given for each geographic region and are based on the total number of responses received from each region.
|
Latin Ameri- |
Euro- |
Middle |
Aus- |
|||
|
Response |
can |
pean |
African |
East |
Asian |
tralian |
|
Available to the specialist in the common language Available to the general public in |
25% |
50% |
27% |
29% |
40% |
|
|
the common language Available to the specialist, but not |
25% |
37% |
13% |
14% |
20% |
|
|
in the common language Available to the general public, |
12% |
4% |
13% |
50% |
21% |
|
|
but not in the common language |
1% |
|||||
|
Generally unavailable |
38% |
8% |
47% |
50% |
36% |
40% |
Literature Availability by Topic
An overview of the responses to each of the questions asked is given in Tables 1-3 for each geographic region. It must be kept in mind that there was often only a small number of responses for any particular country and these may not have been filled out with as much care as might have been desired. Nevertheless, additional library research on the availability of the literature of mammalogy for a country frequently was in accord with the impressions of the in-country specialists as indicated on the returned questionnaires.
The availability of field guides is shown to be quite spotty (Table 1), with Europe and the Australian region having such publications readi-
Table 3. — The availability of natural history literature for children within each of the major geographic regions. Percentages are given for each geographic region and are based on the total number of responses received from each region.
|
Latin Ameri- |
Euro- |
Middle |
Aus- |
|||
|
Response |
can |
pean |
African |
East |
Asian |
tralian |
|
Many in the common language |
60/0 |
62% |
7% |
8% |
38% |
|
|
Few in the common language |
22% |
24% |
36% |
100% |
46% |
56% |
|
Rare in the common language Many, but not in the common language Few, but not in the common lan- |
11% |
2% |
29% |
15% |
||
|
guage Rare, but not in the common |
7% |
8% |
||||
|
language |
7% |
8% |
||||
|
None |
61% |
7% |
14% |
15% |
6% |
|
|
No response |
5% |
166
Annals of Carnegie Museum
VOL. 55
ly available in the common language of the countries queried. In the Australian area, of course, this is English, but in Europe we found that most countries responding had access to field guides that were published in the common language of the country in question. Asia and Africa had a moderate rate of field guide availability and such books were primarily available in the common language of the country. Because of the frequency of having an official language (such as English, for example) listed for a country, however, there arises the problem of availability of such guides to the majority of people who may neither read nor speak the official language. More will be said about this prob- lem when geographic regions are reviewed individually (below). The Middle East and especially Latin America are geographic regions that show a decided scarcity of field guides. Those few that are available are usually not published in the common language. Fully 72% of the respondents in Latin America reported that field guides to mammals are unavailable.
Mares (1982, 1985, in press) has discussed the importance of the technical literature of mammalogy as a basis for the popular literature. A review of the responses concerning the availability of technical lit- erature (Table 1) indicates that such literature is generally more widely available (to the specialist) than is semi-technical literature on mam- mals. All geographic regions reported a fairly broad availability of technical literature on mammals, with Latin American responses re- porting the lowest values. Generally, such literature is available in the common language of the country (except in the Middle East, where our sample size was quite small).
Taxonomic keys are vitally important to the diffusion of mammalogi- cal information beyond a small coterie of specialists. It is keys that allow other biologists to identify their research materials with minimal effort. Such keys are also useful in instructional purposes and to people in government or other fields that need to have a correct determination made for a particular specimen. Most respondents (except for Latin Americans) indicated a fairly good availability of taxonomic keys for use by specialists and generally these were available in the common language of the country (Table 1).
One group of publications that proved to be quite scarce in all geo- graphic regions is collecting and preparation manuals (Table 2). Even in such well-studied areas as Europe and Australia, only 50% or fewer of the respondents indicated the availabilty of such works to the spe- cialist. These types of publications were even less readily available to the general public.
The types of mammalogical publications that turned out to be the least available on a worldwide scale were those that are concerned with
1986
Mares and Braun —Mammalogy Literature
167
the operation and importance of natural history museums (Table 1). The very low values for availability in any language indicate the general lack of literature in this area.
Children’s literature on natural histoiy was shown to be readily avail- able only in Europe (Table 3). In most regions, fewer than 10% of the respondents reported that children’s literature was readily available, while from 20-50% of them said such books were few.
Literature Availability by Region
Middle East. --Only two of five questionnaires sent to countries of the Middle East were returned, so it is not possible to speak in any detail about the status of the literature of this region. Each of the four countries surveyed has had foundational literature published on mam- mals, although most of this literature is available only in European languages. Nevertheless, although there are few guides to the mammals of these countries and apparently almost no museum literature or pop- ular literature published in the native languages, a rather firm foun- dation for future workers has been established.
Latin Twerira.— Eighteen of 32 questionnaires were returned from 10 countries. Unfortunately, this is not a large sample for such a vast region, but responses were received from Mexico, Central America, the Caribbean, and South America, giving a good geographical repre- sentation. In almost all literature areas surveyed, Latin America was ranked at the lowest levels. Most countries lack even the foundational literature, regardless of its language of publication. Few faunal surveys have been published for most countries, taxonomic keys and field guides are generally unavailable, museum literature is practically non- existent, and children’s literature is rare to unavailable. These trends are particularly pronounced for South America, as opposed to Central America, the Caribbean, and Mexico. Because of their proximity to North America, these regions have been fairly well-studied, especially Mexico, the Caribbean, Panama, and Costa Rica. Unfortunately, de- spite the availability of foundational literature (largely published in English), there are as yet relatively few publications available of the nature of field guides, children’s books, and general museum publi- cations. Indeed, even in Mexico, perhaps the best-studied of the Latin nations, semitechnical and popular publications are either uncommon or hard to obtain. For South America, the panorama is much less positive. There, even the foundational literature is lacking and, as has been argued by Mares (in press), without this most basic literature of mammalogy, the preparation of the literature needed to educate the general population is impossible. Among the South American coun- tries, only Suriname, Venezuela, Uruguay, Chile, and Argentina have
168
Annals of Carnegie Museum
VOL. 55
had an important portion of their foundational literature completed. Even in these few countries, however, the semitechnical and popular literature on mammals is largely unavailable.
Europe. —YovXy -two of 66 questionnaires were returned from Eu- ropean collections, for a 64% response rate. Of the regions surveyed, Europe led in all categories of literature availability, reflecting a long history of nature study, museum collection formation, resident spe- cialists on mammals, and so on. Generally, basic literature as well as the semitechnical and popular literature were available in the common language of each country.
4/nca.— Fourteen of 23 (61%) African collections queried returned the questionnaires, with six responses coming from two countries, Na- mibia and the Republic of South Africa. Many African countries were thus not sampled due either to the fact that questionnaires were not returned or that a particular country had no collection-oriented mam- malogist to be queried. There is little doubt that the inclusion of four responses from the Republic of South Africa skews the African data. Still, it is evident that the African area (excluding the Middle East) has a very poor literature base, particularly for works published in non- European languages. Although 50% of the respondents indicated the availability of field guides to the mammals of their particular country, published in the common language, this number is inflated. Many cited Dorst and Dandelot (1970), for example, which is limited to the larger mammals of Africa. Fully 86% of the respondents reported that tech- nical publications were available in the common language of the coun- try, and 83% said that taxonomic keys were readily available (Table 1), but again these are probably overestimations due to the fact that some well-surveyed countries responded (for example, Kenya, Repub- lic of South Africa).
Australian region. — By and large, the Australian area supports a rich and varied literature at the technical, semitechnical, and popular levels. Field guides, technical publications, taxonomic keys, and collecting and preparation manuals were listed as readily available, as was chil- dren’s literature.
In answer to the question requesting a listing of subject areas needing research, responses were understandably mixed. Topics suggested with regularity were surveys and technical publications on a country’s fauna; taxonomic revisions; the production of both field guides and taxonomic keys; educational materials aimed at the general public and children that deal with museum/collection-related topics and natural history; theoretical and applied ecological research (including conservation); collecting and preparation technique manuals; and behavioral research.
1986
Mares and Braun— Mammalogy Literature
169
Socioeconomic Factors
In any broad examination dealing with some facet of life in the diverse nations of the world, it is instructive to examine what country- wide phenomena might be related to the patterns that have been iden- tified. We have shown that the world’s literature on mammalogical subjects has an extremely spotty distribution. We might ask if there are any obvious measures of a country’s socioeconomic profile that might be related to the availability of this type of literature. It takes no special understanding of economics or politics, for example, to see that those countries generally classified among the economically de- veloped nations of the world (DC’s) have a higher rate of literature availability than do those nations generally considered as being under- developed (UDC’s).
We have examined some basic socioeconomic factors of each nation that responded to our questionnaire (Table 4). These data include in- formation on Gross National Product (GNP), per capita income (PCI), the natural rate of population increase (r), percent literacy, population density, and the percent of the population that is urban. Utilizing the SPSS statistical package (SPSS Inc., 1983), we compared these data to an index of literature availability via multiple regression analysis and stepwise multiple regression analysis. In order to do this, it was necessary to score the responses from each country to each of the first six questions on the questionnaire (mean values were used in the case of multiple responses). Scores on each question varied from zero for an unavailability of literature to five for an abundance of literature. Scores for each question were summed to obtain the country’s total score (scores may thus vary from zero to 30). Literature availability scores for those nations that responded fully to a questionnaire are given in Table 4.
When multiple regression analysis was run so that all independent variables (the socioeconomic data of Table 4) were examined for their effect on explaining the variance in the dependent variable (the liter- ature availability scores of Table 4), two factors were shown to have statistical importance in explaining variance in literature availability: the rate of population increase, r, with P = 0.04; and the per capita income, with P = 0.05. The sign of the beta value for r was negative, whereas it was positive for the beta values of PCI (Table 5). This means that the greater the value for r, the lower the availability of literature, and the greater the PCI, the more literature that is available in a country. In this analysis, each variable was examined for its effect on the de- pendent variable, while all other variables were held constant.
In a separate stepwise multiple regression analysis, the single most
Table A. — Gross national product (GNP), per capita income, natural increase in population, percent literacy, population density, percent urban population, and the literature availability score given for most countries that responded to questionnaires. Countries are listed alphabetically within each region. The year in which the data were obtained is given in parentheses. Unnumbered countries lack sufficient
VOL. 55
170
Annals of Carnegie Museum
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1986
Mares and Braun— Mammalogy Literature
171
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172
Annals of Carnegie Museum
VOL. 55
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1986
Mares and Braun— Mammalogy Literature
173
Table 5.— Multiple regression analysis of the socioeconomic data versus the index of literature availability from Table 4 for countries that responded fully to the questionnaire. B = slope; SE B = standard error of B; P = level of significance; Beta is a standardized B.
|
Variable |
B |
SE B |
Beta |
T |
p |
|
Urban population |
-0.014 |
0.031 |
-0.063 |
-0.446 |
0.66 |
|
GNP |
-0.004 |
0.003 |
0.187 |
1.529 |
0.13 |
|
Population density |
0.49 xlO |
0.49 X 10 |
-0.121 |
-1.017 |
0.32 |
|
PCI |
0.55 X 10 |
0.27 X 10 |
0.349 |
1.975 |
0.05 |
|
Literacy |
-0.054 |
0.048 |
-0.223 |
-1.128 |
0.26 |
|
r |
-2.560 |
1.233 |
-0.475 |
-2.077 |
0.04 |
|
(Constant) |
20.421 |
5.497 |
3.715 |
important independent variable in explaining variance in the depen- dent variable was per capita income (P < 0.000 1): all other independent variables were non-significant. In this analysis, = 0.33 and the beta value was 0.91 x 10“^. Since only one independent variable was used, beta values can be used to show the effect of PCI on literature avail- ability. We can see that if per capita income in a country were increased by $ 100/year, the availability of mammal literature would increase by almost 10%. Increasing PCI by $ 1000/year would essentially double the availability of literature in a particular society.
Discussion
Clearly, there is a great need to increase the availability of all types of mammalogical literature throughout the world. In many places (for example, Latin America), even the foundational literature is lacking, thus making the development of a more broad-based and diverse lit- erature more difficult. Many countries lack active mammalogists and do not support a working collection of mammals. In other countries, such collections, if they exist at all, are very poorly supported, ill- housed, and understaffed. Research, such as coordinated faunal surveys and basic taxonomic studies, is not encouraged. Given these conditions, the general mammalogical panorama is bleak and the foundational literature of mammalogy, that most basic literature that is the basis for more advanced topics such as ecology, physiology, behavior, and conservation, is unavailable. The questions that we asked of each cu- rator covered both the basic literature as well as the more advanced and popular literature. It is consistent with the views of Mares (1985, in press) and with the points made earlier in this paper, that every country that reported a rich popular literature also reported a very broad basic literature. No country having a weak foundational literature reported an abundance of field guides, children’s books, or other semi-
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technical or popular works. Very few countries reporting a strong foun- dational literature had a weak popular literature (exceptions are Ghana, Papua New Guinea, Portugal, Republic of South Africa, and Singa- pore). The close relationship between the foundational literature and the extremely important literature of the general public appears ines- capable. It seems unlikely that a country could begin to publish popular faunal works on a large scale without first developing the infrastructure of mammalogy “the collections, the faunal surveys, and the taxonomic research.
We have shown that there is a great disparity in the way in which the literature of mammalogy is dispersed throughout the world. Lit- erature availability does not appear to follow political ideologies, al- though there is a strong geographic component to the data. Holarctic countries generally have a strong literature in mammalogy —this lit- erature is widely available in the common language of the country and includes both technical and popular literature. If a country is located north of 35®N latitude, it shows a strong literature at almost all levels. Countries south of this line, with notable exceptions (for example, the Australian region and some few African, Asian, and Latin American countries), lack a strong literature.
The response to the question dealing with suggested research topics supports our analysis as to the importance of the foundational litera- ture. In countries having a rich literature, suggested research topics were often well-formulated and specific. Often they involved fine tuning of ongoing research efforts, or studies of sweeping scope that were synthetic in outlook. In countries having a poor literature, suggested topics involved the most basic types of research, taxonomic surveys, the construction of taxonomic keys, and so forth.
The analysis of the socioeconomic factors that might affect literature availability points out some interesting patterns. First, although some of the data items utilized may be intercorrelated, and even though all of these factors are the result of complex sociopolitical components, there are only two items of statistical significance that seem to affect literature availability, and these two items are themselves related. Per capita income and the rate of population increase would seem to be, a priori, inversely related. The faster a population is growing, the lower the rate of income per person. It is this latter statistic, PCI, that has such an important effect on literature availability. As per capita income increases, so does the literature of mammalogy become more diverse. We suspect that although we limited our analysis to literature on mam- mals, the same trends will be shown for all natural history literature. The higher the PCI, the greater the availability of literature on nature.
It is instructive to plot literature availability (as shown by the index) and PCI of countries (Fig. 1). There is a cluster of countries that have
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PER CAPITA INCOME ($1000)
Fig. L— A plot of per capita income (PCI) versus an index of literature availability for tbe countries of the world that responded to the questionnaire. Numbers of countries refer to Table 4. The asymptotic curve (fitted by eye) shows that after a PCI of about $4000 per annum, literature availability reaches a relatively stable high level.
both a low per capita income and a low level of literature availability. With few exceptions, most countries that have a diverse literature on mammals have PCFs above $5000 US per year. Only about 1 1 coun- tries show an anomalous relationship of PCI and literature availability. Several socialist nations (that is, Czechoslovakia, USSR, Poland, Ro- mania, and Yugoslavia) show a rather abundant literature with rela- tively low PCI. Given the fact that PCI in a socialist nation is low due to the many social services that are covered through governmental programs, and because scientific education is strongly supported in these countries, it is not surprising that a higher level of literature availability is maintained with a lower PCI.
Singapore, a city/nation only since 1965, might be expected to sup- port a poor literature, because few studies concern themselves with mammals of a densely populated urban area. Another country that deviates from the expected relationship, Iceland, is also easily ex- plained. Although the country has a high PCI, it has a depauperate mammal fauna, thus detailed works on mammals are rare.
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Several other nations that do not fit the observed pattern are the Republic of South Africa, where the PCI is quite low due to the inclu- sion of Black incomes in the overall PCI, while White incomes are quite high. The literature is primarily produced by and for Whites, thus the unusual location of the PCI versus literature value is easily explained.
Finally, Mexico, Portugal, and Papua New Guinea also seem to deviate from the general pattern. Mexico is well-studied due to its proximity to the United States, whose mammalogists have worked in Mexico for almost a century. Similarly, Papua New Guinea has had much of its fauna studied by colonial powers, such as England and Germany, as well as by Australians. Portugal, although having a low PCI at present (partially due to the fact that almost one-third of the labor force is composed of agricultural workers), has had a history of European research efforts on mammals in the recent past.
In general then, it appears that without a marked increase in per capita income, mammal literature will not increase. Obviously, this does not reflect a 1:1 relationship between PCI and literature avail- ability. Apparently, scientific and educational resources in any country can only be allocated after basic societal needs are being met. PCI is a measure of how well such needs are being met, especially in a non- socialist society. If PCI is too low, few people are able to learn to read, to complete school, or to go into a complex field of study such as science. In countries with low PCI’s, higher education will be a luxury, and research on mammals or other animal groups will be uncommon, because such studies are not usually viewed as being vital to the national interest. With few students, fewer professors, and a lack of support for natural history studies, literature availability will perforce be at min- imal levels in these countries.
Can a country decide to reverse this pattern and produce the popular literature that is needed to educate the masses about the importance of their natural environment without increasing PCI? Our results sug- gest that this is not a simple task. The only exceptions to the perceived trend were in countries where PCI is a poor indicator of living standard (socialist countries) or where foundational research had been completed in the past due to historical accident (for example, Mexico). The PCI/ literature relationship seems to be a strong one. Elevating the PCI clearly influences a multiplicity of societal factors in education, com- merce, science, government, and so on. But there appears to be no ready mechanism whereby PCI can be bypassed with a resultant flow- ering of literature. The gradual elevation of the PCI (insofar as it reflects economic health) very likely leads to a society that has taken care of its basic economic needs and has developed the time and inclination
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to dedicate resources to what are, from the standpoint of the basic requirements of life, rather esoteric pursuits.
An important point in this analysis is that even though the best m,echanism for increasing the availability of natural history literature is to raise the PCI, there is another way to elevate the overall level of literature in any country. Supporting basic research by scientists of any nation in a particular countiy will, ultimately, yield a significant amount of foundational literature. This literature will later be used by nationals of the country as building blocks for a more advanced and compre- hensive national literature. Mexico is an excellent example of how a country with a long history of basic research by scientists from another nation, in this case the United States, is able to utilize the work of these foreign scientists to develop its own cadre of biologists and the diversity of its particular literature on nature, despite its low PCI. Papua New Guinea is another example of this same occurrence.
Our results lead us to be cautiously optimistic. If international co- operative research efforts on basic systematics, natural historical, and biogeographical research can be significantly increased, the founda- tional literature of each country can be established. The very positive response by almost all curators to the possibility of cooperative re- search/publication efforts indicates that the door to scientific interac- tion is open. Colleagues throughout the world are eager to participate with foreign scientists who might have expertise working with the fauna of a particular country. Together, these biologists can act to produce the keystone of the science of natural history, the foundational literature upon which all more advanced studies are based and upon which the popular literature of a country rests.
Acknowledgments
We thank Dr. Thomas E. Lacher, Jr., for statistical advice and Mrs. Sonya Johnson for typing the manuscript. We also express our gratitude to the many curators who took the time to respond to our questionnaire.
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Back issues of many Annals of Carnegie Museum articles are available, and a few early complete volumes and parts are listed at half price. Orders and inquiries should be addressed to: Publications Secretary, Carnegie Museum, 4400 Forbes Avenue, Pittsburgh, Pa. 15213.
0/ CARNEGIE MUSEUM
CARNEGIE MUSEUM OF NATURAE HISTORY
4400 FORBES AVENUE « PITTSBURGH, PENNSYLVANIA 15213 VOLUME 55 7 November 1986 ARTICLE 1 1
CATALOG OF THE RECENT MARINE MAMMALS IN THE CARNEGIE MUSEUM OF NATURAL HISTORY
Suzanne B. McLaren
Collection Manager, Section of Mammals
Duane A. Schlitter
Curator, Section of Mammals
Hugh H. Genoways
Research Associate, Section of Mammals
Abstract
The Section of Mammals, Carnegie Museum of Museum of Natural History, houses 6 1 2 specimens of Recent marine mammals, including 34 cetaceans, 44 polar bears, six sea otters, 499 pinnipeds, and 29 manatees. Families represented in the Order Cetacea include Platanistidae, Physeteridae, Monodontidae, Delphinidae, Phocaenidae, and Bal- aenopteridae. Families represented in the Order Pinnipedia include Otariidae, Odobeni- dae, and Phocidae. The single family Trichechidae represents the Order Sirenia in the collection. For each specimen the following data are recorded: date collected, catalog number, sex, age or condylobasal length, nature of specimen, condition of specimen, and comments. The latter category gives information on the condition of the skin and skeletal material, and also explains the availability of the specimen.
INTRODUCTION
The Section of Mammals, Carnegie Museum of Natural History, currently holds 6 1 2 specimens of Recent marine mammals, including 34 cetaceans, 44 polar bears, six sea otters, 499 pinnipeds, and 29 manatees. The majority of the collection of marine mammals consists of specimens preserved as skulls or disarticulated skeletons. However,
Submitted 9 June 1986.
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the collection also contains 151 skins and five fluid-preserved speci- mens as well as body mounts and articulated skeletons. The specimens are housed primarily in the fur vault and large osteological storage area of the Section of Mammals, but all mounted material is on display at this time in the public galleries of the Museum.
The primary purpose of this catalog is to make the availability of this material known to the scientific community. It is extremely difficult to obtain new specimens of any marine mammal because most species are considered to be threatened with extinction (Berger et al., 1979). All species are placed in at least Appendix II (Threatened) under the Convention on International Trade in Endangered Species and are protected under the various Marine Mammal Acts beginning as early as 1967. Therefore, it is necessary for researchers to make maximum use of the materials already stored in museum collections. Because specimens stored in this collection are essentially irreplaceable, none is available for study on loan. However, all qualified researchers are welcome to study any of the specimens at the collection facilities of the Section of Mammals.
All collection data files of the Section of Mammals have been com- puterized. This catalog presents selected data categories of which some are available on the computer file, but others have been specifically generated for this publication. Other categories of data are also available upon request in printout or machine-readable form. The format of this catalog is patterned after Napier’s (1976, 1981, 1985) Catalogue of the Primates in the British Museum (Natural History) and the McLaren et al. (1984) Catalog of the Recent Scandentia and Primates in the Carnegie Museum of Natural History.
Classification
The collection of the Section of Mammals and this catalog follow G. G. Simpson’s (1945) classification in the Principles of Classification and the Classification of Mammals through the level of subfamily. Below subfamily, specimens are arranged alphabetically by genus. Spec- imens stored in the osteological collection and the fur vault are then arranged numerically within the genus. This catalog is arranged alpha- betically by genus, species, and in a few cases subspecies, so that re- searchers may easily see what is available for each taxon. Beyond this taxonomic arrangement all specimens are presented in numerical order. It is hoped that by arranging the catalog in this way that work by researchers in the collection will be facilitated.
Cetaceans in the collection have been examined and identified by C. W. Potter and G. S. Morgan. The Odobenus have been identified, and sexed in some cases, by Francis Fay. Hall (1980) has been used for
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identification of the Pinnipedia, but scientific names follow Honacki et al. (1982) except in the case of Physeter catodon (Schevill, 1986).
Explanation of Headings
Locality. —Collection localities appear as listed on the original skin and skull tags. Latitude and longitude were added to numerous localities as listed in the most recent edition of the Standard Names Gazetteer for localities outside of Canada (printed by the United States Board of Geographic Names). Gazetteer of Canada (printed by the Ca- nadian Permanent Committee on Geographical Names) was used for referencing Ca- nadian localities.
Of the 6 1 2 marine mammals, 3 1 were zoo specimens. In many cases, original localities are not known for these animals. To facilitate recognition of locality information per- taining to captive specimens, these data are followed by an asterisk.
Date of collection.— ¥ or wild taken specimens, this date refers to the day on which the animal was obtained. In nearly all cases for zoo specimens, the only date which is available is the day on which the animal died in captivity. In rare cases when more information is known about a particular captive specimen, those data will be shown under the “Comments” category.
Three symbols are used for denoting sex: F, M, ? (undetermined by original collector). Where not denoted on the original tag, sex has sometimes been inferred based on the examination of adult skulls. This practice has not been used for the preparation of the current catalog, but in instances where such conclusions have been drawn pre- viously, the appropriate symbol will be enclosed in brackets.
Age or condylobasal length.— Dming the preparation of this catalog, we were advised by R, L. Brownell to replace age with condylobasal length. Whenever possible this measurement was taken. If minor damage made precise measurement impossible, a rough measurement is shown in brackets. If skull damage was too severe to make this practical, or when only skins were available, a judgement based on tooth eruption, fusion of sutures, or pelage was sometimes possible. Additionally, original collectors’ notes on specimen tags were sometimes utilized in an effort to provide potential researchers with an idea of whether a specimen was immature [Imm] or adult [Ad]. Immature specimens were recognized with most certainty. When satisfactory judgement was not possible, a question mark (?) appears in this category. In one series of Phoca hispida hispida spec- imens, collector J. K. Doutt noted age approximations (in years) as told to him by local Inuit hunters. These estimates have been included in brackets under this category, if measurements cannot be taken. When condylobasal length can be measured, these es- timates appear in brackets under the “Comments” category.
Nature of specimen. —This category describes the type of preservation used for each specimen and corresponds to the two letter abbreviation system used on our computer file. The following list describes each type of preservation used in the marine mammal collection: SK = skull only; SO = skin only; SS = skin and skull; SB = skin, skull, and body skeleton; SN = complete skeleton; CO = cranium only; AL = alcoholic (preserved in 70% ethyl alcohol); BM = body mount; PS = partial skeleton. If peculiarities exist regarding availability of a specimen these are noted by an asterisk following the two letter code. There are two types of specimens that carry the asterisk: 1) specimens for which some of the parts are missing; 2) specimens on exhibit. These latter specimens are available for examination but work must be planned with the understanding that the specimens are in exhibit areas and are not housed with the remainder of the collection. For further explanations regarding the availability status of a specimen refer to the “Comments” category.
Condition of specimen and comments. — This category is designed to inform the reader of the usefulness of a specimen for systematics research. The term “Good” is used for
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all specimens and specimen parts, which are in useable condition for study. Damage to the skull is described so that measurability can be evaluated. All skins are either tanned or body mounts with the exception of a few newborn polar bears which have been prepared as conventional study skins. Captives are noted; missing parts and exhibit specimens are noted. Inuit age estimates of Phoca hispida hispida specimens are included here.
Taxonomic List
Cetacea
Platanistidae
Iniinae
Inia geoffrensis Physeteridae Physeterinae Physeter catodon Monodontidae
Delphinapterus leucas Monodon monoceros Delphinidae
Delphinus delphis Sotalia fluviatilis Tursiops truncatus Phocaenidae
Phocaena phocaena Balaenopteridae
Balaenoptera musculus Carnivora Ursidae
Ursus maritimus
Mustelidae
Lutrinae
Enhydra lutris lutris Pinnipedia Otariidae
Callorhinus ursinus Eumetopias jubatus Otaria byronia Zalophus californicus
Odobenidae
Odobenus rosmarus diver- gens
Odobenus rosmarus ros- marus Phocidae Phocinae
Erignathus barbatus Halichoerus grypus Phoca fasciata Phoca groenlandica groen- landica
Phoca hispida hispida Phoca sibirica Phoca vitulina concolor Phoca vitulina mellonae Phoca vitulina richardii Phoca vitulina vitulina Phoca vitulina Phoca species Cystophorinae
Cystophora cristata Sirenia Trichechidae
Trichechus inunguis Trichechus manatus lati- rostris
Trichechus senegalensis
Acknowledgments
During the curation of the marine mammal collections and the subsequent updating of the computer file, the position held by Suzanne McLaren was supported by National Science Foundation Grant BSR-81 1 1553. We would like to express our thanks to Dr. Darryl Domning, Dr. Francis Fay, Gary S. Morgan, and Charles W. Potter for their assistance in identification of portions of the marine mammal collection. We also ap-
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predate the assistance of Joseph Bissonnette in helping to pinpoint original collecting localities for several zoo specimens.
Literature Cited
Berger, T, J., A. M. Neuner, and S. R. Edwards. 1979. Directory of federally con- trolled species. Assoc. Syst. Collections, Lawrence, Kansas, iii + 1--6 + MA 1--MA
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Doutt, J. K. 1942. A review of the genus Phoca. Ann. Carnegie Mus., 29:61-125. Hall, E. R. 1980. The mammals of North America. John Wiley and Sons, New York, 2nd ed., 2:601-1181.
Honacki, j. H., K. E. Kinman, and J. W. Koeppl (eds.). 1982. Mammal species of the world. Allen Press, Inc., and Assoc. Syst. Collections, Lawrence, Kansas, ix + 694 pp.
McLaren, S. B., D. A. Schlitter, and H. H. Genoways. 1984. Catalog of the Recent
Scandentia and Primates in the Carnegie Museum of Natural History. Ann. Carnegie Mus., 53:463-525.
Napier, P. H. 1976. Catalogue of Primates in the British Museum (Natural History). Part I: Families Callitrichidae and Cebidae. British Museum (Natural History), London, x + 1 2 1 pp.
— — — . 1981. Catalogue of Primates in the British Museum (Natural History) and elsewhere in the British Isles. Part II: Family Cercopithecidae, Subfamily Cerco- pithecinae. British Museum (Natural History), London, vii + 203 pp.
— — . 1985. Catalogue of Primates in the British Museum (Natural History) and elsewhere in the British Isles. Part III: Family Cercopithecidae, Subfamily Colobinae. British Museum (Natural History), London, x + 1 1 1 pp.
ScHEViLL, W. E. 1986. The International Code of Zoological Nomenclature and a paradigm: The name Physeter catodon Linnaeus 1785. Marine Mammal Science, 2: 153-157.
Simpson, G. G. 1945. The principles of classification and a classification of mammals.
Bull. American Mus. Nat. Hist., 85:i + 1-350.
Williams, S. H. 1928. A river dolphin from Kartabo, Bartica District, British Guiana. Zoologica, 7(4): 105-1 28.
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incomplete for normal animal; zoo specimen
Locality unknown* Jul 1968 CM 60939 F [331] SB Rostrum damaged, brittle; tanned
skin; zoo specimen
CBL Nature of
Locality Date collected Catalog number Sex or age specimen Condition of specimen and comments
Locality unknown* Jui 1968 CM 60940 F [340] SB Skull autopsied; rostram damaged;
244
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VOL. 55
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missing
Locality unknown* 22 Nov 1915 CM 3222 M [Imm] SS Zoo specimen; newborn
Canada: Quebec; Great Whale Aug 1915 CM 3393 ? 280.7 SK Several teeth broken or missing; left
River zygomatic arch broken
CBL Nature of
Locality Date collected Catalog number Sex or age specimen Condition of specimen and comments
1986
McLaren et al.— Catalog of Marine Mammals
245
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James Bay, South Twin Is- land
Canada: Northwest Territories; 23 Jul 1935 CM 11287 M 356.2 SS Braincase damaged; tanned skin
246
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VOL. 55
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1986
McLaren et al.™ Catalog of Marine Mammals
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1986
McLaren et al.— Catalog of Marine Mammals
249
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icality unknown* Dec 1907 CM 1562 F 218.0 SB* Most teeth loose; zoo specimen; skin
missing
CBL Nature of
Locality Date collected Catalog number Sex or age specimen Condition of specimen and comments
250
Annals of Carnegie Museum
VOL. 55
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252
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254
Annals of Carnegie Museum
VOL. 55
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Canada: Quebec; Povungnituk 28 Jun 1945 CM 23307 ? 194.8 SK Left side of rostrum broken
Post 59®40'N; 77°30'W
Canada: Quebec; Povungnituk 6 Jul 1945 CM 23308 ? 230.4 SK Good, but all teeth gone
Post 59°40'N; 77°30'W
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256
Annals of Carnegie Museum
VOL. 55
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Canada: Quebec; 5 mi below Mar 1938 CM 15223 ? [Imm] SO Tanned skin
mouth of Great Whale River
Canada: Quebec; 5 mi south of 28 Feb 1938 CM 15224 ? 108.7 SS Good; tanned skin
Great Whale River
Catalog of Marine Mammais— Continued.
262
Annals of Carnegie Museum
VOL. 55
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Annals of Carnegie Museum
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272
Annals of Carnegie Museum
VOL. 55
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Post 59°40'N; 77°30'W baculum saved
Canada: Quebec; Povungnituk 22 May 1945 CM 23103 F 153.2 SK Most teeth loose; rostrum damaged
Post 59°40'N; 77°30'W
274
Annals of Carnegie Museum
VOL. 55
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CBL Nature of
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1986
|
McLaren et al. — |
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CBL Nature of
Locality Date collected Catalog number Sex or age specimen Condition of specimen and comments
276
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Annals of Carnegie Museum
VOL. 55
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luk Bay 59°40'N; 77°10'W
Canada: Quebec; mouth of Jun 1945 CM 23181 F 159.3 SK Good; cross-section of canine saved
Kogaluk River, near Koga- luk Bay 59®40'N; 77°10'W
Catalog of Marine Continued.
280
Annals of Carnegie Museum
VOL. 55
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282
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283
Canada: Quebec; Povungnituk 27 Jun 1945 CM 23235 ? [Imm] SK Skull fragmentary
Post 59°40'N; 77®30'W
Canada: Quebec; Povungnituk 27 Jun 1945 CM 23236 ? 150.4 SK Good; cross-section of canine saved;
Post 59°40'N; 77°30'W numerous loose teeth
CBL Nature of
Locality Date collected Catalog number Sex or age specimen Condition of specimen and comments
284
Annals of Carnegie Museum vol. 55
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CBL Nature of
Locality Date collected Catalog number Sex or age specimen Condition of specimen and comments
1986
McLaren et al.— Catalog of Marine Mammals
285
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286
Annals of Carnegie Museum
VOL. 55
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1986
McLaren et al.™ Catalog of Marine Mammals
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290
Annals of Carnegie Museum
VOL. 55
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1986
McLaren et al.™ Catalog of Mamne Mammals
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CBL Nature of
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292
Annals of Carnegie Museum
VOL. 55
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McLaren et al.— Catalog of Marine Mammals
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294
Annals of Carnegie Museum
VOL. 55
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Trichechus senegaiensis
Cameroun: 10 mi W Edea 1942 CM 42762 ? 351.3 SK Bullar regions damaged
ANNALS"*-"
0/ CARNEGIE MUSEUM
CARNEGIE MUSEUM OF NATURAL HISTORY
4400 FORBES AVENUE » PITTSBURGH, PENNSYLVANIA 15213 VOLUME 55 7 November 1986 ARTICLE 12
TAXONOMIC AND DISTRIBUTIONAL NOTES ON BATS FROM KENYA
Duane A. Schlitter
Curator, Section of Mammals
ISSA R. Aggundey*
Mazin B. Qumsiyeh^
Rea Postdoctoral Fellow, Section of Mammals
Kimberlyn Nelson^
Rodney L. Honeycutt^
Abstract
Taxonomic and distributional comments are given for six species in three families of bats from Kenya. Two species {Hipposideros camerunensis and Tadarida russata) are reported from Kenya for the first time.
Introduction
During field work on the systematics of small mammals in Kenya from September to November 1985, a number of bats were collected
‘ Curator of Mammals, National Museums of Kenya, P.O. Box 40658, Nairobi, Kenya. ^ Current address: Department of Biological Sciences, Texas Tech University, Lubbock, TX 79409.
^ Address: Museum of Comparative Zoology, Harvard University, Cambridge, MA 02138. Submitted 5 February 1986.
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in various parts of Kenya, Some of these specimens add substantially to the knowledge of the distribution of bats in Kenya and East Africa while others were of taxonomic interest. This paper summarizes these data on six species of bats in three families.
Materials and Methods
All specimens reported herein were captured with mist nets set in the normal manner. Cranial and forearm measurements were taken by means of dial calipers and are given in millimeters. All specimens are deposited in the Carnegie Museum of Natural History, Pittsburgh (CM); Museum of Comparative Zoology, Harvard University, Cambridge (MCZ); and National Museums of Kenya, Nairobi (NMK). Coordinates for localities are listed only for the first reference to the locality.
Account of Species
Family Hipposideridae Hipposideros camerunensis Eisentraut, 1956
Hipposideros camerunensis Eisentraut, 1956. Zool. Jahrb. Abt. Syst. Oekol. Geogr. Tiere, 84:526. Near Buea, Cameroun.
Records (1). — Kenya: Western Prov., Kakamega Dist., Kakamega Forest Station, 3- 1/2 km E Kakamega (0°14'N, 34°52'E) (1 CM).
Measurements. ~Sq\qqXq6. measurements of an adult female, length of forearm, 74.3; condylocanine length, 24.9; zygomatic breadth, 15.7; alveolar length of maxillary toothrow, 9.3; and greatest crown breadth of upper molar toothrows, 11.1.
Remarks. —Hipposideros camerunensis is a rare, large forest species of Hipposideros resembling closely the more common H. cyclops. It has been reported from the type locality, Buea, Cameroun, and else- where only from Shabunda, eastern Zaire (2°42'S, 2r20'E) (Hill, 1963: 81). On 5 November a single adult female was taken in the bottom shelf of a mist net placed along a cleared trail in the Intermediate Evergreen Forest near the Forest Guest House at the Kakamega Forest Station. This specimen from the Kakamega Forest extends the range of this species about 900 kilometers to the northeast and is the first record of occurrence for Kenya. For a description of the capture area, see Zimmerman (1972).
In size, the Kenyan female agrees well with a series of Cameroun specimens in Carnegie Museum and with the measurements given by Hill (1963:80).
Hipposideros cyclops (Temminck, 1853)
Phyllorhina cyclops Temminck, 1853. Esquisses Zool, sur la Cote de Guine, p. 75. Boutry River, Ghana.
Records (2).— Kenya: Coastal Prov. Kwale Dist., Shimba Hills Nature Reserve, Ma- kandara Picnic Site, 7 km S, 8 km W Kwale (4®15'S, 39®23'E) (2 CM).
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SCHLITTER ET AL. — NOTES ON KENYAN BaTS
299
Measurements.— measurements of a male and female, re- spectively, length of forearm, 66.8, 63.7; condylocanine length, 27.1, 27.2; zygomatic breadth, 14.8, 1 5.2; alveolar length of maxillary tooth- row, 8.4, 8.7; and greatest crown breadth of upper molar toothrows,
10.4, 10.6.
Remarks.— Hipposider os cyclop s is known currently from two lo- calities in western Kenya. Hayman (1 935:48) reported this species from the Yala River and later Harrison (1961:290) listed Kaimosi. The species is not known from Tanzania.
On 14 October, two adult individuals of H. cyclops were captured in a mist net set in the cleared Makandara Picnic Site in the Shimba Hills Nature Reserve. The net was set parallel and adjacent to the forest edge. These two specimens extend the range of this species to extreme southeastern Kenya. The geographic range of this species probably reaches this region of Kenya along a corridor of forest blocks extending from southwestern Tanzania to northeastern Kenya. This species should be captured in appropriate forests in this Tanzanian corridor after careful and concentrated mist netting.
Family Vespertilionidae Scotophilus nux Thomas, 1 904
Scotophilus Thomas, 1904. Ann. Mag. Nat. Hist., (7) 4:355. Gambage, Ghana.
Records (4).— Kenya: Western Prov., Kakamega Dist., Kakamega Forest Station, V/i km S, 12 km E Kakamega (2 CM, 1 MCZ, 1 NMK).
Measurements. SQlQcXtd external measurements of four females, length of forearm, 57.8, 57.6, 54.5, 55.3.
Remarks.— Scotophilus nux has been reported in Kenya only from 8 km SE Kakamega (Aggundey and Schlitter, 19^4:138). The eight specimens from Kenya referred to by Robbins (1983:23) are from this locality. Both localities refer to the Kakamega Forest Station. Four additional specimens of this dark-brown colored forest species of Sco- tophilus were netted in the forest adjacent to the station and in the cleared areas around the station complex on 5 and 7 November. These specimens further verify the occurrence of this species in the forests of western Kenya. Specimens of Scotophilus dinganii were not taken to- gether with S. nux at this locality, but the former species was taken in more heavily disturbed areas near the station complex and seems to be the most common species of Scotophilus taken in Kenya.
Kerivoula argentata argentata Tomes, 1861 Kerivoula argentata Tomes, 1861. Proc. ZooL Soc. London, 1861:32. Otjoro, Namibia.
Records (I).— Kenya: Coastal Prov., Kwale Dist., Shimba Hills Nature Reserve, 5
km S, 1 km W Kwale (4°13'S, 39°27'E) (1 CM).
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Measurements. — Selected measurements of an adult male, length of forearm, 35.9; condylobasal length, 13.9; and alveolar length of max- illary toothrow, 5.9.
Remarks.— Kerivoula argentata occurs primarily in southern Africa and ranges northward as far as Kenya. Specimens of Kerivoula are seldom captured and there is a paucity of records for this species in eastern Africa. It has been reported from Li wale, in southern Tanzania (Harrison, 1 958:95), an unspecified locality in Uganda (Kingdon, 1974: 304), and from Kibwezi, southeastern Kenya (Aggundey and Schlitter, 1984:139). A single adult male was taken on 12 October in a mist net set under the trees in the central clearing surrounding the bandas at the overnight facilities in Shimba Hills Nature Reserve. This specimen constitutes the second locality record of the species for Kenya and only the fourth for this region encompassing the northern part of the range of the species.
The Shimba Hills specimen exhibits the normal white-tipped orange- rufous dorsal coloration and the somewhat lighter colored ventral fur. Mensurally it fits within the size range of southern African examples of the species.
Family Molossidae
Tadarida (Chaerephon) mssata (J. Allen, 1917)
Chaerephon mssata J. Allen, 1917. Bull. Am. Mus. Nat. Hist., 37:458, Medje, Zaire.
Records (3). —Kenya: Rift Valley Prov., Naivasha Dist., Hell’s Gate Canyon, 20 km S, 14 km W Naivasha (0°54'S, 36°19'E) (1 CM, 1 MCZ, 1 NMK).
Measurements.— SeXQcXQd measurements of two males and one fe- male, respectively, length of forearm, 45.4, 46.2, 46.2; condylobasal length, 18,0, 18.0, 17.1; zygomatic breadth, 11.2, 11.9, 11.2; alveolar length of maxillary toothrow, 7.2, 7.1, 6.7; and greatest crown breadth of upper molar toothrows, 9.0, 8.7, 8.3.
Remarks.— ThQ taxonomic and distributional status of Tadarida russata was reviewed by Peterson (1971) and Fenton and Peterson (1972). This species is presently known from Ghana, Cameroun, and the type locality in northeastern Zaire (Fenton and Peterson, 1972:20). On 30 September, two adult males were captured in a mist net set across the floor of the northern end of Hell’s Gate Canyon. On 3 October a single adult female was taken in a net set in the same area. These three specimens comprise the first record of the species for Kenya and extend the range of the species more than 1000 kilometers to the southeast.
The Kenyan specimens of this species agree in color of pelage with three Cameroun examples in Carnegie Museum. However, mensurally they appear to be appreciably larger than these examples and the mea-
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SCHLITTER ET AL. — NOTES ON KENYAN BaTS
301
surements given in Fenton and Peterson (1972:21, 22). When more specimens from throughout the geographic range of this species become available, the Kenyan population may prove to be significantly larger and worthy of subspecific recognition.
Tadarida (Tadarida) fulminans (Thomas, 1903)
Nyctinomus fulminans Thomas, 1903. Ann. Mag. Nat. Hist, (7)12:501.
Fianarantsoa, eastern Betsileo, Madagascar.
Records (1).— Kenya: Rift Valley Prov., Nakuru Dist., Njoro River, 3 km S, 3 km W Nakuru (0°19'S, 36°03'E) (1 CM).
Measurements. SQlQCiQd measurements of an adult male, length of forearm, 56.7; condylobasal length, 21.2; zygomatic breadth, 13.4; al- veolar length of maxillary toothrow, 8.3; and greatest breadth of upper molar toothrow, 9.3.
Remarks. — The first record of occurrence for Tadarida fulminans in Kenya was a specimen from Nairobi reported by Harrison (1960:74). This species occurs primarily in southern Africa (Smithers, 1983) and reaches the northern extent of its range in Kenya. On 8 October a single adult male was captured in the gorge of the Njoro River southwest of Nakuru. The specimen was taken in a mist net set across shallow water and among trees on the steep bank of the watercourse.
Acknowledgments
A research permit (CAB 13/001/1 1C38/17) allowing us to do field studies in Kenya was received from the Office of the President, We thank Mrs, C. A. Mwango and Miss L, A, Gwiyo and their staff for their efforts and courtesy in processing this permit on our behalf and the Secretary, National Council for Science and Technology, and his colleagues for approving our research project.
Many individuals in the Ministry of Tourism and Wildlife supported the research project in many ways. We are especially grateful to Mr. D. M. Mbuvi, Assistant Director (Research), Wildlife Conservation and Management Department and his staff for support, Mr. J. P. Oriero, Assistant Director (Administration), for issuing permit No. WCMD/ 7/9/ Vol. VI, and the numerous Wardens and their staff in the Parks and Reserves in Kenya who made our stay comfortable and work easier. Mr. F. N. Pertet, Principal, Wildlife Training College, Naivasha, and his staff, especially Mr. Okumu Mbanda and Mr. Osbourne Mivelo, were gracious hosts and gave invaluable assistance.
We thank Mr. John S. Karmali and the staff at Nakuru Wildlife Trust House, Lake Nakuru National Park and Dr. Stephen G. Njuguna and his staff at the Moana Marine Station, Diani Beach, for logistical assistance.
Finally we are especially indebted to Dr. Richard Leakey and his staff of the National Museums of Kenya, including Mr. James N. Maikweki, Curator in Charge, Kisumu Museum, for their overwhelming support. We are truly appreciative of the assistance of Caroline Plazek and Paul Gathinji who assisted with the field work.
Financial assistance for field work in Kenya was received from the M. Graham Netting Research Fund, Cordelia Scaife May Charitable Trust, Carnegie Museum of Natural History; from Barbour and Richmond funds. Museum of Comparative Zoology; from NIH grant AIO 4242 to Dr. Robert Traub; and the Rea Postdoctoral Fellowship program at Carnegie Museum of Natural History.
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Literature Cited
Aggundey, L R., and D. A. Schlitter. 1984. Annotated checklist of the mammals of Kenya. I. Chiroptera. Ann. Carnegie Mus., 53:1 19-161.
Fenton, M. B., and R. L. Peterson. 1972. Further notes on Tadarida aloysiisabaudiae and Tadarida russata (Chiroptera: Molossidae— Africa). Canadian J. ZooL, 50:19-24.
Harrison, D. L. 1958. Two bats (Microchiroptera) new to the fauna of Tanganyika Territory. Durban Mus. Novit., 5:95-98.
. 1960. Notes on some Central and East African bats. 3. The free-tailed bat
Tadarida Thomas, 1903, in Kenya Colony. Durban Mus. Novit., 6:74-78.
. 1961. A checklist of the bats (Chiroptera) of Kenya Colony. J. East African
Nat. Hist. Soc., 23:286-295 (for 1960).
Hayman, R. W. 1935. A note on Hipposideros cyclops Temminck and its synonym Hipposideros langi Allen. Ann. Mag. Nat. Hist., (10)15:47-50.
Hill, J. E. 1963. A revision of the genus Hipposideros. Bull. British Mus. (Nat. Hist.), Zool., 2(1):1-129.
Kingdon, j, 1974. East African mammals. An atlas of evolution in Africa. Volume II, Part A (Insectivores and Bats). Academic Press, London, xii + 392 pp.
Peterson, R. L. 1971. The African molossid bat Tadarida russata. Canadian J. Zool., 49:297-301.
Robbins, C. 1983. A new high forest species in the African bat genus Scotophilus (Vespertilionidae). Ann. Mus. Roy. Afr. Centr., Sc. Zool., 237:19-24.
Smithers, R. H. N. 1983. The mammals of the southern African subregion. University of Pretoria, Pretoria, xxii + 734 pp.
Zimmerman, D. A. 1972. The avifauna of the Kakamega Forest, western Kenya, in- cluding a bird population study. Bull. American Mus. Nat. Hist., 149:257-339.
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Back issues of many Annals of Carnegie Museum articles are available, and a few early complete volumes and parts are listed at half price. Orders and inquiries should be addressed to: Publications Secretary, Carnegie Museum, 4400 Forbes Avenue, Pittsburgh, Pa. 15213.
o/ CARNEGIE MUSEUM
CARNEGIE MUSEUM OF NATURAL HISTORY 4400 FORBES AVENUE ® PITTSBURGH, PENNSYLVANIA 15213 VOLUME 55 7 NOVEMBER 1986 ARTICLE 13
RESULTS OF THE ALCOA FOUNDATION-SURINAME EXPEDITIONS. XL BATS OF THE GENUS MICRONYCTERIS (MAMMALIA: CHIROPTERA)
IN SURINAME
Hugh H. Genoways
Research Associate, Section of Mammals Stephen L. Williams
Collection Manager, Section of Mammals
Abstract
Of the 10 recognized species in the genus Micronycteris, seven species {brachyotis, daviesi, hirsuta, megalotis, minuta, nicefori, and sylvestris) have been reported to occur in Suriname. Micronycteris daviesi is easily distinguished from the other species by its large size (forearm over 50 mm) and massive dentition. The next largest species (forearm over 43 mm)--M hirsuta— is distinguished by having the upper inner incisors separated at the base but in contact at the tip and having lower incisors that are high and wedged between canines so that the canines are in contact behind the incisors.
Micronycteris sylvestris has dorsal pelage that is tricolored. The upper incisors of this species are similar in length to the canines and the first upper premolar possesses accessory cusps.
The other four species form two species pairs. Micronycteris megalotis and minuta are the smallest members of the genus in Suriname. They can be distinguished from each other by the more deeply notched interauricular band in minuta and by the first upper premolar being smaller than the second premolar in minuta but of about equal size in megalotis.
Micronycteris brachyotis, which was not encountered during our work in Suriname, has short ears (less than 1 6 mm from notch) and lacks the faint gray line usually present on the lower back of specimens of M. nicefori. These species are also distinguished by
Submitted 24 April 1986.
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304
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Fig. 1. — Stylistic representation of variation in the interauricular band of members of the genus Micronycteris. Upper, interauricular band complete (exemplified by M. hir- suta). Middle, interauricular band deeply notched in middle {M. minuta). Lower, inter- auricular band absent (M. nicefori). Not to scale.
their upper incisors, which are chisel-shaped and in line with the canines in brachyotis, but project forward and, therefore, are out of line with the canines in nicefori.
Introduction
The genus Micronycteris is a member of the subfamily Phyllostom- inae of the New World leaf-nosed bat family Phyllostomidae. Micro- nycteris is currently recognized as containing 10 small- to medium-
1986
Genoways and Williams —Suriname Micronycteris
305
sized species of bats. The genus was reviewed by Andersen (1906) and Sanborn (1949).
The genus is divided into six subgenera based primarily upon the work of Sanborn (1949). We suggest that this level of classification be carefully examined in the future because the genus seems oversplit at the subgeneric level. Andersen (1906) recognized seven species divided into two genera {Micronycteris and Gly phony cteris), whereas Sanborn (1949) recognized 10 species in one genus. Subsequent to Sanborn’s work, one species (platyceps) was placed as a junior synonym of brachy- Otis (Goodwin and Greenhall, 1961) and another species, daviesi, was described in a separate genus {Barticonycteris; Hill, 1 964), but was later placed into Micronycteris (Koopman and Cockrum, 1967).
During our field studies in Suriname, six species of the genus Mi- cronycteris {daviesi, hirsuta, megalotis, minuta, nicefori, and sylvestris) were captured. Husson (1978) reported a seventh species {brachyotis) from the country. This means that only three of the currently recognized species of Micronycteris {behni, pusilla, and schmidtorum) have not been taken in Suriname. With this large number of species from this complex genus present in such a small geographic area, we have taken the opportunity to gain a better insight into the morphological and ecological relationships of species within the genus.
Methods and Materials
Specimens were taken with mist nets and preserved as skins and skulls or in fluid. Field weights were taken with Pesola scales, accurate to 1 g. Measurements of forearm and cranial dimensions were taken with dial calipers accurate to 0. 1 mm. Only adult specimens (phalangeal epiphyses completely fused) were measured in this study. Mea- surements were taken as described by Genoways and Williams (1984). Reproductive condition of the skin and skull specimens was determined by gross dissection in the field, whereas fluid preserved specimens were dissected in the laboratory. Specimens listed in each account were deposited in the Section of Mammals, Carnegie Museum of Natural History.
Acknowledgments
We gratefully acknowledge the logistical support, assistance in acquiring permits, and field assistance of Dr, loop Schulz, Mr. Henry Reichart, and Mr. Ferdinand Baal. Mr. Kris Mohadin, Ms. Muriel Hand, and other staff members of STINASU contributed to the success of field work in Suriname. We are particularly thankful to Mr. Leo Roberts for accompanying and assisting us with most of the field work. Ms. Marga Werkhoven and Mr. 1. Douglas provided housing and lab facilities in Paramaribo. Other individuals who helped collect specimens used in this study include Ms. Jane Casne, Mr. Michael Arnold, Dr. Rodney Honeycutt, Mr. Ben Koop, Ms. Paisley Seyfarth, Mr. Murray de la Fuente, Dr. Carleton Phillips, Dr. Robert Baker, and Mr. Keith Studholme.
Financial support for field work in Suriname was received from the Alcoa Foundation (Charles L. Griswold, President) and the M. Graham Netting Research Fund established by a grant from the Cordelia S. May Charitable Trust.
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vou 55
Fig, 2. — Dental characteristics of members of the genus Micronycteris. A, lower incisors bifid (exemplified by M. megalotis); B, lower incisors trifid (exemplified by M. sylvestris). C, upper and lower incisors of M. hirsuta, showing long, narrow lower incisors with unexpanded crowns and awl-shaped upper incisors; D, upper and lower incisors of M. megalot is showing short lower incisors with expanded crowns. E, upper premolars (P3,
1986
Genoways and Williams —-Suriname Micronycteris
307
Key to the Species of Micronycteris in Suriname (see also Medellin et al., 1985)
1 . Interauricular band present (possibly notched or not as broad
in the middle (Fig. lA, B); lower incisors bifid (Fig. 2 A) ..... 2
1'. Interauricular band not present (Fig. 1C); lower incisors trifid
(Fig. 2B) 4
2. Forearm less than 40; greatest length of skull less than 22;
lower incisors with expanded crowns (Fig. 2D) 3
2'. Forearm greater than 40; skull more than 22; upper incisors awl-shaped; lower incisors long, narrow, and lacking expanded
crown (Fig. 2C); 2n = 30, FN = 32 ....... .Micronycteris hirsuta
3. Calcar longer than foot (claws included); length of interfemoral membrane more than twice the length of tail; band of skin between ears with shallow notch in middle; upper premolars (P3, P4) about the same height (Fig. 2E); 2« = 40, FN =68
Micronycteris megalotis
3 ' . Calcar shorter than foot (claws included); length of interfemoral membrane less than twice the length of tail; band of skin be- tween ears deeply notched in middle (Fig. IB); first upper premolar (P3) distinctly shorter than second upper premolar (P4) (Fig. 2F); 2n = 28, FN = 52 Micronycteris minuta
4. First upper incisors similar to canines in length; first upper premolar (P3) having accessory cusps on lingual and posterior
margins 5
4'. First upper incisors distinctly shorter and narrower than ca- nines; first upper premolar (P3) lacking accessory cusps, only
the main cusps present 6
5. Forearm less than 50; greatest length of skull less than 25; dorsal hair tricolored; two pairs of upper incisors; 2n = 22,
FN = (40) Micronycteris sylvestris
5'. Forearm greater than 50; greatest length of skull more than 25; dorsal hair brownish throughout; sagittal crest straight; one pair of upper incisors; 2n = 28, FN = 52 . . Micronycteris daviesi 6. Length of ear (to notch) less than 16; calcar about the same length as foot; first pair of upper incisors chisel-shaped (Fig.
2G) and in line with canines; second pair of upper incisors
P4) of M. megalotis (anterior is to the right), note that premolars of about equal size; F, upper premolars of M. minuta (anterior is to the right), note that P3 is distinctly shorter than P4. G, chisel-shaped upper incisors of M. brachyotis; H, upper incisors of M. nicefori which are nearly as broad as they are tall.
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bifid with elongated inner cusp; 2n = 32, FN = 60 .........
Micronycteris brachyotis
6'. Length of ear (to notch) greater than 16; calcar shorter than length of foot; faint gray line often present on lower back; first pair of upper incisors not chisel-shaped (Fig. 2H); upper in- cisors projected forward and out of line with canines; In = 28,
FN = 52 Micronycteris nicefori
Generic Account Micronycteris Gvdiy , 1866
1866. Micronycteris Gray, Proc. Zool. Soc. London, p. 113, May.
1856. Schizostoma Gervais, Mammiferes in Castelnau Exped. dans les parties centrales de I’Amer. du Sud . . . pt. 7, p. 38. Type species, Schizostoma minuta Gervais. Preoccupied by Schizostoma Bronn, 1835, a genus of Mollusca.
1896. Glyphonycteris Thomas, Ann. Mag. Nat. Hist., ser. 6, 18:302, October. Type species, Glyphonycteris sylvestris Thomas.
1907. Xenoctenes Miller, Bull. U.S. Nat. Mus., 57:124, 29 June. Type species, Schizosto- ma hirsutum Peters.
1964. Barticonycteris Hill, Mammalia, 28:556, December. Type species, Barticonycteris daviesi Hill.
Type species.— Phyllophora megalotis Ovdcy , 1842.
Diagnosis.— K genus of small- to medium-sized bats with a well- developed noseleaf and a tail extending only to the middle of the interfemoral membrane. In the subfamily Phyllostominae, the dental formula of i 2(1 )/2, c 1/1, p 2/3, m 3/3 is shared only with the genera Macrotus and Vampyrum. The one species with only one pair of upper incisors is M. daviesi. Rostrum not as long as braincase; auditory bullae small; middle lower premolar approximately same size as last lower premolar.
Micronycteris {Glyphonycteris) daviesi (Hill, 1964)
Specimen examined (1).— Saramacca: Raleigh Falls, 1.
The species M. daviesi is easily distinguished from other members of the genus Micronycteris in Suriname by its large size (Table 1; Figs. 3A, 4A) and massive dentition. This species was originally described as the sole representative of the genus Barticonycteris by Hill (1964). Shortly thereafter, Koopman and Cockrum (1967) treated Bartico- nycteris as a synonym of Micronycteris. Most recent authors have fol- lowed this arrangement (see for example Jones and Carter, 1976), al- though LaVal (1977) and Hall (1981) are exceptions. Koopman (1978) treated Barticonycteris as a subgenus of Micronycteris, citing as his reasons that the characteristics of Barticonycteris “are simply those of M. {Glyphonycteris), the subgenus including sylvestris and behni carried one step further.” Hill (1964) had earlier recognized that the closest
1986
Genoways and Williams —Suriname Micronycteris
309
relatives of Barticonycteris were members of Gly phony cteris. We agree with these assessments of the relationship of daviesi based upon our own studies, but we believe that the relationships of the taxon are represented best by placing it as a member of the subgenus Glyphonyc- teris.
Only one specimen of this rare species was taken during our work in Suriname (Fig. 5). It was an adult male taken on an island in the Coppemame River that serves as the headquarters of the Raleigh Falls Nature Reserve. The bat was netted along a trail on the western side of the island, about 200 m northeast of the park headquarters and about 50 m from the river. Vegetation in the area consisted of near- mature lowland rainforest. Our specimen, weighing 1 8 and with testes measuring 3, was captured on 24 August at about 2000 hours following a short rainstorm. Sixteen other species of bats were taken in this area (Table 2).
Our specimen was found to have a In = 28 and FN = 52. The X-chromosome was submetacentric, whereas the Y-chromosome was acrocentric (Honeycutt et al., 1980).
Micronycteris (Glyphonycteris) sylvestris (Thomas, 1896)
Specimens examined (14). — Brokopondo: Brownsberg Nature Park, 8 km S, 2 km W Brownsweg, 14.
Our specimens were the first of this species reported from Suriname (Williams and Genoways, 1980). This taxon can be recognized exter- nally by having tricolored dorsal hair and ears that are about as broad as they are high. Cranially this species resembles M. daviesi with upper incisors about the same length as the canines (Figs. 3, 4). Currently M. sylvestris is considered to be monotypic (Jones and Carter, 1976), al- though not enough specimens have been available for a proper analysis of infraspecific variation.
Our 14 specimens were collected from a hollow tree in a mature tropical hardwood forest on the Brownsberg highlands (Fig. 5). The opening to the hollow was located about 3 m above the ground. Eight of the specimens were taken on 24 September and the other six on the following day. Four males weighed 6, 6, 7, and 7; each had testes that measured 3. Eight females had weights ranging from 7.5 to 11 with a mean of 9.3. None of these females evinced gross reproductive activity. Only 10 other species of bats were captured in nets set along trails near the hollow tree (Table 2).
The specimens of M. sylvestris from Suriname had a diploid number of 22 and a probable fundamental number of 36. The fundamental number could not be determined with certainty because only females were available for chromosomal analysis. It was supposed that the X-chromosome was biarmed (Honeycutt et al., 1980).
310
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1cm
VOL, 55
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Annals of Carnegie Museum
Fig. 3.— Dorsal view of the crania of the seven species of Mkronycteris occurring in Suriname. A, M. daviesi (CM 63573); B, M. syivestris (CM 63597); C, M. brachyotis (from Trinidad); D, M. megalotis (CM 68390); E, M. minuta (CM 63584); F, M nicefori (CM 76771); G, M. hirsuta (CM 68388).
1986
Genoways and Williams "“Suriname Micronycteris
313
Micronycteris (Lampronycteris) brachyotis (Dobson, 1879)
Previous Brokopondo: Gros (about 100 km S Paramaribo on railroad from
Paramaribo to the interior) [5°06'N, 55°15'W] (Husson, 1978).
Husson (1978) first reported this species from Suriname based upon six males from Gros (Fig. 5). The specimens were taken from an old goldmine in a savannah area. We did not encounter this species during our work in Suriname.
Most of the characteristics of the specimens listed by Husson —fore- arm 40.2 to 42.9, no interauricular band, and second phalanx of third digit much longer than first— seem to match M. brachyotis closely; however, the Suriname specimens had the fourth metacarpal the short- est, whereas in brachyotis the fifth metacarpal is the shortest. The exact meaning of this difference must await further examination of these specimens.
Goodwin and Greenhall (1961) were the first to recognize that M. platyceps, described by Sanborn in 1949, was a junior synonym of the long described, but poorly known, M. brachyotis.
Micronycteris {Micronycteris) megalotis megalotis (Gray, 1842)
Specimens examined (12).— Brokopondo: 1 km N Rudi Kappelvliegveld, 1; P/2 km W Rudi Kappelvliegveld, 1; 3 km SW Rudi Kappelvliegveld, 2; Brownsberg Nature Park, 3 km S, 20 km W Afobakka, 1 . Commewune: Nieuwe Grond Plantation, 1 . Maro- wune: Oelemarie, 1; Perica, 2. Nickerie: Kayserberg airstrip, 2. Para: Zanderij, 1.
Previous records (Husson, 1978).— Suriname: Plantation Kwatta, near Rijweg; Para- maribo. No specific district or locality.
Among Surinamese representatives of the genus Micronycteris, M. megalotis is distinguished by the presence of an interauricular band which is only slightly notched, a broad interfemoral membrane, upper premolars (P3, P4) about the same size (Fig. 2E), and bifid lower incisors (Fig. 2A). The species is polytypic with the nominate subspecies oc- curring in Suriname and surrounding areas (Jones and Carter, 1976).
M. megalotis has been reported from localities in Suriname previ- ously and these together with our records indicate that the species may be expected in most forested situations in the country (Fig. 6). Many of the capture sites were described by the collectors as being in mature tropical forest or lowland tropical rainforest. However, at the Nieuwe Grond Plantation a specimen was taken in a net set along an orchard path that was bordered on one side by a canal and on the other by alternating rows of citrus trees and secondary tropical vegetation. At Perica, the specimens were netted on the edge of secondary growth forest. M. megalotis was taken at more than half of its nine capture sites with six species (Table 2)— Saccopteryx bilineata, S. leptura, To- natia bidens, Carollia perspicillata, Rhinophylla pumilio, and the larger
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VOL. 55
1986
Genoways and Williams —Suriname Micronycteris
315
58 57 56 55 54
Fig. 5,— Map of the geographic distribution of four species of Micronycteris in Suriname. Closed circle, M. daviesi; open circle, M. sylvestris; closed triangle, M. brachyotis; open triangle, M. hirsuta.
species of Artibeus. It was also taken together with two other species of the gQmis—minuta and although only at one and two sites,
respectively.
Our 12 specimens consist of 10 males and two females. Testes mea- surements for the males are as follows (date of capture in parentheses):
Fig. 4.— Lateral view of the crania of the seven species of Micronycteris occurring in Suriname. A, M. daviesi (CM 63573); B, M. sylvestris (CM 63597); C, M. brachyotis (from Trinidad); D, M. megalotis (CM 68380); E, M. minuta (CM 63584); F, M. nicefori (CM 76771); G, M. hirsuta (CM 68388).
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58 57 56 55 54
Fig. 6.— Map of the geographic distribution of Micronycteris megalotis in Suriname. Closed circles, specimens examined; open circles, previous records.
3, 3 (4 May); 3 (7 July); 2 (13 September); 2 (30 September); 2 (1 October); 2 (3 October); 3, 4.5 (24 October); 2.5 (23 November). A female taken on 1 October evinced no reproductive activity, whereas no data are available for the other female. Seven of the males had an average weight of 5.7 (range, 5—7) and the one female for which data are available weighed 6,
The karyotype of a male from Suriname had a 2« = 40 and FN == 68. The X-chromosome was subtelocentric and the Y-chromosome was acrocentric (Honeycutt et al., 1980).
Micronycteris (Micronycteris) minuta (Gervais, 1856)
Specimens examined (15).-— Brokopondo: 1 km N Rudi Kappelvliegveld, 1; Browns- berg Nature Park, 8 km S, 2 km W Brownsweg, 2. Commewune: Nieuwe Grond Plan-
1986
Genoways and Williams —Suriname Micronycteris
317
58 57 56 55 54
Fig. 7.— Map of the geographic distribution of Micronycteris minuta in Suriname.
tation, 1. Marowune: 3 km SW Albina, 1; 10 km N, 24 km W Moengo, 1. Nickerie: Avanavero, 1; Sipaliwini airstrip, 4. Saramacca: Voltzberg, 3. Suriname; Powaka, 1.
Prior to our work in Suriname (Genoways and Williams, 1979; Wil- liams and Genoways, 1 980) M. minuta was unknown from the country; however, we took the species in all major regions of the country (Fig, 7). In Suriname, M. minuta would most likely be confused with M. megalotis; however, minuta can be distinguished by the deeply notched interauricular band (Fig. 1) and a first upper premolar (P3) that is distinctly smaller than the second premolar (P4) (Fig. 2F). M. minuta has some individuals smaller than any individuals of M. megalotis, but the two species cannot be separated consistently on size alone (Table 1). M. minuta currently is considered to be monotypic (Jones and Carter, 1976). A comparison of our material with specimens from
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Trinidad and Brazil deposited in the American Museum of Natural History revealed no notable differences; however, there may be slight karyotypic differences between specimens from Trinidad and Suriname (see discussion below).
Micronycteris minuta usually was netted in association with lowland rainforest. However, at several places such as 1 km N Rudi Kappel- vliegveld, 10 km N, 24 km W Moengo, and Powaka, this species was taken in nets set in open areas along or near the forest. At the Nieuwe Grond Plantation, M. minuta was captured in a series of nets set through the lawns and gardens surrounding the plantation buildings. Nine species of bats were collected in association with M. minuta at more than half of the nine capture sites (Table 2). An interesting association is that between M. nicefori and M. minuta, which were taken together at five capture sites. M. minuta was also captured together with two other members of the genus (megalot is and sylvestris) at single localities.
Our 15 specimens consist of six males and nine females. Testes measurements for males were as follows (date of capture in paren- theses): 3 (12 May); 2 (24 September); 2 (28 September); 3 (23 October); 2 (16 November). A female taken on 30 July was carrying a 13 mm fetus, and another was lactating on 24 September. Females netted on the following dates evinced no gross reproductive activity: 26 May; 6 August; 28 August; 12 September; 16 November (3 individuals). Four of the males had weights of 5, 5, 6, and 6 and three females weighed 5, 5, and 8.
The diploid number for this species in Suriname was 28 and the fundamental number was 52 (Baker et al., 1981). This karyotype dif- fered slightly from that reported for the species from Trinidad (Baker, 1979). In the Suriname specimen, the smallest pair of autosomes was biarmed, whereas in the material from Trinidad this pair was acro- centric.
Micronycteris (Trinycteris) nicefori Sanborn, 1 949
Specimens examined (31). — Brokopondo: 1 km N Rudi Kappelvliegveld, 1. Comme- wune: Nieuwe Grond Plantation, 1. Marowune: 3 km SW Albina, 1; 10 km N, 24 km W Moengo, 1; Perica, 2. Nickerie: Avanavero, 3; Kabalebo, 20. Saramacca: Tafelberg, SE side of Arrowhead Basin (3®54'N, 56°10'W), 600 m, 1. Suriname: Powaka, 1.
This species had not been reported in Suriname before our work (Genoways and Williams, 1979; Williams and Genoways, 1980), but we found it to be widespread in the country (Fig. 8). Micronycteris nicefori is a medium-sized member of the genus (Table 1) that is most likely to be confused with M. brachyotis. M. nicefori can be distin- guished by its upper incisors, which are shorter and narrower than the canines and project forward out of line with the canines (Fig. 2H), long
1986
Genoways and Williams —Suriname Micronycteris
319
58 57 56 55 54
Fig. 8.— Map of the geographic distribution of Micronycteris nicefori in Suriname,
ears, and a faint gray line often present on the lower back. We compared our specimens with material from Trinidad and Colombia deposited in the American Museum of Natural History. We could detect no consistent differences in size or morphology, which supports the idea that this species is monotypic (Jones and Carter, 1976).
The large sample from Kabalebo is composed of all males. These individuals were captured in nets set along a newly cut trail, which passed through the moderate undergrowth of a secondary forest along a road, which eventually led into the larger trees of a mature rainforest. Elsewhere the species usually was collected in either secondary or pri- mary lowland rainforest. The exceptions to this were in the highlands of Tafelberg where the typical vegetation was lower montane forest
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and at Nieuwe Grond Plantation where a single specimen was taken in nets set over the lawns and gardens surrounding the plantation headquarters. M. nicefori shows a high correlation with the distribu- tions of only seven other species; five of these— Lonchophylla thomasi, Carollia perspicillata, Rhinophylla pumilio, Sturnim lilium, and Arti- beus (large species)— were common, widespread species (Table 2). Glos- sophaga soricina was not a particularly common species in Suriname. The most interesting distributional correlation was with M. minuta, which was taken at five of the nine localities where M. nicefori was captured.
Length of testes of the 20 males from Kabalebo taken on 28 May averaged 3.5 (range, 3-4). Other males had the following testes lengths (date of capture in parentheses): 3, 4, 5 (26 May); 5 (6 August); 4 (23 October); 3 (24 October). None of the three females for which data are available evinced reproductive activity (netted on 30 July, 12 Septem- ber, and 30 September). Two of the males weighed 7.3 and 8 and three females weighed 7, 8, and 8.5.
Surinamese specimens of M. nicefori had a 2/t = 28 and FTSF = 52. The X-chromosome was submetacentric and the Y-chromosome was acrocentric (Honeycutt et al., 1980; Baker et al., 1981).
Micronycteris (Xenoctenes) hirsuta (Peters, 1869)
Specimens examined (3).— Nickerie: Kabalebo, 1. Para: Zanderij, 2.
Micronycteris hirsuta is the sole member of the subgenus Xenoctenes (Figs. 3, 4). Miller (1907) originally gave this taxon generic status, but Sanborn (1949) reduced it to subgeneric level. The species M. hirsuta can be distinguished from other members of the genus in Suriname by the following characteristics: ears connected across the forehead by a low unnotched band (Fig. 1); upper inner incisors separated at base, but in contact near tip (Fig. 2C); upper outer incisors small; lower incisors high and wedged tightly between canines (Fig. 2C); lower ca- nines in contact, or nearly so, behind incisors; and lower incisors bifid (Fig. 2C). The species is considered to be monotypic (Jones and Carter, 1976).
Our specimens were the first members of the species (Fig. 5) to be reported from Suriname (Genoways et al., 1981). The specimens from Zanderij, a reproductively inactive adult female and an immature fe- male, were netted on 18 May. The specimen from Kabalebo was a reproductively inactive adult female. The area at Kabalebo was covered by mixed primary and secondary lowland rainforest, whereas the vi- cinity of Zanderij was secondary lowland forest associated with a rub- ber plantation. M. hirsuta was taken in association with 24 other species of bats (Table 2); however, only four species— Phyllostomus elongatus,
1986
Genoways and Williams “Suriname Micronycteris
321
P. hastatus, Carollia perspicillata, and large Artibeus species— were captured at both localities. All of these are common, widespread species in Suriname.
The specimens from Suriname had a 2« = 30 and FN = 32 (Baker et al,, 1981). This karyotype appears to be identical to the one found in Middle American populations of M. hirsuta but differs from that found on Trinidad where 2n = 28 (Baker et al., 1973; Baker, 1979).
Discussion
Seven of the 1 0 recognized species of Micronycteris are known from the small country of Suriname. As a general rule, specimens were taken in association with forested habitats, particularly mature lowland rain- forest. However, there was a relatively low correlation between the occurrence of any one species of Micronycteris and any of the other species of the genus (Table 2). Only M. minuta and M. nicefori were taken together at more than half of their collecting sites. There are two possible explanations for this fact. First, there can be, and probably are, subtle differences in the microhabitat required by each of the species within the forest. The other factor affecting this correlation may be that the distribution of each species may be clumped around available roost- ing sites. We saw this phenomenon in at least two places in Suriname. At Brownsberg Nature Park, specimens of M. sylvestris were taken only from a hollow tree. At Kabalebo, a large number of male M. nicefori was captured only in one set of nets. If the occurrence of species of Micronycteris is clumped around available roost sites and individuals have relatively small home ranges, then there definitely should be a reduction in the places that species co-occur.
The seven species of Micronycteris from Suriname form a gradient in size starting with the small M. minuta and M. megalotis and pro- gressing through the large M. daviesi at the opposite end of the scale. The small species really form a species pair based upon size, although M. minuta probably averages slightly smaller than M. megalotis for most characters. Between these extremes fall (beginning with the small- est) M. nicefori, M. sylvestris, and M. brachyotis, and finally M. hirsuta. Micronycteris sylvestris and M. brachyotis cannot be distinguished on size alone but there are numerous other useful characters to separate them. Size is not the only character needed to separate several of the species of Micronycteris in Suriname, but it is useful in narrowing the number of comparisons that need to be made.
We stated earlier that the genus Micronycteris seemed to be oversplit at the subgeneric level. The current arrangement was proposed by Sanborn (1949) based primarily upon characters of the wings and ears. Deviations from this arrangement have been suggested by Arnold et
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Table 2.— Species of bats taken in association with specimens o/Micronycteris in Suri- name. The numbers in each column represent numbers of localities.
|
Species of bats |
M. daviesi |
M. hirsuta |
M. megalotis |
M. minuta |
M. nicefori |
M. sylvestris |
|
No. of localities for which data are available |
1 |
2 |
9 |
9 |
9 |
1 |
|
Saccopteryx bilineata |
0 |
1 |
5 |
1 |
1 |
0 |
|
Saccopteryx canescens |
0 |
0 |
2 |
0 |
0 |
0 |
|
Saccopteryx leptura |
1 |
1 |
5 |
0 |
1 |
0 |
|
Cormura brevirostris |
0 |
1 |
0 |
1 |
1 |
0 |
|
Pteronotus parnellii |
1 |
1 |
3 |
4 |
3 |
1 |
|
Noctilio leporinus |
0 |
0 |
0 |
1 |
1 |
0 |
|
Chrotopterus auritus |
1 |
0 |
2 |
0 |
0 |
0 |
|
Micronycteris megalotis |
0 |
0 |
— |
1 |
2 |
0 |
|
Micronycteris minuta |
0 |
0 |
1 |
— |
5 |
1 |
|
Micronycteris nicefori |
0 |
0 |
2 |
5 |
~ |
0 |
|
Micronycteris sylvestris |
0 |
0 |
0 |
1 |
0 |
-= |
|
Tonatia bidens |
0 |
0 |
5 |
1 |
2 |
0 |
|
Tonatia brasiliense |
0 |
0 |
1 |
1 |
0 |
0 |
|
Tonatia carrikeri |
0 |
1 |
0 |
2 |
0 |
0 |
|
Tonatia schulzi |
0 |
0 |
2 |
0 |
1 |
0 |
|
Tonatia silvicola |
0 |
1 |
1 |
3 |
2 |
1 |
|
Mimon crenulatum |
0 |
0 |
1 |
1 |
1 |
0 |
|
Lonchorhina aurita |
0 |
0 |
0 |
1 |
1 |
0 |
|
Phyllostomus discolor |
1 |
1 |
2 |
1 |
0 |
1 |
|
Phyllostomus elongatus |
1 |
2 |
4 |
6 |
3 |
1 |
|
Phyllostomus hastatus |
1 |
2 |
3 |
5 |
3 |
0 |
|
Phyllostomus latifolius |
0 |
0 |
1 |
2 |
0 |
1 |
|
Trachops cirrhosus |
1 |
1 |
2 |
0 |
0 |
0 |
|
Anoura caudifer |
0 |
0 |
3 |
0 |
2 |
0 |
|
Lonchophylla thomasi |
1 |
1 |
4 |
7 |
6 |
1 |
|
Glossophaga soricina |
0 |
0 |
2 |
4 |
5 |
0 |
|
Carollia brevicauda |
1 |
0 |
0 |
1 |
0 |
0 |
|
Carollia perspicillata |
1 |
2 |
7 |
5 |
6 |
0 |
|
Rhinophylla pumilio |
1 |
1 |
5 |
6 |
5 |
1 |
|
Ametrida centurio |
0 |
0 |
0 |
1 |
0 |
0 |
|
Sturnira lilium |
1 |
1 |
4 |
6 |
6 |
0 |
|
Sturnira tildae |
0 |
0 |
1 |
2 |
2 |
1 |
|
Artibeus cinereus |
1 |
0 |
1 |
5 |
4 |
0 |
|
Artibeus concolor |
0 |
0 |
1 |
1 |
1 |
0 |
|
Artibeus (large species) |
1 |
2 |
7 |
9 |
8 |
1 |
|
Uroderma bilobatum |
1 |
1 |
1 |
3 |
3 |
0 |
|
Chiroderma trinitatum |
0 |
1 |
0 |
1 |
1 |
0 |
|
Vampyressa bidens |
0 |
0 |
2 |
0 |
0 |
0 |
|
Vampyressa brocki |
0 |
0 |
0 |
1 |
1 |
0 |
|
Vampyrops aurarius |
0 |
0 |
0 |
0 |
1 |
0 |
|
Vampyrops brachycephalus |
0 |
0 |
2 |
1 |
2 |
0 |
|
Vampyrops helleri |
1 |
1 |
2 |
4 |
3 |
0 |
|
Vampyrodes caraccioli |
0 |
0 |
0 |
1 |
1 |
0 |
|
Mesophylla macconnelli |
0 |
1 |
1 |
2 |
3 |
0 |
|
Desmodus rotundus |
0 |
1 |
0 |
0 |
0 |
1986
Genoways and Williams —Suriname Micronycteris
323
Table 2. — Continued.
|
Species of bats |
M. daviesi |
M. hirsuta |
M. megalotis |
M. minuta |
M. nicefori |
M. sylvestris |
|
Thyroptera tricolor |
0 |
0 |
1 |
2 |
1 |
0 |
|
Natalus tumidirostris |
0 |
0 |
0 |
1 |
0 |
0 |
|
Myotis nigricans |
0 |
1 |
1 |
3 |
3 |
0 |
|
Eptesicus brasiliensis |
0 |
1 |
4 |
1 |
3 |
0 |
|
Molossus ater |
0 |
1 |
0 |
0 |
0 |
0 |
|
Molossus molossus |
0 |
1 |
1 |
1 |
1 |
0 |
|
Total species (51) |
16 |
24 |
36 |
41 |
35 |
10 |
al. (1983) based on electrophoretic studies. For phenotypic criteria a useful classification might be devised by using characters of the ears and the teeth at the front part of the dental arcade (canines and incisors). The genus can be split into two major groups— in the first the ears are connected by an interauricular band and the lower incisors are bifid, and in the second the ears are not connected by an interauricular band and the lower incisors are trifid. We suggest that future investigation of subgeneric classifications of this genus examine these groupings and any subgroupings within them.
Literature Cited
Andersen, K. 1906. On the bats of the genera Micronycteris and Gly phony cteris. Ann. Mag. Nat. Hist., ser. 7, 18:50~-63.
Arnold, M. L., R. J. Baker, and R. L. Honeycutt. 1983. Genic differentiation and phylogenetic relationships within two New World bat genera. Biochem. Sys. Ecol., 11:295=303.
Baker, R. J. 1979. Karyology. Pp. 107=155, in Biology of bats of the New World family Phyllostomatidae, Part III (R. J. Baker, J. K. Jones, Jr., D. C. Carter, eds.). Spec. Publ. Mus., Texas Tech Univ., 16:1=441.
Baker, R. J., H. H. Genoways, W. J. Bleier, and J. W. Warner. 1973. Cytotypes and morphometries of two phyllostomatid bats, Micronycteris hirsuta and Vam- pyressa pusilla. Occas. Papers Mus., Texas Tech Univ., 17:1-10.
Baker, R. J., H. H. Genoways, and P. A. Seyfarth. 1981. Results of the Alcoa Foundation-Suriname Expeditions. VI. Additional chromosomal data for bats (Mammalia: Chiroptera) from Suriname. Ann. Carnegie Mus., 50:333-344. Genoways, H. H., and S. L. Williams. 1979. Records of bats (Mammalia: Chiroptera) from Suriname. Ann. Carnegie Mus., 48:323-335.
— . 1984. Results of the Alcoa Foundation-Suriname Expeditions. IX. Bats of the
genus Tonatia (Mammalia: Chiroptera) in Suriname. Ann. Carnegie Mus., 53:327- 346.
Genoways, H. H., S. L. Williams, and J. A. Groen. 1981. Results of the Alcoa Foundation-Suriname Expeditions. V. Noteworthy records of Surinamese mam- mals. Ann. Carnegie Mus., 50:319-332.
Goodwin, G. G., and A. M. Greenhall. 1961. A review of the bats of Trinidad and Tobago. Amer. Mus. Nat. Hist, 122:187=302.
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Annals of Carnegie Museum
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Hall, E. R. 1981. The mammals of North America. John Wiley & Sons, New York, l:xv + 1-600 + 90.
Hill, J. E. 1964. Notes on bats from British Guiana, with the description of a new genus and species of Phyllostomatidae. Mammalia, 28:553-572.
Honeycutt, R. L., R. J. Baker, and H. H. Genoways. 1980. Results of the Alcoa Foundation-Suriname Expeditions. III. Chromosomal data for bats from Suriname. Ann. Carnegie Mus., 49:237-250.
Husson, a. M. 1978. The mammals of Suriname. Zool. Monog., Rijksmuseum Nat. Hist., 2:xxiv + 1-569.
Jones, J. K., Jr., and D. C. Carter. 1 976. Annotated checklist, with keys to subfamilies and genera. Pp. 7-38, in Biology of bats of the New World family Phyllostomatidae, Part I (R. J. Baker, J. K. Jones, Jr., and D. C. Carter, eds.). Spec. Publ. Mus., Texas Tech Univ., 10:1-218.
Koopman, K. F. 1978. Zoography of Peruvian bats with special emphasis on the role of the Andes. Amer. Mus. Novitates, 2651:1-33.
Koopman, K. F., and E. L. Cockrum. 1967. Bats. Pp. 109-150, in Recent mammals of the World (S. Anderson and J. K. Jones, Jr., eds.), The Ronald Press Co., New York, viii + 453 pp.
LaVal, R. K. 1977. Notes on some Costa Rican bats. Brenesia (Museo Nacional de Costa Rica), 10/11:77-83.
Medellin, R, A., D. E. Wilson, and D. Navarro L. 1985. Micronycteris brachyotis. Mamm. Species, 251:1-4.
Miller, G. S., Jr. 1907. The families and genera of bats. Bull. U.S. Nat. Mus., 57: xvii + 1-282.
Sanborn, C. C. 1949. Bats of the genus Micronycteris and its subgenera. Fieldiana: Zook, 31:215-233.
Williams, S. L., and H. H. Genoways. 1 980. Results of the Alcoa Foundation-Surina- me Expeditions. II. Additional records of bats (Mammalia: Chiroptera) from Su- riname. Ann. Carnegie Mus., 49:213-236.
Back issues of many Annals of Carnegie Museum articles are available, and a few early complete volumes and parts are listed at half price. Orders and inquiries should be addressed to: Publications Secretary, Carnegie Museum, 4400 Forbes Avenue, Pittsburgh, Pa. 15213.
AS
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5 0 /, 73
/
ISSN 0097-4463'
ANNALS*
of CARNEGIE MtlSEHiBl
CARNEGIE MUSEUM OF NATURAT"' HISTORY '
4400 FORBES AVENUE • PITTSBURGH, PENNSYLVANIA 15213 VOLUME 55 31 DECEMBER 1986 ARTICLE 14
ANNOTATED CHECKLIST OF THE MAMMALS OF KENYA. II. INSECTIVORA AND MACROSCELIDEA
ISSA R. Agoundey'
Resident Museum Specialist, Section of Mammals
Duane A. Schlitter
Curator, Section of Mammals
Abstract
Four families of Insectivora (Tenrecidae, Chrysochloridae, Erinaceidae, Soricidae) and one of Macroscelidea (Macroscelididae) are known from Kenya. Of these families, only the Soricidae, with 31 species, and the Macroscelididae, with five species, are known from more than a single species. Records of occurrence are annotated by museum spec- imens or with references to the literature. Suncus murinus seems to be the only introduced species in the insectivore fauna of Kenya. Distributional records of insect! vores in Kenya are poor as 1 3 species of the 39 reported are known from only a single locality in Kenya. A gazetteer of collecting localities is included.
Introduction
This is the second in a series of annotated checklists on Kenya mam- mals, The first checklist (Aggundey and Schlitter, 1984) covered the bats. This checklist includes the insectivoran orders Insectivora and Macroscelidea. Our treatment of this group follows Yates (1984). In this checklist we cover four Lipotyphlan families, namely the Tenreci- dae, Chrysochloridae, Erinaceidae, and Soricidae, and a single Men- otyphlan family, the Macroscelididae, of Kenya.
‘ Address: Curator of Mammals, National Museums of Kenya, P.O. Box 40658, Nairobi,
Kenya.
Submitted 27 January 1986.
325
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Annals of Carnegie Museum
VOL. 55
In a series of papers, Dollman (1915<2, 1915^?, 1915c, \9\5d, 1915c, 1915/ 1916) reviewed the African species of Crocidura. This review was followed by Hollister’s (1918) review of Kenyan insectivores in the United States National Museum. In his massive checklist, Allen (1939) covered all of the species of Kenyan insectivores recognized at that time. This work is still important as a starting point in under- standing the taxonomic relationships of these species. Since these early papers, there have been no subsequent reviews in a wider context of Kenyan insectivores except elephant shrews. Corbet and Neal (1965) and Corbet and Hanks (1968) reviewed the taxonomy and distribution of Kenyan species in their broader studies of elephant shrews. This second annotated checklist attempts to synthesize the pertinent liter- ature on all of the insectivores known from Kenya. The taxonomy of many groups of insectivores, and particularly the family Soricidae in Africa, is still in a state of confusion. Undoubtedly many taxonomic changes and additional distributional records will be required before a satisfactory arrangement is possible.
We have followed the format from the first part of the series (Ag- gundey and Schlitter, 1984). Specimen records are included from the following museums with their accepted acronyms in parentheses.
Carnegie Museum of Natural History, Pittsburgh (CM)
National Museums of Kenya, Nairobi (NMK)
National Museum of Natural History, Smithsonian Institution, Washington, D.C. (USNM)
As with the checklist on bats, we point out that the distributional records of this group are also undoubtedly incomplete. Nevertheless, it is our sincere hope that the information given will stimulate addi- tional interest and research on the insectivores of Kenya.
Acknowledgments
We were unable to spend as much time gathering distributional records from museum collections for this checklist as we were for the checklist on bats. Nevertheless, we would like to thank Charles O. Handley, Jr., for allowing us to record data at the Smithsonian Institution and for sending photocopies of numerous old and difficult to find articles. In Kenya, Mrs. Gooderis helped compile the list of the specimens of insectivores in the collection in Nairobi. R. Hutterer critically reviewed the manuscript. Winnie Woodland and Mary Ann Schmidt patiently and expertly typed numerous drafts of the manuscript. We thank them all for their important part in completing this checklist.
This checklist resulted from the first author’s tenure as a Resident Museum Specialist at Carnegie Museum of Natural History. This visit was made possible by the International Program, endowed by the Richard King Mellon Foundation. Lastly, the senior author wishes especially to thank Richard E. Leakey, Director of the National Museums of Kenya, for making it possible for him to take six months study leave in the United States.
1986 Aggundey AND ScHLiTTER— Kenyan Insectivora AND Macroscelidea 327
Checklist
Order Insectivora Family Tenrecidae Genus Potamogale Du Chaillu, 1860 Potamogale velox Du Chaillu, 1860
Potamogale velox Du Chaillu, 1860. Proc. Boston Soc. Nat. Hist., 7:361. Gabon. Records.— A km S Kakamega (Aggundey, 1977:368, NMK).
Family Chrysochloridae Genus Lacepede, 1799
Chrysochloris stuhlmanni fasten (St. Leger, 1931)
Chlorotalpa fosteri St. Leger, 1931. Ann. Mag. Nat. Hist., (10)8:605, Dec. Mt. Elgon, 9300 ft, Uganda.
Records.— MX. Elgon (NMK); Cherangani Hills (Duncan and Wrangham, 1971:149; Meester, 1974:3); southwest side Cherangani Hills, 6500 ft (Duncan and Wrangham, 1971:157); Labot, Cherangani Hills (Duncan and Wrangham, 1971:157).
Family Erinaceidae Genus Atelerix Pomel, 1848 Atelerix albiventris (Wagner, 1841)
E{rinaceus) albiventris Wagner, 1841. Schreber’s Saugethiere, Suppl., 2:22. Type locality unknown.
Erinaceus albiventris atratus Rhoads, 1896. Proc. Acad. Nat. Sci., Philadelphia, p. 544, 8 Dec. Lake Rudolf, Ngare Nocbor, Marsabit Dist., Kenya (ca. 2°45'N, 36°45'E). Erinaceus hindei Thomas, 1910. Ann. Mag. Nat. Hist., (8)5:193, Feb. Kitui, 3500 ft, Kenya.
Erinaceus sotikae Heller, 1910. Smithsonian Misc. Coll., 56(15):!, 23 Dec. Southern Guaso Nyiro, Sotik Dist., Kenya.
Ngare Nocbor (Rhoads, 1896:544); Kitui (Peters, 1878:198; Anderson, 1895: 420; Thomas, 1910<3:193); Southern Guaso Nyiro (Heller, 1910^:1; Hollister, 1918:26; J. Allen, 1922:16); Kapiti Plains (Hollister, 1918:26; J. Allen, 1922:16); Loita Plains (Hollister, 1 9 1 8:26); Mt. Lololokwi (Hollister, 1 9 1 8:26; J. Allen, 1 922: 1 6); Taveta (True, 1892:469, 480; Hollister, 1918:26); Ulukenia Hills (Hollister, 1918:26; J. Allen, 1922: 16); Upper Ura River (J. Allen, 1922:16); Lokori, Southern Turkana (NMK); Naivasha (NMK); Nairobi (Harmsen and Jabbal, 1968:158; Kingdon, 1974:32; Kock, 1978:116, NMK); Busia (CM); Voi (Allen and Lawrence, 1936:39); Lodwar (St. Leger, 1937:526); Olorgasailie (Toschi, 1949:28); River Kerio Suk (Ruxton, 1926:29); Narro Surra River (Kollmann, 1914:319); Machakos (Lonnberg, 1912Z?:48).
Family Soricidae Genus CwciWwm Wagier, 1832 Crocidum allex Osgood, 1910
Crocidura allex Osgood, 1910. Publ. Field Mus. Nat. Hist., Zool. Ser., 10(3):20, 7 Apr. Naivasha, Kenya.
Crocidura alpina Heller, 1910. Smithsonian Misc. Coll., 56(9):5, 22 July. West slope of Mt. Kenya, 10,000 ft, Kenya.
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^FCor^/5.~Naivasha Station (Osgood, 1910^7:20; Hollister, 1918:67, USNM); West slope Mt. Kenya (Heller, 1910^2:5; Hollister, 1918:68, USNM); Aberdare Range (Hol- lister, 1918:67, USNM); Oljoro O Nyon River (Hollister, 1918:67, USNM); Man Forest (Toschi, 1947:12, NMK); Ngong (NMK); Loita (NMK); Amala River (NMK); Selangai (NMK); near Lake Olbollossat (Dollman, 1915/:513); Solai, 8000 ft (Dollman, 1915/: 514); Nanyuki (Southern and Hook, 1963:511); Mt. Kenya, 12,500 ft (Harmsen and Jabbal, 1968:160); north slope Mt. Kenya (Coe and Foster, 1972:8).
Remarks. —WQim de Balsac and Meester (1977:9) regard both C a. allex and C a. alpina as valid subspecies in Kenya.
Crocidura bottegi Thomas, 1898
Cwcidura bottegi Thomas, 1898. Ann. Mus. Civ. Stor. Nat. Genova, (2): 18, 677, 21
Feb. Between Badditu and Dime, northeast of Lake Rudolf, Ethiopia.
Records. — MditsdihiX (Heim de Balsac and Meester, 1977:10).
Crocidura elgonius Osgood, 1910
Crocidura bicolor elgonius Osgood, 1910. Ann. Mag. Nat. Hist., (8)5:369. Kirui’s, south- ern foothills of Mount Elgon, 6000 ft, Kenya (restricted by Moreau et al., 1946:394).
RFCor<7s'. — Kirui’s (Osgood, 1910c:369; Allen and Lawrence, 1936:44; Loveridge, 1937: 519); Kisumu (Hollister, 1918:65); Lukosa River (Hollister, 1918:65, USNM); Kapiti Plains (Hollister, 1918:65, USNM); Kaimosi (Hollister, 1918:65; Allen and Lawrence, 1936:44; Allen and Loveridge, 1942:159, NMK, USNM); Muguga (NMK); Lake Nakuru (Hutterer, 1983:225); Nairobi (Hutterer, 1983:225); Cherangani Hills (Ruxton, 1926: 29); Eldoret (Loveridge, 1923:698); Mt. Elgon (Granvik, 1924:9).
Remarks. — Both Heim de Balsac and Meester (1977) and Hutterer (1984) regard C elgonius as a distinct species.
Crocidura fischeri Pagenstecher, 1885
Crocidura fischeri Pagenstecher, 1885. Jahrb. Wiss. Anst., Hamburg, 2:34, pi. 1, f 1--3. Nguruman, Kenya.
JRFCorflf5.““Nguruman (Pagenstecher, 1885:34; Hutterer, 1986:26).
Remarks. —ThQ type locality for the species was given by Pagen- stecher (1885:34) as Nguruman with no other qualifier than Massai- land. Neumann (1900) also visited Nguruman from 22-27 December 1893 and plots the locality on his map as slightly on the Tanzanian side of the border (which seems to be the same as the present border). He labels the locality on his map as “Nguruman (Bagasse)” which leads us to believe he was referring to a village near Lake Natron or “Ngu- ruman Salz-See” from his map. Lake Natron seems to have been at a low level in 1893. Swynnerton and Hayman (1951:284) list the type locality of Crocidura fischeri as Nguruman, west of Lake Magadi, Kenya Colony [between L50' and 2®S, 36°5'E; 2500 to 6000 feet]. This locality reaches from the Rift Valley floor to nearly up to the top of the eastern face of the Nguruman Escarpment. Hutterer (personal communication)
1986 Aggundey AND ScHLiTTER— Kenyan Insectivora AND Macroscelidea 329
has rechecked this type locality as part of a study (Hutterer, 1986) of C fischeri and concluded that it is in Kenya based on Fischer’s (1884, 1885) accounts of his explorations and especially the photographs and maps in these accounts. Fischer camped at the foot of a mountain range northwest of “Natron Swamp” or Lake Natron at 650 meters, near the foot of Mt. Sambo, where acacia steppe occurs and a flowing stream came down from the mountains. Mt. Sambo, although on the Tan- zanian side of the border, is visible far in the background of a pho- tograph of the campsite. It seems best to follow Swynnerton and Hay- man (1951) and Hutterer (1986) in placing the type locality on the Kenyan side of the present border.
Crocidura flavescens nyansae Neumann, 1900
Crocidura flavescens nyansae Neumann, 1900. Zool. Jahrb., Syst., Geog., Biol., 13:544, 10 Oct. Fort Lubwa’s in Ussoga, outlet of Victoria Nyanza, Uganda (restricted to Fort Thruston, 10 mi E Jinja, Busoga Dist., Uganda, by Moreau et al., 1946:396). Crocidura kijabaej. A. Allen, 1909. Bull. Amer. Mus. Nat. Hist, 26:173. Kijabe, Kenya.
Records. — KAjabe (J. Allen, 1909:173); Elgeyo Forest (J. Allen, 1914:343); Aberdare Mountains, 11,000 ft (Dollman, 1915^:568; Hollister, 1918:43); Laikipia (Hollister, 1918:43); Mt. Kenya (Dollman, 19156:568; Hollister, 1918:43); Mt. Umengo (Hollister,
1 9 1 8:43); Naivasha Station (Hollister, 1 9 1 8:43); Nakutishu River, Naivasha Plains (Hol- lister, 1918:43); Nyeri (Dollman, 19156:568; Hollister, 1918:43); Kaimosi (Hollister, 1918:42; Allen and Lawrence, 1936:41); Kakamega (Hollister, 1918:42); Kisumu (Hol- lister, 1918:42); Sergoit Lake (Hollister, 1918:42); Jombeni (Dollman, 19156:568); Mweru (Dollman, 19156:568); Lake Olbollosat (Dollman, 19156:568); Kirui’s (Dollman, 19156: 567); Londiani (Lonnberg, 1918:175); Lake Elmenteita (Osgood, 1936:221); Molo (Os- good, 1936:221); Mianzini (Thomas, 1891:182); Meru Country (Lonnberg, 19126:52); Mt. Elgon (Granvik, 1924:8).
Crocidura fulvastra (Sundevall, 1 843)
Sorex fulvaster SnndQvaW, 1843. Kongl. Svenska Vet.-Akad. Handl., Stockholm, p. 172, for 1842. Bahr-el-Abiad, Sudan.
North of Lokichokio (Hutterer, 1984:215).
Remarks. species includes C. sericea (Sundevall, 1843) as a synonym according to Hutterer (1984:21 1, 215).
Crocidura fumosa fumosa Thomas, 1 904
Crocidura fumosa Thomas, 1904. Ann. Mag. Nat, Hist., (7)14:238, Sept. Western slope
of Mt. Kenya, 2600 m, Kenya.
Crocidura alchemillae Heller, 1910. Roosevelt’s African Game Trails, American ed., p. 480, London ed., p. 491. Summit of Aberdare range, Kenya.
Records.— WQsXQvn slope ofMt. Kenya, 2600 m (Thomas, 1904:238; Dollman, 191 5£’: 369, 370); summit of Aberdare range (Heller, 1910c:480; Hollister, 1918:55); Kinangop (Kollmann, 1913:1 39); west side Mt. Kenya (Hollister, 1918:55, NMK); Fort Hall (Thomas, 1904:238; Kollmann, 1913:139; Hollister, 1918:55, NMK); Chyulu Hills (Osgood, 19106: 21, NMK); Nairobi (NMK); Ngong (NMK); Chania River (NMK); Naro Mom (NMK);
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01 Arabel (NMK); Kikuyu (NMK); Thika (NMK); Nyeri (Hollister, 1918:55); Upper Nzoia River (Hollister, 1918:55); Mt. Elgon (Dollman, 1915^:369, 370); Jombeni Range (Dollman, 1915e:369, 370); Aberdare Mountains (Dollman, 1915^:369, 370); Machakos (Thomas, 1904:238; Kollmann, 1913:139); Nandi (Thomas, 1904:238; Kollmann, 1913: 139); Kakamega (Thomas, 1904:238; Kollmann, 1913:139); Donya Sabuk (Lonnberg, 1916:5); Juja Farm (Lonnberg, 1916:5); Nanyuki (Southern and Hook, 1963:512); Mt. Kenya (Southern and Hook, 1 963:5 1 2; Duncan and Wrangham, 1 97 1 : 1 6 1); 2^bra Farm, Athi Plains (J. Allen, 1909:173); Lagari (Thomas, 1904:238); Blue Post (Lonnberg, 1912Z?: 53); Kagio (Lonnberg, 1912Z):53); Embu (Lonnberg, 1912Z?:53); Kanyakeni (Lonnberg, 1912Z?:53); Meru (Lonnberg, 1912Z):53); Mt. Kenya, 3800 m (Coe and Foster, 1972:8).
Remarks. — Y^onnhQrg (1 9 1 6:5) considered the series of 10 specimens from Donya Sabuk to approach C / schist acea, now considered a synonym of C luna. It is doubtful that all of these listed records rep- resent C. fumosa as the differences between C fumosa, C. luna, and C. zaodon are subtle ones. Many of these records could prove to be C. luna upon reexamination.
Crocidura fuscomurina (Heuglin, 1865)
S.{orex) fusco murinus Heuglin, 1865. Leopoldina, 5:36, in Nova Acta Acad. Caes.
Leop.-Carol., Dresden, June. Meshra-el-Req, Bahr-el-Ghazal Prov., Sudan. Crocidura ^/co/or Bocage, 1889. Jom. Sci. Math., Phys. e Nat., Lisboa, (2)1:29, March.
“Gambos, dans I’interieur de Mossamedes,” Angola.
Crocidura bicolor cuninghamei Thomas, 1904. Ann. Mag. Nat. Hist., (7)14:240, Sept. Vumba Island, 1 mi N Sagitu Island, Lake Victoria, Uganda.
Nairobi (Osgood, 1936:230).
Remarks. Hutterer (1 983) reviewed the taxonomy and distribution of C fuscomurina but did not report any records from Kenya. His preliminary distribution map (Hutterer, 1983:224) included only rec- ords for which he had some degree of certainty of identification.
Crocidura hildegardeae Thomas, 1 904
Crocidura hildegardeae Thomas, 1904. Ann. Mag. Nat. Hist., (7)14:240, Sept. Fort Hall, Kenya.
Crocidura lutreola Heller, 1912. Smithsonian Misc. Coll., 60(1 2):8, 4 Nov. Mt. Mbololo, Taita Hills, 5000 ft, Kenya.
Crocidura hildegardeae procera Heller, 1912. Smithsonian Misc. Coll., 60( 1 2): 1 0, 4 Nov.
Mt. Lololokwi, 6000 ft, northern Guaso Nyiro, Kenya.
Crocidura ibeana Dollman, 1915. Ann. Mag. Nat. Hist., (8)15:514, May. Ann. Mag.
Nat. Hist., (8)16:362, Oct. Olgerei River, Kenya.
Crocidura hildegardeae altae Heller, 1912. Smithsonian Misc. Coll., 60(1 2):9, 4 Nov. Mt. Garguez, Mathews Range, 6000 ft, Kenya.
Records. — Fort Hall (Thomas, 1904:240; Dollman, 1915/:508; Hollister, 1918:64, USNM); Mt. Garguez (Heller, 1912:9, 10; Hollister, 1918:65, USNM); Mt. Lololokwi (Heller, 1912:10; Hollister, 1918:64); Mt. Mbololo (Heller, 1912:8, 9; Hollister, 1918: 64; Allen and Lawrence, 1936:43, USNM); Olgerei River (Dollman, 1915<2:514); Voi (Hollister, 1 9 1 8:64, NMK); Ngong (NMK); Narosura River (Kollmann, 1 9 1 4:3 1 9, NMK); Lemek Valley (NMK); Amala River (NMK); Nyeri (Hollister, 1918:64, NMK); Meru (Hollister, 1 9 1 8:64, NMK); Amboseli (NMK); Engare Narok (Hollister, 1 9 1 8:64, NMK);
1986 Aggundey AND ScHLiTTER— Kenyan Insectivora AND Macroscelidea 331
Isiola River (Hollister, 1 9 1 8:64); Kapiti Plains (Hollister, 191 8:64); Mayo River, Laikipia (Hollister, 1 9 1 8:64, USNM); Mt. Kenya (Hollister, 1 9 1 8:64); Mt. Sagalla (Hollister, 1918: 64, USNM); Mt. Umengo (Heller, 1912:9; Hollister, 1918:64, USNM); Naivasha Station (Hollister, 1918:64, NMK); Ndi (Hollister, 1918:64, USNM); Oljoro O Nyon River (Hollister, 1918:64); Wambugu (Hollister, 1918:64); Taveta (Dollman, 1915^:379); Tsa- vo River (Dollman, 1915^:380); Mt. Elgon (Dollman, 1915/:508); Baringo (Dollman, 1915/:509); Kaimosi (Allen and Lawrence, 1936:43); Peccatoni (Allen and Lawrence, 1936:43); Wema (Allen and Lawrence, 1936:43); Kazere (Lonnberg, 1912Z):54); Blue Post (Lonnberg, 1912^:54); Kutu (Lonnberg, 1912Z?:54).
Remarks. —We follow Demeter and Hutterer (1986) and Hutterer (personal communication) in recognizing C hildegardeae for these
Kenyan records rather than C gracilipes Peters, 1870, an apparently distinct species.
Crocidura hirta Peters, 1852
Crocidura hirta Peters, 1852. Reise nach Mossambique, Saugeth., p. 78, pi. 18, f. 2. Tette, Mozambique (17°S).
Records.— I^OVLQ found.
Remarks.— Yieim de Balsac and Meester (1977:17) list C h. velutina Thomas, 1904 to occur in . . presumably also Kenya and southern Somalia.”
Crocidura Jacksoni Thomas, 1 904
Crocidura jacksoniThom^s, 1904. Ann. Mag. Nat. Hist., (7)14:238, Sept. Ravine Station,
Kenya.
Crocidura jacksoni amalae Dollman, 1915. Ann. Mag. Nat. Hist, (8)15:516, May; Ann. Mag. Nat. Hist, (8)16:376, Oct. 1915. Amala River, NyanzaProv., Kenya (restricted by Moreau et at, 1946:396 to Amala River, 30 mi N Kenya-Tanzania border, 5500 ft, Kenya).
Records.— Station (Thomas, 1904:239); Amala River (Dollman, 1914^:309, 1 9 1 5<3:5 1 6, NMK); Isiola River (Hollister, 1 9 1 8:60); Kaimosi (Hollister, 1 9 1 8:60, USNM); Kapiti Plains (Hollister, 1 9 1 8:60); Mtito Andei (Hollister, 1 9 1 8:60, USNM); Neumann’s Boma (Hollister, 1918:60); Southern Guaso Nyiro (Hollister, 1918:60); Ulukenia (Hol- lister, 1918:60); Voi (Hollister, 1918:60, Allen and Lawrence, 1936:43, USNM); Na- rosura River (Dollman, 1914/7:309, NMK); Loita Plains (NMK); Ngong (NMK); Fort Hall (NMK); Tsavo River (Dollman, 1914^z:88, NMK); Yala River (NMK); Sultan Hamud (NMK); Shimba Hills (NMK); Peccatoni (Allen and Lawrence, 1936:43); Gol- banti (Allen and Lawrence, 1936:43); Zuwani (Dollman, 19 14^z:88); Lengototo (Dollman, 1914/7:309).
Crocidura littoralis Heller, 1910
Crocidura littoralis Heller, 1910. Smithsonian Misc. Colt, 56(1 5):5, 23 Dec. Butiaba, east shore of Albert Nyanza, Uganda.
— Kaimosi (Hollister, 1918:68; Dippenaar, 1980:129).
Remarks.— Hollister (1918:68) referred this series to C maurisca but Dippenaar (1980:130) regards this species as known only by the holotype and that the specimens from Kaimosi are C. littoralis.
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Crocidura luna Dollman, 1910
Crocidura luna Dollman, 1910. Ann. Mag. Nat. Hist., (8)5:175, Feb. Bunkeya River, Katanga, Zaire.
Crocidura fumosa schistacea Osgood, 1910. Publ. Field Mus. Nat. Hist., Zool. Ser., 10(3): 20, 7 April. Lukenya Mountain, Kenya.
Crocidura raineyi Heller, 1912. Smithsonian Misc, Coll., 60(1 2):7, 4 Nov. Mt. Gargues, Kenya.
Crocidura fumosa selina Dollman, 1915. Ann. Mag. Nat. Hist., (8)15:510, May; Ann.
Mag. Nat. Hist., (8)16:371, Oct., 1915. Mabira Forest, Chagwe, Uganda.
Crocidura luna umbrosa Dollman, 1915. Ann. Mag. Nat. Hist., (8)15:514, May; Ann. Mag. Nat. Hist., (8)16:360, Oct., 1915. Machakos, 5400 ft, Kenya.
Records. —Lukenya Mountain (Osgood, \9\0b:2Q)\ Mt. Gargues (Heller, 1912:7; Doll- man, 1915^:373; Hollister, 1918:60); Machakos, 5400 ft (Dollman, 1915^:514, \9\5e: 361); Ulukenia Hills (Hollister, 1918:59); Kapiti Plains (Hollister, 1918:59); Kaimosi (Hollister, 1918:59); Mt. Elgon (Rode, 1935:167).
Remarks. de Balsac and Meester (1977:18) list only C. /.
schistacea occurring in Kenya and point out that C. raineyi could be a valid large subspecies. Hollister (1 9 18:59) listed specimens from Kai- mosi in western Kenya as C /. selina.
Crocidura macarthuri St. Leger, 1934
Crocidura macarthuri St. Leger, 1934. Ann. Mag. Nat. Hist., (10)13:559, May. Merifano, 20 mi from mouth of Tana River, Kenya.
Recor^/5.— Merifano (St. Leger, 1934:559; Hutterer, 1986:28); lOmiEMoyal, onMurri Road, 1500 m (Heim de Balsac and Meester, 1977:19; Hutterer, 1986:28); Ijara (Percy; et al., 1953Z?:11; Heim de Balsac and Meester, 1977:19; Hutterer, 1986:28); Rojewero Plains (Hutterer, 1986:28).
Crocidura macowi Dollman, 1915
Crocidura macowi Dollman, 1915. Ann. Mag. Nat. Hist., (8)15:515, May; Ann. Mag. Nat. Hist., (8)16:378, Oct. 1915. Mt. Nyiro, south of Lake Rudolf, Kenya.
Records.-MX. Nyiro (Dollman, 1915^:515, 1915^:378).
Crocidura monax monax Thomas, 1910
Crocidura monax Thomas, 1910. Ann. Mag. Nat. Hist., (8)6:310, Sept. Rombo, Mt. Kilimanjaro, 6000 ft, Tanzania.
Records.— River (Dollman, 1914Z?:309).
Crocidura nanilla Thomas, 1 909
Crocidura nanilla Thomas, 1909. Ann. Mag. Nat. Hist., (8)4:99, Aug. Probably Entebbe, Uganda.
Crocidura denti St. Leger, 1932. Ann. Mag. Nat. Hist., (10)9:240, March, Koliokwell River, North Turkana Dist., Kenya (not C. jacksoni denti Dollman, 1915). Crocidura rudolfi St. Leger, 1932. Ann. Mag. Nat. Hist., (10)10:487, Nov. (renaming of C. denti St. Leger, 1932).
— Koliokwell River (St. Leger, 1932:241).
1986 Aggundey AND ScHLiTTER— Kenyan Insectivora AND Macroscelidea 333
Remarks.— de Balsac and Meester (1977:20) place C rudolfl as a synonym of C. nanilla.
Crocidum parvipes Osgood, 1910
Crocidura parvipes Osgood, 1910. Publ. Field Mus. Nat. Hist., Zool. Ser., 10(3): 19, 7
Apr. Voi, Kenya.
Crocidura parvipes nisa Hollister, 1916. Smithsonian Misc. Coll., 66(8):2, May. Kibabe, Kisumu, Kenya.
Eusso Nyiro Post (NMK); Voi (Osgood, 1910Z?:19; Hutterer, 1986:31); Ki- babe (Hollister, 1 9 1 66:2; Hollister, 1 9 18:47, USNM); Embu (NMK); Fort Hall (Hollister, 1918:47); Mt. Sagalla (Heller, 1912:9; Hollister, 1918:60; Heim de Balsac and Meester, 1977:12, USNM).
Remarks. — Three specimens of shrews from Mt. Sagalla are reported as C. parvipes by Heller (1912:9) but are referred to C jacksoni by Hollister (1918:63). Heim de Balsac and Meester (1977:12) refer this record again to C. cyanea parvipes. Both C. c. parvipes and C c. nisa are recognized as valid in Kenya by Heim de Balsac and Meester (1 977: 12). Hutterer (1986:31) considered C. parvipes to be distinct from C. cyanea.
Crocidura ultima Dollman, 1915
Crocidura ultima Dollman, 1915. Ann. Mag. Nat. Hist., (8)15:517, May; Ann. Mag. Nat. Hist., (8)17:204, Feb., 1916. Jombeni Range, Nyeri Dist., 5000 ft, Kenya.
Records. —iomhQni Range, 5000 ft (Dollman, 1915<2:517, 1916:205).
Remarks.— de Balsac and Meester (1977:20) plaee C ultima as a synonym of C. monax, but Dippenaar (1980:130) considers C. ultima to be a distinct species known only from the holotype.
Crocidura viaria (1. Geoffroy Saint-Hilaire, 1834)
Sorex viarus 1. Geoffroy Saint-Hilaire, 1834. Voyage aux Indes-Orientales par C. Belan- ger, ZooL, p. 1 27. Senegal (restricted by Hutterer, 1984:209, to region between Dakar and St. Louis).
Crocidura hindei Thomas, 1904. Ann. Mag. Nat. Hist., (7)14:237, Sept. Machakos, Kenya.
Crocidura suahelae Heller, 1912. Smithsonian Misc. Coll., 60(1 2):6, 4 Nov. Mazeras, Kenya.
Crocidura beta Dollman, 1915. Ann. Mag. Nat. Hist., (8)15:513, May; Ann. Mag. Nat. Hist., (8)16:78, July, 1915. Chania River, Kenya (restricted to Chania River, near Nyeri and Fort Hall, Kenya by Allen, 1939:31).
Machakos (Thomas, 1904:237; Hutterer, 1984:211); Mazeras (Heller, 1912; 6; Hollister, 1918:50, USNM); Chania River (Dollman, 1915^3!:513, 1915c:78); Chan- gamwe (Hollister, 1918:50, UNSM); Juja Farm (Lonnberg, 19126:54; Hollister, 1918: 46); Ulukenia Hills (Hollister, 1918:46); Nairobi (Loveridge, 1923:698, NMK); Kajiado (NMK); Potha (NMK); Ngatana (Allen and Lawrence, 1936:42); Mombasa (Hutterer, 1984:211); Karati (Thomas, 1904:237).
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Remarks. (1984) placed C. sericea under C. fuivastra (Sundevall, 1 843) but regarded C hindei and C suahelae as conspecific with C viaria.
Crocidura voi Osgood, 1910
Crocidura voi Osgood, 1910. Publ. Field Mus. Nat. Hist., ZooL Sen, 10(3): 18; 7 April. Voi, Kenya.
Crocidura percivali Dollman, 1915. Ann. Mag. Nat. Hist, (8)15:513, May; Ann. Mag. Nat Hist, (8)16:126, Aug., 1915. Jombeni Range, Nyeri Dist, 3500 ft, Kenya.
Records.— Yoi (Osgood, 1910Z?:18; Hutterer, 1986:30) Jombeni Range (Dollman, 1915a: 5 1 3; Hutterer, 1 986:30); Lakiundu River (Hollister, 1 9 18:50); Mt. Suswa (Hutterer, 1 986: 30).
Remarks.— Hutterer (1986) recognized Crocidura voi as a distinct species, and included C. butleri from Sudan, C percivali from Kenya, and C aridula from Sudan as synonyms.
Crocidura xantippe Osgood, 1910
Crocidura xantippe Osgood, 1910. Publ. Field Mus. Nat. Hist., Zool. Sen, 10(3): 19, 7 April. Voi, Kenya,
Records. — Voi (Osgood, 19 10/?: 19; Dollman, 1915^:375; Heim de Balsac and Meester, 1977:25); Taveta (Dollman, 1915^:375); Nyiru (Heim de Balsac and Meester, 1977:25); Tsavo (Heim de Balsac and Meester, 1977:25),
Crocidura yankariensis Hutterer and Jenkins, 1980
Crocidura yankariensis Hutterer and Jenkins, 1980. Bull. British Mus. (Nat. Hist.), Zool., 39:305. Futuk, 16 km E Yankari Game Reserve boundary, Nigeria (9®50'N, 10®55'E).
Records. — West of Lake Rudolf, Kakuma, 50-60 mi NW Lodwar (Hayman, 1937: 531; Hutterer and Jenkins, 1983:195).
Crocidura zaodon Osgood, 1910
Crocidura turba zaodon Osgood, 1910. Publ. Field Mus. Nat. Hist., Zool. Sen, 10(3):21, 7 Apr. Nairobi, Kenya.
Crocidura turba provoeax Thomas, 1910. Ann. Mag. Nat, Hist., (8)6:1 12, July. Aberdare Mountains, 1 1 ,000 ft, Kenya.
Crocidura turba iakiundae Heller, 1912. Smithsonian Misc. Colt, 60(1 2):6, 4 Nov. Lak- iundu River, near junction with Northern Guaso Nyiro, Kenya,
Crocidura turba kempi Dollman, 1915. Ann, Mag. Nat, Hist,, (8)15:511, May; Ann. Mag. Nat. Hist., (8)16:134, Aug., 1915. KimPs, southern foothills of Mount Elgon, 6000 ft, Kenya (restricted by Moreau et aL, 1946:397).
Records.— Nmmhi (Osgood, 1910Z?:21; Thomas, 1910^:113; Lonnberg, 1918:175, NMK); Aberdare Mountains, 1 1,000 ft (Thomas, 1910Z):1 13; Dollman, 19 15a: 133; Hol- lister, 1918:54, NMK); Lakiundu River (Heller, 1912:6; Lonnberg, 1912l?:54; Hollister, 1918:54); Kinangop (Kollmann, 1913:140); Kirafs, Mt. Elgon, 5000-6000 ft (Dollman, 191 5^7: 134); Sirgoit Lake (Hollister, 1918:54); Sirgoit (Hollister, 1918:54); Kakamega (Hollister, 1918:54); Kibabe (Hollister, 1918:54); Naivasha Plains (Hollister, 1918:54); Mt. Kenia, west slope (Hollister, 1918:54); Isiola River, head (Heller, 1912:7; Hollister, 1918:54); Archer's Post (Heller, 1912:7; Hollister, 1918:54); Mt. Mbololo (Hollister, 1 9 1 8:54); Mt. Umengo (Hollister, 1 9 1 8:54); Nzoia River, Guas Ngishu plateau (Hollister,
1986 Aggundey and Schlitter— Kenyan Insectivora and Macroscelidea 335
1918:54); Mt. Sagalla (Hollister, 1918:54); Yala River (NMK); Nyeri (NMK); Kaimosi (Hollister, 1 9 1 8:54; Allen and Lawrence, 1 936:42, NMK); Kenna (NMK); Kericho (NMK); Maua (NMK); Man Forest (Toschi, 1 947: 1 2, NMK); Kasigau (NMK); Kisumu (Hollister, 1918:54, NMK); Chania River (NMK); Amala River (Dollman, 1915d:132, NMK); Kabete (NMK); Fort Hall (Hollister, 1918:54, NMK); Laikipia plateau, 15 mi N Nyeri (Hollister, 1 9 1 8:54); Lukosa River (Hollister, 1 9 18:54); Northern Guaso Nyiro (Dollman, 1915€/:132); Jombeni Range (Dollman, 191 5£/: 132); Donya Sabuk (Lonnberg, 1916:6); Cherengani Hills (Ruxton, 1926:29); Zuwani Swamp (Dollman, 1914^2:88); Nanyuki (Southern and Hook, 1963:512); Thomson’s Falls (Southern and Hook, 1963:512); Mt. Kenya, 1 2,500 ft (Harmsen and Jabbal, 1 968: 1 59); Mt. Kenia, 2450 m (Lonnberg, \9\2b: 54); Luazomela River (Lonnberg, 1912^:54); acacia steppe south of Guaso Nyeri (Lonn- berg, 1912Z):54).
Crocidura zaphiri simiolus Hollister, 1916
Crocidura simiolus Hollister, 1916. Smithsonian Misc. Coll., 66(8):3, May. Kisumu, Kenya.
Records. —Kisumu (Hollister, 19166:3; Hollister, 1918:51); Kaimosi (Hollister, 1918: 51); Kibabe (Hollister, 1918:51).
Genus Suncus Hemprich and Ehrenberg, 1832 Suncus infinitesimus infinitesimus (Heller, 1912)
Pachyura infinitesima Heller, 1912. Smithsonian Misc. Coll., 60(1 2):5, 4 Nov. Rumruti, Laikipia Plateau, 7000 ft, Kenya.
Records.— KummXi (Heller, 1912:5).
Suncus lixus aequatorius (Heller, 1912)
Pachyura lixus aequatoria Heller, 1912. Smithsonian Misc. Coll., 60(1 2):4, 4 Nov. Sum- mit of Mt. Sagalla, 4000 ft, Taita Hills, Kenya.
Recor<75.— Summit of Mt. Sagalla, 4000 ft (Heller, 1912:4; Hollister, 1918:41).
Suncus murinus (Linnaeus, 1766)
Sorex murina Linnaeus, 1766. Syst. Nat., 12th ed., 1:74. Java, Indonesia.
Records. (Heller, 1912:5); Witu (Heller, 1912:5).
Remarks. — This introduced species probably has a more widespread distribution, especially along the coast, than these few records repre- sent.
Genus Sylvisorex Thomas, 1 904 Sylvisorex grand mundus Osgood, 1910
Sylvisorex mundus Osgood, 1910. Publ. Field Mus. Nat. Hist., Zool. Sen, 10:18, 7 Apr. Kijabe, Kenya.
Records.— YA^dibQ (Osgood, 19106:18); west side Mt. Kenya, 7000, 8500, 10,000 ft (Hollister, 1918:39); Mt. Kenya (Duncan and Wrangham, 1971:161).
Remarks.— Tv^o additional locality records, perhaps Mt. Elgon and Cherangani Hills, are plotted by Kingdon (1974:81). The former may be Butandiga, Uganda, reported by Allen and Lawrence (1936:41).
336
Annals of Carnegie Museum
VOL. 55
Sylvhorex megalum gemmeus Heller, 1910
Sylvisorex gemmeus Heller, 1910. Smithsonian Misc. Coll., 56(1 5):7, 23 Dec. Rhino Camp, Lado Enclave, Uganda.
Sylvisorex sorelloides Lonnberg, 1912. Ann. Mag. Nat. Hist., (8)9:67, Jan.; Kungl. Svenska Vet.-Akad. Handl., Stockholm, (2)48(5):51, pi. 3, f. 1, 1912. Steppe near Itiolu River, Northern Guaso Nyiro, Kenya.
Records. — Acacia steppe, near Itiolu River, south of Northern Guaso Nyiro (Lonnberg, 191 2^2:67, 1912Z?:51); Kaimosi (Hollister, 1918:39; Allen and Lawrence, 1936:41); Kirui (Allen and Lawrence, 1936:41),
Genus Myosorex Gray, 1838 Myosorex (Surdisorex) norae (Thomas, 1906)
Surdisorex norae Thomas, 1906, Ann. Mag. Nat. Hist., (7)18:223, Sept. East side of Aberdare Range, near Nyeri, Kenya.
Records.— EsisX side of Aberdare Range, near Nyeri (Thomas, 1906:224); Aberdare Mountains, 10,000 to 11,000 ft (Hollister, 1918:37); Nr Kiandongoro Gate, Aberdare Mountains, 8400 ft (Duncan and Wrangham, 1971:160).
Myosorex (Surdisorex) polulus (Hollister, 1916)
Surdisorex polulus Hollister, 1916. Smithsonian Misc. Coll., 66(1):1, 10 Feb. West side of Mount Kenya, 10,700 ft, Kenya.
Records.— side of Mt. Kenya, 10,700 ft (Hollister, 1916<a!:l); west side of Mt. Kenya, 9000 ft to 12,000 ft (Hollister, 1918:37); Mt. Kenya, 12,500 ft (Harmsen and Jabbal, 1968:160); Mt. Kenya, 3960 m (Coe and Foster, 1972:8); Naro Mom track, Mt. Kenya, 10,500 ft (Duncan and Wrangham, 1971:160).
Order Macroscelidea Family Macroscelididae Genus Petrodromus Peters, 1 846 Petrodromus tetradactylus sangi Heller, 1912
Petrodromus sultani sangi Heller, 1912. Smithsonian Misc. Coll., 60(12):12. Mt. Mbol- olo, Taita Hills, 4000 ft, Kenya.
Records. -Mt. Mbololo, 4000 ft (Heller, 1912:12; Hollister, 1918:29; Allen and Law- rence, 1936:39; Loveridge, 1937:526); Voi (Corbet and Neal, 1965:68).
Remarks. — Corbet and Neal (1965:68) reported a skull in the Berlin Museum labelled “Kibwezi” but were uncertain if the record originated from the Chyulu Hills in Kenya or on Mt. Meru in Tanzania.
Petrodromus tetradactylus sultani Thomas, 1897
Petrodromus sultani Thom^is, 1897. Proc. Zool. Soc. London, p. 435. Mombasa, Kenya.
Records.— R.i\QT Mombaca (Guenther, 1881:164); Mombasa (Thomas, 1897:435; Neumann, 1900:541; Davis et al, 1968:844, CM, NMK); Mazeras (Heller, 1912:12; Hollister, 1918:29); Gede (Corbet and Neal, 1965:67, Rathbun, 1979:16, NMK); Msa- baha (NMK); Sokoke Forest (CM, NMK); Watamu (NMK); Jilari (Corbet and Neal, 1965:67); Taveta (Thomas, 1910c:309; Corbet and Neal, 1965:68; Corbet and Hanks,
1986 Aggundey and Schlitter — Kenyan Insectivora and M acroscelidea 337
1968:70); Shimba Hills (Heller, 1912:13); Rabai Hills (Thomas, 1897:435); Mrima Hill, 30 mi SW Mombasa (Harmsen and Jabbal, 1968:158^
Genus Rhynchocyon Peters, 1847 Rhynchocyon chrysopygus Guenther, 1881
Rhynchocyon chrysopygusGuQnthQr, 1881. Proc. Zool. Soc. London, p. 164, pi, 14. River Mombaca, Kenya [=Mombasa, Kenya, according to Moreau et aL, 1946:392].
Records. — 'R.vvqt Mombaca (Guenther, 1881:164); Gede (Corbet and Hanks, 1968:65; Rathbun, 1978:11, 1979:16, NMK); Mida (NMK); Msabaha (NMK); Sokoke Forest (Corbet and Hanks, 1968:65; Rathbun, 1978:12, CM, NMK); Takaunga (Corbet and Hanks, 1968:65); Arbagundi, Golana River (Corbet and Hanks, 1968:65); Malindi (Cor- bet and Hanks, 1968:65).
Remarks.-— yioYQ2iU et al. (1946:392) restricted Guenther’s type lo- cality of River Mombaca to Mombasa, Kenya. Corbet and Hanks (1968:65) further qualify the type locality by suggesting a rather vague interpretation of Mombasa as the type locality. R. chrysopygus is pres- ently known only from north of Mombasa. Rathbun (1979:9) reports observations at Kombeni River and Boni Forest.
Rhynchocyon petersi petersi Bocage, 1880
Rhynchocyon petersi Bocage, 1880. J. Sci. Math., Phys. Nat., Lisboa, (1)7:159, pi. 4, f. 2 (“Envoye de Zanzibar,” restricted to mainland of East Africa, region opposite of Zanzibar, by Dollman, 1912:131).
Records. — MKZQTSiS (Hollister, 1918:28); Rabai Hills (Corbet and Hanks, 1968:64); Shimba Hills (Corbet and Hanks, 1968:64).
Genus Elephantulus Thomas and Schwann, 1906 Elephantulus {Elephantulus) rufescens (Peters, 1878)
Macroscelides rufescens Peters, 1878. Monatsb. K. Preuss. Akad. Wiss,, Berlin, p. 198, pi. 1, f. 3. Ndi, Kenya.
Macroscelides boranosThomdiS, 1900. Proc. Zool. Soc. London, 1900:802. Mega, western Boran Galla, southeast of Lake Rudolf, Kenya.
Elephantulus dundasi Dollman, 1910. Ann, Mag. Nat. Hist., (8)5:95, Jan, Harich, near Lake Baringo, 3000 ft, Kenya.
Elephantulus phaeus Heller, 1910. Smithsonian Misc. Coll., 56(1 5):8, 23 Dec. Njoro O Solali, Sotik Dist., Kenya.
Elephantulus delicatus Dollman, 1911. Ann. Mag. Nat. Hist., (8)8:652. Orr Valley, Mt. Nyiro, Kenya.
Elephantulus puicher rendilis Lonnberg, 1912. Kungl. Svenska Vet.-Akad. Hand!., (2)48(5):
49, 26 June. Thera, below Chanler Falls, Northern Guaso Nyiro, Kenya. Elephantulus rufescens mariakanae Heller, 1912. Smithsonian Misc. Coll., 60(1 2): 10, 4 Nov. Mariakani, Kenya.
Records.— ^di (Peters, 1878:199); Mega (Thomas, 1900:803); Njoro O Solali, Sotik (Heller, 1910Z?:8; Hollister, 1918:33); Harich (Dollman, 1910:96); Orr Valley, Mt. Nyiro
(Dollman, 191 1:653; Hollister, 1918:36); Thera (Lonnberg, 1912Zj:51); Mariakani (Hel- ler, 1912:10; Hollister, 1918:33); Mtito Andei (Hollister, 1918:33); Voi (Hollister, 1918: 33; Allen and Lawrence, 1936:40; Corbet and Hanks, 1968:86, NMK); Kabalolot Hill, Sotik (Hollister, 1918:33); Lime Springs, Sotik (Hollister, 1918:33); Loita Plains (Hoi-
338
Annals of Carnegie Museum
VOL. 55
lister, 1918:33, NMK); Southern Guaso Nyiro (Hollister, 1918:35); Telik River, Sotik (Hollister, 1918:35); North Loroghi (Hollister, 191 8:35); NyamaNyango (Hollister, 1918: 35); Northern Guaso Nyiro River (Hollister, 1918:36); Longaya Water, Marsabit Road (Hollister, 1918:36); Archer’s Post (Corbetand Hanks, 1968:86); Taveta (Thomas, 1910c: 309; Corbet and Hanks, 1968:86); ^thangaini (NMK); Kilungu (NMK); Limoni (NMK); Ngari Nyiro (NMK); Southern Kidong (NMK); Lemek (NMK); 1 1 mi N Entesekera (NMK); Emali (NMK); Sultan Hamud (NMK); Samburu (NMK); Lokori (NMK); Ka- ruiru (NMK); Tam Desert (NMK); Golbanti (NMK); Kampi ya Samaki (NMK); Kan= jangareng(NMK); Mt. Mbololo (Allen and Lawrence, 1936:40); Lodwar (St. Leger, 1937: 525); Wenje (Percy et ah, 1 953a: 1 16, 1 18); River Kerio Suk (Ruxton, 1926:29); Baringo (Thomas, 1910c:310); Zuwani Swamp (Dollman, 1914a:88); Nanyuki (Southern and Hook, 1963:51 1); 12 mi NW Kerio River (Dollman 1914Z?:309); Kerio River (Lonnberg, 1918:175); below Chanler’s Falls (Lonnberg, 1912^:51); Bushwackers (Rathbun, 1979: 16).
Elephantulus (Nasilio) bmchyrhynchus (A. Smith, 1836)
Macroscelides brachyrhynchus A. Smith, 1836. Report of the Expedition for Exploring Central Africa, p. 42. Country between Lake Lakatoo and the Tropic. Macroscelides delamerei Thomas, 1901. Ann. Mag. Nat. Hist., (7)8:155. Athi River, 6000 ft, Kenya.
Nasilio brachyrhynchus albiventer Os%oo(X, 1910. Publ. Field Mus. Nat. Hist., Zool. Ser., 10(2): 13. Lake Elementeita, Kenya.
Records. ~ Axhi River (Thomas, 1901:155); Engare Narok River (Hollister, 1918:31, NMK); Loita Plains (Hollister, 1918:31, NMK); Southern Guaso Nyiro (Hollister, 1918: 31); Ulukenia Hills (Hollister, 1918:31); Bargunett River (Hollister, 1918:31); Engare Ndare River (Hollister, 1918:31); Lesiweru River, Mem Road (Hollister, 1918:31); Nai- vasha Station (Hollister, 1918:31, NMK); Nyuki River (Hollister, 1918:31); Olorgesailie (Toschi, 1949:27, NMK); Lemik Valley (NMK); Rumumti (NMK); Wame Hill, Konza (NMK); Amala River (NMK); Voi (Allen and Lawrence, 1936:40); Narrosurra River (Kollmann, 1914:319); Suswa (Kollmann, 1914:319); Guasso Nyero (Kollmann, 1914: 319); Lengototo (Dollman, 1914^:309); Lake Elementeita (Osgood, 19 10a: 13).
Gazetteer
Locality names are listed in alphabetical order with variant names cross-referenced to the standard names. Standard names are taken from the second edition of the official standard names gazetteer for Kenya published in 1978 and approved by the United States Board on Geographic Names. Most of the entities can be identified and located on the 1978 version of the Kenya and Northern Tanzania Route Map published in English, French, and German by the Survey of Kenya.
Coordinates for locality names were taken mostly from the Kenyan gazetteer listed above. In addition, Loveridge (1937), Moreau et al. (1946), Chapin (1954), and Davis and Misonne (1964) were consulted together with place modifiers in the original refer- ences for published records. In the case of rivers, when no place modifiers were available for the published records or on the specimen labels, coordinates are given for the river mouth or confluence.
In a number of instances, more than one entity exists in Kenya for a place name. This generally does not cause a real problem but does in the case of the locality cited at Ewaso Ngiro and its variants, especially the older specimens labeled Guasso Nyiro. In this latter instance, we have given coordinates for both the southern and northern Ewaso Ngiro rivers in the gazetteer.
Aberdare Mountains 0°25'S, 36°38'E
Aberdare Range 0°25'S, 36°38'E
1986 Aggundey and Schlitter — Kenyan Insecti vora and M acroscelidea 339
|
Amala River |
r02'S, 35N4'E |
|
Amboseli |
2°40'S, 37N7T |
|
Arbagundi Archer’s Post |
0°39'N, 37“41T |
|
Athi River |
r27'S, 36®59'E |
|
Bargunett River [=Burguret River] |
0°01'S, 36°56'E |
|
Baringo [=Mukiitan] |
0°38'N, 36°16'E |
|
Blue Post Burguret River |
0°0rS, 36°56'E |
|
Busia |
0°28'N, 34®06'E |
|
Chanler’s Falls |
0°47'N, 38®05'E |
|
Changamwe |
4‘’0rS, 39°38'E |
|
Chania River |
r02'S, 37W'E |
|
Cherangani Hills |
riS'N, 35°27'E |
|
Chyulu Hills |
2°35'S, 37°50'E |
|
Donya Sabuk [=ol Doinyo Sapuk] |
r06'S, 37N5'E |
|
Eldoret |
0°3rN, 35°17'E |
|
Elgeyo Forest |
0°46'N, 35°3rE |
|
|
2®05'S, 37®28'E |
|
Embu |
0°32'S, 37“27'E |
|
Engare Nanyuki |
0°2FN, 36®55'E |
|
Engare Narok |
r09'N, 36®35'E |
|
Engare Ndare River [=Engare Ondare] |
0“35'N, 37®23'E |
|
Engare Ondare |
0°35'N, 3T23'E |
|
Eusso Nyiro Post [=Archer’s Post] |
0‘’39'N, 37®4rE |
|
Entasekera |
rsrs, 35®5rE |
|
Entesekera [= Entasekera] |
rSl'S, 35“5rE |
|
Ewaso Ngiro (Northern) |
0°37'N, 36"55'E~ |
|
Ewaso Ngiro (Southern) |
0°28'N, 39°55'E 0°35'S, 35M7'E== |
|
Fort Hall [=Muranga] |
2W'S, 36®07'E 0°43'S, 37“09'E |
|
Cede |
3H8'S, 40°0rE |
|
Golbanti |
2®27'S, 40®12'E |
|
Guasso Nyero [=Ewaso Ngiro] Guaso Nyiro River, Sotik District [=Ewaso Ngiro (Southern)] Harich [=Marich] |
1®32'N, 35®27'E |
|
Horr Valley |
2®10'N, 36“55'E |
|
Ijara |
r36'S, 40®3FE |
|
Ilkaputiei |
r38'S, 37®00'E |
|
Isiola River [=Isiolo River] |
0®34'N, 37°35'E |
|
Isiolo River |
0°34'N, 37°35'E |
|
Itiolu River [=Isiolo River] |
0‘’34'N, 37“35'E |
|
Jilari [=Jilore] |
3°1FS, 39“54'E |
|
Jilore |
3nrS, 39°54'E |
|
Jombeni [=Nyambeni] |
0°13'N, 57‘’52'E |
|
Jombeni Range [=Nyambeni Range] |
0°20'N, 37®57'E |
|
Juja Farm |
TIPS, 37°07'E |
|
Kabalolot Hill |
ca. rOO'S, 35°23'E |
|
Kabete |
ri6'S, 36°43'E |
|
Kagio |
0°40'S, 37N3'E |
|
Kaimoni [=Kaumoni] |
r44'S, 37“35'E |
|
Kaimosi |
OW'N, 34®5rE |
|
340 Annals of Carnegie Museum |
VOL. 55 |
|
|
Kajiado |
1°5LS, |
36®47'E |
|
Kakamega |
0°17'N, |
34°45'E |
|
Kakuma |
3°43'N, |
34®52'E |
|
Kampi ya Samaki |
0°36'N, |
36®01'E |
|
Kanyakeni [=Kanyekine] |
0°08'S, |
37040'E |
|
Kanyangareng |
1°47'N, |
35'08'E |
|
Kanyekine |
0°08'S, |
37°40'E |
|
Kapiti Plains [=Ilkaputiei] |
1°38'S, |
37“00'E |
|
Kapsabet |
0°12'N, |
35®06'E |
|
Karati |
0‘’26'S, |
37°27'E |
|
Karuiro |
0°37'S, |
37°07'E |
|
Karuiru [= Karuiro] |
0°37'S, |
37°07'E |
|
Kasigau |
3°50'S, |
38°40'E |
|
Kathera |
0°03'S, |
37°35'E |
|
Kaumoni |
1°44'S, |
37°35'E |
|
Kazere [= Kathera] |
0°03'S, |
37°35'E |
|
Kenna [=Kinna] |
0°19'N, |
38°12'E |
|
Kericho |
0°22'S, |
35°17'E |
|
Kerio River |
2°59'N, |
36°34'E |
|
Kibabe [=Kibabet] |
0°11'N, |
35°15'E |
|
Kibabet |
O^ll'N, |
35°15'E |
|
Kijabe |
0°56'S, |
36°34'E |
|
Kikuyu |
1°15'S, |
34°40'E |
|
Kilungu |
R48'S, |
37°22'E |
|
Kinangop |
0°44'S, |
36°40'E |
|
Kinna |
0°19'N, |
38°12'E |
|
Kirui, Mt. Elgon [=Kirui’s] |
0°50'N, |
34M0'E |
|
Kirui’s |
0°50'N, |
34°40'E |
|
Kisumu |
0°06'S, |
34°45'E |
|
Kithangaini |
1°29'S, |
37°23'E |
|
Kitui |
1°22'S, |
38°01'E |
|
Koliokwell River |
||
|
Kutu |
m o O |
37°19'E |
|
Lagari [=Lugari] |
0°39'N, |
34°53'E |
|
Laikipia |
0°25'N, |
36°45'E |
|
Laikipia Plateau |
0°25'N, |
36°08'E |
|
Lake Elementeita [=Lake Elmenteita] |
0'’27'S, |
36°15'E |
|
Lake Elmenteita |
0°27'S, |
36“15'E |
|
Lake Ilpolosat |
0°09'S, |
36°26'E |
|
Lake Nakuru |
0°22'S, |
36°05'E |
|
Lake Olbollosat [=Lake Ilpolosat] |
0°09'S, |
36®26'E |
|
Lake Olbollossat [=Lake Ilpolosat] |
0“09'S, |
36®26'E |
|
Lake Sergoi |
0°42'N, |
35°25'E |
|
Lakiundu River [=Ngaramara River] |
0°36'N, |
37°37'E |
|
Lamu |
2°16'S, |
40“54'E |
|
Lemek |
1°06'S, |
35°23'E |
|
Lemek Valley |
ro9's, |
35®19'E |
|
Lemik Valley [=Lemek Valley] |
1°09'S, |
35°19'E |
|
Lengototo [=Lenkutoto] |
1°39'S, |
35°58'E |
|
Lenkutoto |
r39'S, |
35‘’58'E |
|
Lesiweru River |
||
|
Lime Springs [=Maji Moto] |
1°20'S, |
35°42'E |
|
Lodwar |
3°07'N, |
35°36'E |
|
1986 Aggundey and Schlitter — Kenyan Insecti vora |
AND Macroscelidea 341 |
|
|
Loita |
r30'S, |
35®4rE |
|
Loita Plains |
r20'S, |
35°32'E |
|
Lokichokio |
4®2rN, |
34“2rE |
|
Lokori |
r57'N, |
36®0rE |
|
Londiani |
OHO'S, |
35°36'E |
|
Longaya Water |
ca. r07'N, |
37®38T |
|
Lorogi |
roo'N, |
36“5rE |
|
Luazomela River |
0°29'N, |
37®40'E |
|
Lugari |
0°39'N, |
34°53'E |
|
Lukenya |
r3rs, |
36"58'E |
|
Lukenya Hills |
r28'S, |
37°03'E |
|
Lukenya Mountain [=Lukenya Hills] |
r28'S, |
37°03'E |
|
Lukosa River |
0°12'N, |
34°56'E |
|
Machakos |
rsrs, |
37H6'E |
|
Maji Moto |
r20'S, |
35"42'E |
|
Malindi |
40“07'E |
|
|
Mariakani |
3°52'S, |
39“28'E |
|
Marich |
r32'N, |
35®27'E |
|
Marsabit |
2°20'N, |
37°59'E |
|
Mau Forest |
0°20'SA)M0'S, |
|
|
35°25'E- |
G6°05'E |
|
|
Maua |
0°14'N, |
37°56'E |
|
Mayo River |
OHO'S, |
37^0 1'E |
|
Mazeras |
BOSS'S, |
39033, E |
|
Mega |
||
|
Merifano |
2H9'S, |
40®08'E |
|
Meru |
0°03'N, |
37"39'E |
|
Mianzini |
ca. 0“55'S, |
36®25'E |
|
Mida |
3H9% |
39®58'E |
|
Molo |
OHS'S, |
35°44'E |
|
Mombasa |
4“03'S, |
39M0'E |
|
Mt. Elgon |
r08'N, |
34°33'E |
|
Mt. Garguez [=Warges] |
0°57'N, |
37®24'E |
|
Mount Lololokwi [=01 Doinyo Sabachi] |
0«50'N, |
37°32'E |
|
Mt. Kenia [=Mt. Kenya] |
OHO'S, |
37°20'E |
|
Mt» Kenya |
OHO'S, |
37“20'E |
|
Mt. Mbololo |
3®17'S, |
38°28'E |
|
Mt. Nyiro [=0 1 Doinyo Ngiro] |
2°08'N, |
36"5rE |
|
Mt. Sagalla |
3“27'S, |
38°35'E |
|
Mt. Umengo |
ca. 3“18'S, |
38°19'E |
|
Moyale |
3"32'N, |
39°03'E |
|
Mrima Hill |
4‘»29'S, |
39H6'E |
|
Msabaha |
3H6'S, |
40°03'E |
|
Mtito Andei |
2°4rs, |
38H0T |
|
Muguga |
ril'S, |
36°39'E |
|
Mukutan |
0°38'N, |
36°16'E |
|
Muranga |
0®43'S, |
37“09'E |
|
Mweru |
0“40'S, |
37°05'E |
|
Nairobi |
ri7'S, |
36®49'E |
|
Naivasha |
0M3'S, |
36°26'E |
|
Naivasha Plains |
OMl'S, |
36°27'E |
|
Naivasha Station |
0°43'S, |
36®26'E |
|
Nakatishu River |
0®33'S, |
36°38'E |
|
342 Annals of Carnegie Museum |
VOL. 55 |
|
|
Nandi [=Kapsabet] |
0°12'N, |
35°06'E |
|
Nanyuki |
0°0LN, |
37°04'E |
|
Naro Moru |
OHIO'S, |
37°0rE |
|
Narosura River |
1°33'S, |
35°53'E |
|
Narrosurra River [=Narosura River] |
1°33'S, |
35°53'E |
|
Ndi |
3®14'S, |
38°30'E |
|
Neumann’s Boma [=Samburu Game Lodge] |
0°34'N, |
37035, E |
|
Ngari Nyiro [=Ewaso Ngiro] |
||
|
Ngaramara River |
0°36'N, |
37°37'E |
|
Ngare Nocbor |
ca. 2°45'N, |
36°45'E |
|
Ngatana |
2°13'S, |
40°11'E |
|
Ngong |
1°22'S, |
36°39'E |
|
Njoro O Solali |
0°28'S, |
35°04'E |
|
North Laroghi [=Lorogi] |
FOO'N, |
36°51'E |
|
Northern Guaso Nyiro [=Ewaso Ngiro] |
0®37'N, 36°55'E~ |
|
|
0°28'N, |
39°55'E |
|
|
Nyahururu |
0°02'N, |
36°22'E |
|
Nyama Nyango [=Samburu Game Lodge] |
0°34'N, |
37°35'E |
|
Nyambeni |
0°13'N, |
37°52'E |
|
Nyambeni Range |
0°20'N, |
37°57'E |
|
Nyeri |
0°25'S, |
36°57'E |
|
Nyiru [=01 Doinyo Ngiro] |
2°08'N, |
36°51'E |
|
Nyuki River [=Engare Nanyuki] |
0°21'N, |
36°55'E |
|
Nzoia River |
0°03'N, |
33°57'E |
|
01 Arabel |
0“18'N, |
36°18'E |
|
01 Doinyo Ngiro |
2®08'N, |
36°51'E |
|
01 Doinyo Sabachi |
0®50'N, |
37°32'E |
|
01 Doinyo Sapuk |
ro8's, |
37°15'E |
|
Olgerei River |
r43'S, |
35°18'E |
|
Olijoro O Nyon River |
ca. 0°57'S, |
35°55'E |
|
Olorgasailie |
r34'S, |
36°27'E |
|
Orr Valley [=Horr Valley] |
2°10'N, |
36°55'E |
|
Peccatoni |
2°25'S, |
40°43'E |
|
Potha |
r34'S, |
37°10'E |
|
Rabai Hills |
||
|
Ravine Station |
0°01'N, |
35°43'E |
|
River Kerio Suk |
2‘’59'N, |
36°07'E |
|
River Mombaca [=Mombasa] |
4®03'S, |
39°40'E |
|
Rojewero Plains |
0°il'N, |
38°10'E |
|
Rumruti [=Rumuruti] |
onh'N, |
36°32'E |
|
Rumuruti |
0“16'N, |
36®32'E |
|
Samburu |
3M6'S, |
39°17'E |
|
Samburu Game Lodge |
0®34'N, |
37°35'E |
|
Selengai |
2®irN, |
37°10'E |
|
Sera |
LOl'N, |
37°53'E |
|
Sergoi |
0°39'N, |
35°23'E |
|
Sergoit Lake [=Lake Sergoi] |
0M2'N, |
35°25'E |
|
Shimba Hills |
4®13'S, |
39°25'E |
|
Sirgoit [= Sergoi] |
0®39'N, |
35°23'E |
|
Sirgoit Lake [=Lake Sergoi] |
0“42'N, |
35°25'E |
|
Sokoke Forest |
3‘’29'S, |
39°50'E |
|
Solai |
o°orN, |
36®09'E |
|
Southern Guaso Nyiro [=Ewaso Ngiro] |
2®04'S, |
36'’07'E |
|
1986 Aggundey and Schlitter— Kenyan Insectivora and Macroscelidea 343 |
|
|
Southern Kedong Valley |
ca. 1°24'S, 36°27'E |
|
Southern Kidong [= Southern Kedong Valley] |
ca. r24'S, 36®27L |
|
Sultan Hamud |
2°0rS, 37°22'E |
|
Takaungu |
3“4rS, 39®5LE |
|
Talek River |
r26'S, 35W'E |
|
Tara Desert |
3“45'S, 39“08'E |
|
Taveta |
3“24'S, 37°4rE |
|
Teiek River [=Talek River] |
r26'S, 35®04'E |
|
Thera [=Sera] |
rOl'N, 37®53'E |
|
Thika |
r03'S, 37°05'E |
|
Thomson's Falls [=Nyahuraru] |
0°02'N, 36“22'E |
|
Tsavo |
2®59'S, 38°28'E |
|
Tsavo River |
2°59'S, 38“3rE |
|
Ulukenia [=Lukenya] |
1°31'S, 36°58'E |
|
Ulukenia Hills [=Lukenya Hills] |
1°28'S, 37°03'E |
|
Upper Nzoia River |
ca. 0®53'N, 35®22'E |
|
Upper Ura River |
ca. 19U0'N, 37®59'E |
|
Voi |
3«23'S, 38®34'E |
|
Wambugu |
0°35'S, 37®02'E |
|
Wame Hill [-Wami Hill] |
r39'S, 37“08T |
|
Wami Hill |
r39'S, 37°08L |
|
Warges |
0°57'N, 37“24'E |
|
Watamu |
3®21'S, 40°0rE |
|
Wema |
2U3'S, 40UrE |
|
Wenje |
1M7'S, 40“06'E |
|
West Slope Mt. Kenya |
OUO'S, 37°10'E |
|
Witu |
2®23'S, 40®26'E |
|
Yala River |
0°04'N, 34®09'E |
|
Ziwani |
3'’23'S, 37'’47'E |
|
Ziwani Swamp |
3°16'S, 37°47'E |
|
Zuwani [=Ziwani] |
3°23'S, 37°47'E |
|
Zuwani Swamp [=Ziwani Swamp] |
3°16'S, 37M7'E |
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— — — . 1910c. List of mammals from Mount Kilimanjaro, obtained by Mr. Robin Kemp, and presented to the British Museum by Mr. C. D. Rudd. Ann. Mag. Nat. Hist, (8)6:308-316.
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True, F. W. 1892. An annotated catalogue of the mammals collected by Dr, W, L.
Abbott in the Kilima-njaro region, East Africa. Proc. U.S. Natl. Mus., 15:445-480. Yates, T. L. 1984. Insectivores, elephant shrews, tree shrews, and dermopterans. Pp. 1 17-144, in Orders and families of Recent mammals of the world (Sydney Anderson and J. Knox Jones, Jr., eds.), John Wiley and Sons, New York, viii + 453 pp.
Back issues of many Annals of Carnegie Museum articles are available, and a few early complete volumes and parts are listed at half price. Orders and inquiries should be addressed to: Publications Secretary, Carnegie Museum, 4400 Forbes Avenue, Pittsburgh, Pa. 15213.
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CARNEGIE MUSEUM OF NATURAE HISTORY
4400 FORBES AVENUE ® PITTSBURGH, PENNSYLVANIA 15213 VOLUME 55 31 DECEMBER 1986 ARTICLE 15
DEVONIAN AND MISSISSIPPIAN CONULARIIDS OF NORTH AMERICA. PART A. GENERAL DESCRIPTION AND CONULARIA
Loren E. Babcock
Rodney M. Feldmann^
Research Associate, Section of Invertebrate Fossils
Abstract
The systematics, morphology and paleoecology of the new phylum Conulariida pro- posed here, from the Devonian and Mississippian rocks of North America are reviewed in this two-part work. Conulariids are identified by an elongate, pyramidal exoskeleton comprising a framework of calcium phosphate rods, with or without spines and nodes, covered by an integument made of thin layers of calcium phosphate and protein. Con- ulariids were gregarious invertebrate animals that were attached to substrata by means of calcium phosphate or chitinous stalks. These animals were exclusively marine and limited to rocks of the Ordovician through the Triassic. Conulariids are found in all types of marine facies and most probably had either a pseudoplanktonic or a benthonic lifestyle.
Three valid genera of conulariids are recognized in the Devonian and Mississippian of North America. These are Conularia Miller in Sowerby, 1821, Paraconularia Sinclair, 1940a and Reticulaconularia Babcock and Feldmann, n. gen. The genus Diconularia Sinclair, 1952 is considered to be a junior synonym of Conularia. Prior to 1986, 69 species-level taxa of conulariids were described from the Devonian and Mississippian of North America. Herein, and in Part B, 28 species are recognized as valid. Eleven species are assigned to Conularia and are described in Part A.
‘ Address: Department of Geology, Kent State University, Kent, Ohio 44242.
2 Present address: Department of Geology, University of Kansas, Lawrence, Kansas 66045.
Submitted 2 June 1986.
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Introduction
Conulariids, members of an extinct phylum of invertebrates, have been described in the literature for a span of nearly 200 years. In that time, well over 400 species, subspecies or varieties, and over 40 genera have been named.
The purpose of this paper, and Part B of the same work, is to present taxonomic and morphologic information obtained from an examina- tion of conulariids from Devonian and Mississippian strata of North America. Approximately 1 2% of all species-level conulariid taxa have been described based upon specimens collected from these rocks. Spec- imens described herein were collected only from the United States and Canada; conulariids are not known to have been collected in Mexico. This work involves: 1 , studies of intraspecific and interspecific variation in morphology, ideally based upon large numbers of specimens; 2, studies of the stratigraphic and geographic distributions of species; and 3, analyses of conulariid anatomy, functional morphology and paleo- ecology, as the fossil record permits. Each taxon identified from De- vonian or Mississippian rocks of North America is accompanied by a new, or in some cases, the first, illustration, and a description high- lighting points of morphology now considered of greatest taxonomic value.
In the course of examining the Devonian and Mississippian forms, it became clear that it was necessary to revise the terminology related to morphology. In so doing, it was considered essential that conulariids from a much wider range of geological and stratigraphical occurrences had to be examined than those treated systematically herein. Thus, the detailed morphology and terms are intended to be applicable to all organisms referable to the Conulariida.
Part A of this work comprises a general description of the hard- and soft-part morphology of conulariids, occurrences and paleoecology of Devonian and Mississippian taxa of North America, a summary of the taxa treated both in this paper and in Part B, a key to the Devonian and Mississippian conulariid taxa of North America, and descriptions of species of that group which are referred to the genus Conuiaria. Part B of this work contains descriptions of species referable to the genera Paraconularia and Reticulaconularia n. gen. and specimens described in the literature as conulariids but which are here rejected from the phylum. Locality descriptions and measurements of selected specimens are included as appendices to Part B. Figures are numbered consecu- tively in both Parts A and B in order to avoid cross-reference confusion.
Morphology
General. --When preserved in three dimensions, the exoskeleton of a conulariid generally has a four sided, bilaterally symmetrical, elongate
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pyramidal shape (Fig, 1.1). The profile may be modified by the de- velopment of one or more exoskeletal constrictions (Fig. LI). The exoskeleton, as preserved, generally ranges from 2 to 10 cm in length in full grown individuals. In a few species, however, the exoskeleton may attain a length in excess of 20 cm (Fletcher, 1938; Lamont, 1946; Sinclair, 1948; herein, Fig. 1 1.2). In nearly all instances, the conulariid exoskeleton diverges in width gradually and uniformly from a closed apical end to an open apertural end. The apical end may be closed either by a (morphological) apex (Fig. 1.1) or by an apical wall (Fig. 1.2). Presumed soft-parts consist of a single tubular structure that runs internally along the length of the exoskeleton and at least one globular body (Fig. 2.1).
Morphologic terms. --Tht literature on conulariid morphology in- cludes important review papers by Slater (1907), Boucek and Ulrich (1929), Kiderlen (1937), Richardson (1942), Sinclair (1948, 1952), Moore and Flarrington (1956a, 1956/?) and Babcock and Feldmann (1986). Much misunderstanding of conulariid morphology has arisen from terms that have been improperly defined or undefined, from terms that are ambiguous and from terms that imply systematic affinities. Babcock and Feldmann (1986) proposed a set of morphological terms for conulariids that described morphological features without intro- ducing unnecessary connotations of genetic affinities. Below is a list of morphological terms applied to conulariids, modified from Babcock and Feldmann (1986), together with terms here considered synony- mous and which relate to the morphology of members of the phylum but are inappropriate in the light of advances made during this study.
AD APERTURAL SPINE —long spine projecting from near the adapertural side of a rod, in the direction of the aperture.
AD APICAL SPINE— short spine projecting from near the internal adapical side of a rod, in the direction of the apex.
ALIMENTARY TRACT— narrow, elongate, essentially tubular soft-part structure run- ning the length of the central cavity. Synonyms: axial element, intestine.
ANGULATED CIRCULAR CURVE — style of rod articulation in which two abutting rods on a face form a broad arcuate, adapically concave ridge, interrupted by a slight adapertural point at the midline, and by gentle adapertural turns in the vicinity of the facial margins.
APERTURAL CONSTRICTION— exoskeletal constriction located nearest the aper- ture. Synonym: wrinkle.
APERTURE— opening at widest end of exoskeleton. Synonyms: base, mouth, opening.
APERTURAL TERMINATION— rounded or bluntly subtriangular extension of exo- skeleton on each face at widest end of exoskeleton.
APEX (MORPHOLOGICAL APEX)— narrowest termination of exoskeleton, where the four faces Join at a closed point. Synonyms: (biological) apex, summit. Compare with hypothetical apex.
APICAL ANGLE— hypothetical angle formed by one face of the exoskeleton; measured at the intersection of two lines each identified by tracing positions on the exoskeleton
exoskeleton
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aperture
ridge
interridge area midline
corner groove corner angle
^^>«exoskeletal constriction
rod angle
face
apical wall apical angle
hypothetical apex
Fig. 1.— Conulariid morphology, exhibited by a generalized Pamconularia. 1.1; exo- skeleton with stalk attached. 1.2; apical region with stalk removed. Morphological terms are explained in the text.
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Fig. 2.~Conulariid morphology, exhibited by a generalized Paraconularia. 2.1; cutaway view of exoskeleton showing internal soft-parts. Structure of the soft-parts in the apertural region is problematic. 2.2; detailed view of two rods. Morphological terms are explained in the text.
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tangential to the facial margins and defining the maximum angle of separation. Synonym: facial angle. See major apical angle and minor apical angle.
APICAL WALL —broadly rounded, adapically convex, portion of integument lacking rods which completely covers the apical end of the exoskeleton when the apex itself is missing. Synonyms: apical septum, apical diaphragm, basal limitation, diaphragm, internal partition, Schott, septum.
CENTRAL CAVITY — region located internal to the four faces of the exoskeleton. Syn- onym: body cavity.
CORNER ANGLE— longitudinal line in the marginal region of a face connecting points of greatest inflection of the rods. Synonym: shoulder.
CORNER GROOVE — longitudinal invagination of exoskeleton connecting points where pairs of rods from adjacent faces cross near the marginal terminations of those rods. Synonyms: angular furrow, articulating suture, edge, lateral channel, longitudinal channel, marginal furrow, marginal groove, side furrow.
EXOSKELETAL CONSTRICTION— depression, restricted in the longitudinal direc- tion, traceable on all four faces of the exoskeleton in the same relative position. Compare with apertural constriction.
EXOSKELETON (SKELETON)— four sided pyramidal structure, open at the widest end and closed at the narrowest end, comprising rods joined by integument. Syn- onyms: periderm, pyramid, shell, test.
FACE— one of four sides of the exoskeleton crossed by ridges; it is delimited by the aperture, by the apex or the apertural wall and by two comer grooves. Synonyms: side, surface, wall. See major face and minor face.
GLOBULAR BODY— large internal soft-part stmcture, subovoid in outline, located near the aperture. Synonym: esophagus.
GOTHIC ARCH — style of rod articulation in which two adjacent rods on a face form ridges that meet at an obtuse, adapically concave angle at the midline and proceed away from the midline along lines subtly curved adapically.
HYPOTHETICAL APEX— point in space where two lines, traced along the mean di- rection of the comer angles; meet; the hypothetical apex may or may not coincide with the position of the (morphological) apex.
INFLECTED CIRCULAR CURVE — style of rod articulation in which two adjacent rods on a face form a broadly arcuate, adapically concave ridge except in the vicinity of the facial margins, where they turn gently adaperturally.
INFLECTED GOTHIC ARCH— style of rod articulation in which two adjacent rods on a face form ridges that meet at an obtuse, adapically concave angle at the midline and proceed away from the midline along lines subtly curved adapically except in the vicinity of the facial margins, where they turn gently adaperturally.
INTEGUMENT— multilayered, presumably flexible, stmcture composed of calcium phosphate and protein, within which rods and spines were embedded and held in position. Synonyms: periderm, test.
INTERRIDGE AREA— roughly transverse band of integument located between two facial ridges. Synonyms: intercostal space, interspace, transverse furrow, transverse sulcus, space.
INTERRIDGE CREST —raised area, usually a linear ridge, located in an interridge area and positioned at a right angle to a ridge; formed by integument covering an ad- apertural or adapical spine. Synonyms: bar, intercostal longitudinal striation, vertical striation, longitudinal bar, longitudinal striation.
INTERRIDGE FURROW— low area, usually linear, located in an interridge area, and between two interridge crests.
INTERROD AREA— open region located between two rods; exposed only when integ- ument is absent.
MAJOR APICAL ANGLE— apical angle subtended by a major face.
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MAJOR FACE —wider of two adjacent faces.
MARGIN (FACIAL MARGIN)— longitudinal edge of a face, or a line connecting points where two faces meet in a comer groove.
MIDLINE— longitudinal line connecting points where either two adjacent rods on a face meet, or central to the facial terminations of each pair of adjacent rods if the rods do not meet. The midline can be expressed as either a thin groove or a raised line if the integument is preserved. The midline seems to be pigmented in some speci- mens. Synonyms: central face furrow, central facial groove, facial groove, facial midline, longitudinal carina, median groove, median line, mesial furrow, mid-line, middle line, parietal line, septum, stmctural channel.
MINOR APICAL ANGLE— apical angle subtended by a minor face.
MINOR FACE— narrower of two adjacent faces.
NODE— minute, subcircular, raised surface on a rod or ridge. Synonyms: granule, papilla, tubercle, pustule, wart.
RIDGE (FACIAL RIDGE)— raised line crossing a face from a comer groove to the midline area, and formed by integument covering a rod. Synonyms: costa, crenu- lation, ornamental rib, plica, plication, rib, riblet, transverse line, transverse rib, transverse ridge, transverse striation.
ROD —narrow, elongate stmcture that is subcircular in cross section, composed of cal- cium phosphate, and embedded within the integument; it is thickened near the marginal termination, and tapers very gradually to a blunt point at the facial ter- mination.
ROD ANGLE— angle subtended by a line connecting the two most distant points of a rod along a longitudinal line and a line constmcted perpendicular to the facial margin at the point where that ridge intersects the comer angle. Synonym: angle at the mid- line.
ROD PAIR— two rods on a face whose distal ends meet, or approach closely, at the midline.
SKELETON— See exoskeleton.
SPINE— solid, narrow, short or elongate stmcture, projecting from, and whose axis is at a right angle to, a rod; tapers gradually to a sharp point distally. See adapical spine and adapertural spine.
STALK— elongate stmcture, possibly chitinous, phosphatic, or chitinophosphatic, which articulates proximally with a conulariid apex; distally, the stmcture seems to attach to a substratum of uncertain nature.
Abandoned morphological terms. ~T\iQ following terms, previously used in connection with the description of conulariids, are here con- sidered inappropriate for various reasons including: 1, the structures have been shown to be taphonomic in origin; 2, the structures were described from organisms which should be excluded from the phylum Conulariida; 3, the structures have been shown to be absent in conu- lariids; or 4, the structures were described from dubious fossil material.
ANUS.
APPENDIX.
APERTURAL LOBE, Synonyms: apertural flap, apeitural lip, flap, laterales, lobe, lip, mouth flap.
ATTACHMENT DISC.
BODY WALL.
EYE LENS.
HINGE.
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FEATHER EDGE.
KEEL.
LATERAL LINE. Synonym: longitudinal line, line near midline.
MOUTH PANEL.
NERVE CENTER.
NET-SHAPED EPITHELIUM.
ORNAMENTATION, Synonym: sculpture.
PLATE. Synonym: transverse plate,
SEPTUM: Synonyms: carina, internal longitudinal rib, internal rib, internal rod, internal septum, longitudinal septum,
SKIN-MUSCLE LINING.
URULA.
WALL OPENING,
Aperture. --ThQ aperture (Fig. LI) appears to have been a simple opening at the larger termination of the pyramidal exoskeleton. This region has been the subject of much speculation. ‘"Flaps” or “lappets,” partially or wholly closing the apertural region, were first described by Miller {in Sowerby, 182 IX and subsequently by Etheridge (1901), Las- eron (1912), Richter and Richter ( 1 930), Reed (1933), Kowalski (1935), Kiderlen (1937), Sinclair (1948), Moore and Harrington (1956Z?), Bran- isa (1965) and others. Kiderlen (1937), Termier and Termier (1949) and Moore and Harrington (1956fi) proposed elaborate mechanisms for closure of the apertural region involving the infolding of a flexible exoskeleton. They assumed the line of flexure to be a straight line normal to the midline. In one mechanism of closure, the exoskeleton not only folded along a straight line perpendicular to a face, but also collapsed like a bellows in the vicinity of the comer angles (Moore and Harrington, 1956/?, Fig. 43.1), Moore and Harrington (1956/?, p. F57) suggested that, in order for a conulariid to have been so flexible in the apertural region, the line of flexure at the base of each “apertural flap” was chitinophosphatic, while the remainder of the exoskeleton was phosphatic. No chemical data were presented in support of this hy- pothesis.
Specimens exhibiting closed or partially restricted apertures are com- mon. Over 200 such specimens were observed in the course of this study (for example, Figures 8.1, 10.1, 28.2). Among these, there is no evidence of a consistent line of flexure (for example, Fig. 7. 1-7.2, 8.7=- 8.9, 10.2, 18.2, 22.2-22.3). Typically, the line along which a flap is developed is not straight. Instead, the line of flexure often mimics the style of rod articulation or seems to be arbitrary. No two adjacent faces on the same specimen necessarily fold inward at the same position. Within the same species, there is no consistency from individual to individual either in the placement of a line of flexure or in the mode of closure, as defined by Kiderlen (1937), Termier and Termier (1949) or Moore and Harrington (1956/?). Furthermore, “apertural flaps” have
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been observed at multiple sites on single conulariid exoskeletons (Bab- cock and Feldmann, 1986, fig. IE). Their presence at various places on the exoskeleton of different specimens indicates that they are taph- onomic phenomena. Richter and Richter (1930), noted the extreme flexibility of the holotype of Conularia tulipa from the Hunsruckshiefer (Lower Devonian) of Germany. In their opinion, the conulariid exo- skeleton was flexible enough to have collapsed under its own weight. Infoldings of exoskeleton commonly found in the apertural region, and less commonly elsewhere on a conulariid, are probably taphonomic structures resulting from collapse of the exoskeleton after death. The exoskeleton may not be quite as weak as suggested by Richter and Richter, but it is certainly not as rigid as a mollusk shell.
Apex.— T\iQ apical end of a conulariid may be: 1, a narrow, blunt point (Fig. 1.1); 2, truncated (Fig. 7.6); or 3, truncated, but having the end covered by a smooth, convex, imperforate apical wall (Fig. 1.2). The apex of a conulariid has been interpreted as a sharp point, as a bluntly rounded structure, as a smooth, imperforate wall, or as a smooth wall with a centrally located hole. Since the work of Kiderlen (1937), conulariids have been thought of as metazoans having a sharp point in the juvenile state. Presumably, the point was attached by an attach- ment disc to a hard substratum. Support for Kiderlen’s argument was provided by supposed conulariids which were previously described by Ruedemann (1896i2, 1896/?). The specimens described by Ruedemann seem to be tubes of Sphenothallus (Feldmann et al., 1986). Sphenoth- alius has recently been shown to be unrelated to conulariids (Mason and Yochelson, 1985).
Apices of conulariids are exceedingly rare. The apices are not pointed as once interpreted, but are slightly rounded (Figs. 7.6, 16.6, 33. l- 33.2). During life, most or all conulariids were attached by an elongate stalk (Figs. 1.1, 24.1-24.2, 32.5) to some substratum, during at least part of the life cycle. The apex was sheathed by the proximal portion of the stalk. Breakage at the proximal end of the stalk may possibly explain why so few conulariids are observed that have their apices intact.
Authentic conulariids with attachment discs have never been de- scribed. Small, round, black, and presumably chitinous or chitino- phosphatic, bodies attached to bryozoans or brachiopods have been identified in various museum collections as conulariid attachment discs. Often, the presumed base of a tube is preserved connected to such a structure. Such tubes are circular or subcircular in cross section. These problematic fossils probably represent attachment devices of some type of organism, but a relationship to conulariids has not been demon- strated.
Smooth, imperforate apical walls have been noted by many authors
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(Miller in Sowerby, 1821; Hall, 1876, 1879; Slater, 1907; Richter and Richter, 1930; Sinclair, 1948; Moore and Harrington, \956b\ Babcock and Feldmann, 1984, 1986, herein. Figs. 4.1, 10.4, 11.2, 14.1, 14.4, 1 5.4, 19.1). These structures have been termed septa, apical diaphragms or Schotten. None of these terms seems appropriate, either because of its genetic implications or because of an incomplete description of the morphology. Apical walls are single units of integument that cover the convergent end of a conulariid close to the apical terminus. An apical wall is not located at the apical terminus itself, but seems to be attached to the interior of the faces on the exoskeleton, slightly adaperturally of this region. The line of juncture of the apical wall with the remainder of the integument always seems to occur near the narrowest portion of an exoskeletal constriction. Apical walls appear to lack support from rods or other structures, and may be bowed slightly in the adapical direction. Multiple apical walls may be present in single individuals (Eichwald, 1860; Steinmann and Doderlein, 1890; Slater, 1907; Sin- clair, 1948).
References to conulariids with centrally perforated apical walls in- clude Slater (1907), Richardson (1942), and Swartz and Richardson (1945). A collapsed specimen of Conularia congregata exhibiting a subcircular structure located centrally on the apical wall is illustrated in Figure 4.1. This subcircular structure is interpreted as having resulted from the compression of a thin, flexible apical wall against the apical portion of some soft-part morphologic feature such as the alimentary tract.
The function of the apical wall may have been to seal off the portion of the central cavity in which the conulariid lived from the stalk and older, unused portions of the body. In specimens preserving apical walls, the most apicad portions of exoskeleton are not smooth (Figs. 1 1.2, 14.1). This indicates that conulariid exoskeletons like these may have been tom from their stalks by current forces. It is also possible that some few conulariids periodically may have shed unused portions of the exoskeleton (Babcock and Feldmann, 1984).
Apical angle. — An apical angle is measured at the intersection of two lines projected by tracing tangent lines along the trend of two adjacent comer grooves (Fig. 1.2). Apical angles are typically in the range of 8° to 26°.
The apical angle, as measured on a large segment of a specimen, may differ, by several degrees, from the apical angle as measured on a small section of the skeleton (see Appendix B in Part B). Small segments, particularly at exoskeletal constrictions and near the apex, usually yield somewhat larger apical angles than generalized apical angles, measured over a large segment of an exoskeleton.
A difference in the acute apical angle between adjacent sides of a
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conulariid, mostly attributed to compression, has been noted by nu- merous authors, including Barrande (1867), Hall (1879), Slater (1907), Boucek (1939), and Sinclair (1948). Studies of compressed and pre- sumably uncompressed materials indicate that opposite sides of a con- ulariid exoskeleton are paired (Babcock and Feldmann, 1984, 1986). In cross section, a conulariid is typically rectangular, if only slightly so (Figs. 30.4, 33.4). Each face subtends an apical angle equal to that of the face opposite it, but different from either adjacent face. This suggests that conulariids are bilaterally symmetrical, rather than tetramerally symmetrical, metazoans (Babcock and Feldmann, 1984, 1986). Rhom- boid-shaped conulariids may exist, but most forms that are thought to have a rhomboidal cross section probably were described from subtly compressed specimens.
Faces. —In most cases, the four faces of the conulariid skeleton are essentially planar (Fig. 1.1). At least two forms, Mesoconularia ca~ huanotensis, from the Devonian of Bolivia (Branisa and Vanek, 1973) and M. solitaria, from the Silurian of Czechoslovakia (Sinclair, 1 948), have faces that are markedly curved. In Anaconularia anomala from the Ordovician of Czechoslovakia, Kiderlen (1937) described a clock- wise torsion of the exoskeleton, up to 40°. However, examination of eleven specimens referable to this taxon (GSC 85063-85073) indicates that these fossils, preserved in quartzite, are not twisted or compressed in a uniform fashion. Therefore, the “torsion” which Kiderlen observed may, in fact, be related to post-mortem diagenetic effects.
The two faces on a conulariid exoskeleton which subtend apical angles equal to each other, but smaller than the remaining two faces, are termed minor faces. Those two faces having larger apical angles are termed major faces.
Integument.— T\iQ thin walls, or faces, of the exoskeleton are made up of a multilayered calcium phosphate and protein integument (Ed Landing, personal communication, 1984; based upon electron micro- probe analyses of specimens of Paraconularia byblis and P. subulata from locality 1 90). The precise number of layers and the extent to which this number is consistent from species to species has yet to be determined. In one example of Conularia desiderata, analyzed under the scanning electron microscope, at least thirty very thin, but discrete, layers of calcium phosphate were observed (Fig. 3.9). This stands in marked contrast to previous interpretations of the histology of the conulariid integument (for example, Sinclair, 1 940Z?, 1 948; Richardson,
1 942), in which only two or three layers were observed through use of standard light microscopy.
— Rods (Fig. 2.2), embedded in the integument, are support structures which cross each face transversely; they are composed of calcium phosphate and are subcircular in cross section.
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Each rod crosses one half of each face transversely until its distal end abuts with, or alternates with, the distal end of an adjacent rod. Proximally, rods of adjacent faces articulate in a comer groove. When covered by integument, a rod forms a thin ridge which traverses one half of each face of a conulariid skeleton.
A rod may be equipped with numerous spines (Fig. 2.2) which project adaperturally or adaperturally and adapically. When both types of spines are present, adapertural spines seem to be longer than adapical spines. The function of a spine was probably to provide a stronger framework to support the integument (Figs. 2.2, 3.8). Most conulariid species possess rods which have spines. Some, however, such as Paraconularia planicostata (for example. Fig. 3.1) and P. subulata (for example. Figs. 3.2, 3.6), have integument supported by rods alone.
Rods may or may not possess small nodes on the external side of the exoskeleton (Fig. 2.2). Nodes, if present, are arranged in a single row along a rod. These structures occur slightly adapically of each adapertural spine in all of the taxa reported herein (for example, Fig. 3.7). Thus, the number of adapertural spines and the number of nodes are equal. In Paraconularia sorrocula, the nodes seem to be fused with the adapertural spines, forming a single structure (Fig. 28.2).
The manner of rod articulation has long been used as a diagnostic character at the species level (Hall, 1859; Barrande, 1867; Holm, 1893). To delineate a ‘‘natural grouping” of the conulariids. Holm (1893) identified four species groups, based partially upon ridge characteristics. Richardson (1942) identified four modes of ridge arching among or- ganisms which were then considered conulariids, presumably including
Fig. 3. — 3. 1-3.4; rod articulation styles. 3.1; USNM 33785, Paraconularia planicostata (Dawson) showing gothic arch style; locality 165. 3.2; NYSM 3491, P. subulata (Hall) showing inflected circular curve style; locality 203. 3.3; AMNH 33018, Conularia pyr- amidalis Hall showing inflected gothic arch style; locality 1 17. 3.4; AMNH CU 282G, C. elegantula Meek showing angulated circular curve style; locality 174. 3.5-3.10; scan- ning electron micrographs; arrows point in apertural direction. 3.5; USNM 395833, P. subulata (Hall); view along midline showing integument draped loosely over rods; locality 190;. 3.6; USNM 395834, P. subulata (Hall), rods at midline, integument lacking; locality 72. 3.7; USNM 395830, P. byblis (White); integument closely draped over rods; locality 190. 3.8; USNM 395832, C desiderata Hall, view of ridges, interridge crests and inter- ridge furrows; rods have been broken away; locality 135. 3.9; USNM 395832, same specimen as in Fig. 3.8, C. desiderata Hall, ridge with rod removed, showing multilayered integument; locality 135. 3.10; USNM 395832, same specimen as in Fig. 3.8, C desi- derata Hall, view showing ridges with rods broken away, interridge crests and interridge furrows; locality 135. Bar scales equal 1 mm for Figs. 3. 1-3.4 and 0.1 mm for Figs. 3.5- 3.10.
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Fig. 4. — Conularia congregata Hall. 4.1; NYSM 3483, several paralectotypes from slab exhibiting lectotype and twelve paralectotypes, preserved in black shale. Note that shells
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Sphenothallus, now considered a worm (Mason and Yochelson, 1985; Feldmann et al., 1986). Babcock and Feldmann (1986) recognized four modes of rod articulation which produce patterns of ridges useful as species-level taxonomic criteria. Their terminology for rod articu- lation patterns is followed herein; examples of each style are given in Figs. 3. 1-3.4.
One additional feature of rod articulation at the midline is of interest for species-level determination of taxa. The two rods on a face which meet, or approach closely, at the midline are termed a rod pair. If the right rod of a rod pair is closest to the aperture when the specimen is examined with the aperture upward, the pattern of articulation is right superior (for example. Figs. 3.1, 3.2, 3.6). Conversely, if the left rod of a rod pair is closest to the aperture, the pattern is termed left superior (for example. Fig. 3.5). Some species can be distinguished from others, in part, by the relative proportions of right superior to left superior and to abutting rods on the major and minor faces.
An angle subtended by lines connecting the distal point of a rod with the point at which it joins the comer angle and a line projected across a face, perpendicular to the trend of the comer angle, from the point where a rod joins the comer angle, is termed a rod angle (Fig. 1 .2). The description of a rod angle, combined with information on the style of rod articulation, is used herein as an effective way to help distinguish conulariid species. Rod angles generally vary across an exoskeleton within a few degrees. Notable changes in rod angles are often observed at the narrowest points of exoskeletal constrictions.
A complex of structures is produced on the external surface of the exoskeleton when the integument is draped over a framework of rods and spines. When integument covers a spine, an interridge crest is formed; when integument is draped between two adjacent spines, an interridge furrow is produced. A specimen preserved with the integu- ment draped loosely over the rod and spine framework may appear to be of a different species than a specimen having integument diagenet- ically compressed close to the framework. It may also appear to be different from a specimen which lacks the integument altogether.
Midline.— A line, of variable distinctness, runs longitudinally down the middle of each face (Fig. 1.1). This line, termed the midline, may be a raised structure or a groove, and seems to be pigmented in some
of the inarticulate brachiopod Discina humilis Hall are attached to the conulariid exo- skeletons; locality 154. 4.2; NYSM 3484, detail of ridge structure as preserved in a counterpart specimen. 4.3; NYSM 3484, same specimen as in Fig. 4.2; locality 154. 4.4; NYSM 3483; lectotype and three paralectotypes exposed on same slab as specimens in Fig. 4.1; lectotype is indicated by an arrow. Bar scales represent 1 cm.
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cases (Fig. 23.5). Wiman (1895), Knod (1908), and Liu (1981) have published hand-drawn figures of three separate structures radiating inward from the midlines of conulariids. Wiman (1895) described large T- or Y-shaped ‘‘septa,” Liu (1981) figured elongate “septa” and Knod (1908) figured localized thickenings of integument. Among specimens examined from Devonian and Mississippian rocks of North America, there seems to be no evidence of an elongate, inwardly-directed struc- ture associated with the midline. However, a thickening of the midline, on the internal side of the exoskeleton, has been observed on specimens of Paraconularia ulrichana from the Devonian of Bolivia. In some cases, there appears to be an invagination of the exoskeleton from the interior at the midline (Fig. 14.2).
Corner groove. — A comer groove (Fig. 1 . 1) is the facial invagination at a comer of the exoskeleton where two faces meet at right angles. An individual rod articulates proximally with an adjacent rod in a comer groove; rods on adjacent faces alternate in position along the comer groove. No evidence of localized thickenings of integument interior to the comer grooves, as described by Knod (1908), has been observed. The points of articulation of the rods in the comer grooves seem to be nodose swellings (Fig. 20.4).
Exoskeletal constriction. ■— An exoskeletal constriction (Fig. 1.1) is a slight depression in the conulariid exoskeleton which is identifiable at the same relative position on all faces. An exoskeletal constriction appears to be continuous about the four faces of the exoskeleton (for example. Figs. 5.3, 9. 3-9. 5, 10.1, 27.1-27.3). Trends of rod angles may change slightly (Figs. 8.3, 8.5), rods may converge (Figs. 8.5, 1 1.3) or an apical wall may be attached internally to the exoskeleton at the narrow end of such a structure (Figs. 1 1.2, 14.1). Exoskeletal constric- tions may be indications that conulariids grew by the incremental ad- dition of new integument and rods at the aperture. Because the apertural constriction, the last formed of the exoskeletal constrictions, is always located slightly adapically of the aperture (for example. Fig. 10.1), it is presumed that growth temporarily ceased near the widest portion of the exoskeleton, located between two adjacent exoskeletal constric- tions.
"" Septa.'' —ThQ term “septum” has been applied to three separate morphologic features in conulariids: 1 , apical walls (for example, Miller in Sowerby, 1821; Slater, 1907); 2, ridges (Slater, 1907); and 3, T- or Y-shaped structures or unmodified elongate structures radiating inward from the faces at their midlines (Wiman, 1895; Kiderlen, 1897; Liu, 1981). None of these seems to be an appropriate application of the term. Most commonly, the word “septum” as applied to conulariids means large T- or Y-shaped structures of the exoskeleton that project inward from the midlines. These supposed hard-part structures were described in Conularia loculata, from the Silurian of Sweden by Wiman
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(1895) and were thought by Kiderlen (1 937) to be homologous to septa composed of endodermal tissue in living scyphomedusans. These struc- tures have not been observed, at least not to such a marked extent, in any specimens other than those of Wiman. Wiman’s material was illustrated only by drawings, and the specimens are now lost (W. A. Oliver, Jr., personal communication). Thus, Wiman’s observations cannot be replicated.
The cross sectional view of a specimen of Pamconularia subulata illustrated in Fig. 33.4 exhibits a pattern of limonitic staining in the central cavity which roughly approximates Wiman’s figures. The stain- ing in this example is probably related to the preservation of incom- pletely decomposed internal viscera. It is suspected that Wiman and others may have been misled by some taphonomic feature such as this.
Soft-part morphology . — Much, speculation has surrounded the study of the soft-parts of conulariids. Since the work of Kiderlen (1937), Knight (1937), Moore and Harrington (1956^z, \956b) and Werner (1966, 1967, 1969), conulariids have been interpreted as tentacled creatures. Support for such interpretation is weak, being based on a presumed homology of conulariids to medusoid cnidarians or upon a grouping of true medusoid cnidarians with the conulariids (Kiderlen, 1937; Knight, 1937; Moore and Harrington, 1956<2, \956b).
Remains of presumed conulariid soft-parts were independently de- scribed from European Devonian conulariids by Steul (1984) and from North American Mississippian conulariids by Babcock (1985a) and Babcock and Feldmann (1986). Babcock and Feldmann (1986), work- ing only with exceptionally preserved three-dimensional specimens, identified a single elongate tube that extends the length of the central cavity and a large globular shaped structure near the aperture (Fig. 2. 1). Steul’s (1984) work, based upon x-ray analyses of collapsed specimens preserved in the Hunsruck Slate, revealed other structures which may be preserved soft-parts, though the evidence is ambiguous.
The tubular and globular internal structures (Figs. 30.2=30.6), pre- sumably representing remains of organ systems, appear to be reduced in size compared to expected living organs and show no details of soft- part anatomy. These structures may be somewhat contracted masses of internal tissues. These structures may have been preserved, in outline at least, as altered remains of partially digested food matter and/or sediment left in the intestinal tract when the animals died. Other organs that were originally present in the North American studied specimens may have decayed.
Occurrences and Paleoecology
Conulariids have been reported exclusively from marine rocks rang- ing in age from the Late Precambrian to the Recent (Caster, 1957).
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Fig. S. — Conularia desiderata Hall. 5.1; AMNH 2697, holotype; major face, preserved in limestone; locality 121. 5.2; NYSM 3487, holotype of C continens var. rudis Hall,
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This range, however, includes a variety of taxa now referable to other groups. Occurrences here considered valid include specimens from Lower Ordovician (Sinclair, 1948) through Upper Triassic (Gou and Yang, 1985) rocks. While articulated conulariids are uncommon or rare in many instances, conulariids are, nonetheless, pervasive faunal elements in Middle Ordovician through Permian marine rocks. They are rare in the Lower Ordovician and in the Triassic. Some Paleozoic occurrences yield abundant conulariids.
Fossils identified as conulariids have been identified from all con- tinents. However, the sole report of a conulariid from Antarctica (Cor- dini, 1955) has been received with skepticism and may represent plant material (Dalziel et aL, 1981).
The conulariid skeleton, or exoskeleton, is composed of a calcium phosphate framework, made of rods, and usually having spines or nodes. The framework is set in an interlayered integument made of thin sheets of calcium phosphate and protein. Overall, the exoskeleton was probably fairly delicate, and upon death of the animal, was readily subject to collapse, occasional attachment of epibionts and decom- position accompanied by disarticulation (Feldmann and Babcock, 1986).
Most occurrences of articulated conulariids involve rapid burial and often, early diagenesis. For example, conulariids are abundant in the Meadville and Wooster members of the Cuyahoga Formation in north- eastern and central Ohio. Specimens in these units are usually found in presumed tempestite beds or in siderite concretions. Specimens col- lected from tempestite beds often seem to be current aligned (Fig. 30.7). Specimens collected from siderite concretions, such as that illustrated in Fig. 10.4, are weakly current aligned, if at all. In the Cuyahoga Formation, concretions were probably produced through localized in- creases in pH and lowerings of eH, causing precipitation of iron car- bonate. The onset of siderite precipitation probably occurred soon after burial of the animals.
Some occurrences of articulated conulariids involve “prefossilized” specimens, or ones which have undergone early diagenesis, and which have later been exhumed through bioturbation, winnowing of sedi- ments by currents, or both. The specimens were later deposited in beds
major face, preserved in siltstone; locality 132. 5.3; NYSM 3487, same specimen as in Fig. 5.2, comer view. 5.4; AMNH 2697, same specimen as in Fig. 5.1, minor face. 5.5; NYSM 3487, detail of minor face. 5.6; NYSM unnumbered, detail of specimen with integument draped loosely over rods; locality 124. 5.7; NYSM 3485, syntype of C. continens Hall, preserved in black shale. 5.8; NYSM 3486, syntype of C. continens Hall; detail of minor face; locality 125. 5.9; NYSM 3486, same specimen as in Fig. 5,8, nearly complete, flattened specimen. Bar scales represent 1 cm.
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Fig. 6. — 6. 1-6.3; Conularia delphiensis (Maroney and Orr) preserved in phosphatic con- cretions at a stratigraphic discontinuity surface (Maroney and Orr, 1974). 6.1; lUPC
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representing stratigraphic discontinuities (Maroney and Orr, 1974; Baird, 1981; Baird and Brett, 1981; herein. Figs. 6. 1-6.3).
The general scarcity of articulated conulariid remains, at most lo- calities, may be related to rapid disarticulation of the multielement skeleton, predation and scavenging, and the constancy of water move- ment near the sediment-water interface. The frequent occurrence of articulated conulariids in low diversity faunas (Ruedemann, 1934; Jux, 1960; Babcock and Feldmann, 1986; Feldmann and Babcock, 1986) may be largely a function of low biotic activity in these environments.
Many conulariids show indications that some breakdown of the exo- skeleton has occurred, especially in the vicinity of the aperture (for example. Figs. 5.7, 7.5, 7.7), where the integument may be greatly reduced in thickness or lacking. In rare instances, poorly bioturbated stratigraphic units yield specimens of conulariids in which the integ- ument is lacking over much of the exoskeleton and disarticulated rods have been displaced from their original positions (Fig. 21.4). In sedi- ments that were probably well bioturbated, or sediments where water movement took place rather constantly, articulated conulariids are rare. It is possible that conulariid rods will be found in some Paleozoic and Mesozoic rock units that are sampled for microfossils by insoluble residue or other techniques. Disarticulated conulariid rods have been occasionally misidentified as fossil fish bones (Feldmann and Babcock, 1986).
Many species of conulariids seem to have been geographically wide- spread. For example, the Mississippian species, Paraconularia ches- terensis, has a known geographic range from Alabama to British Co- lumbia and, as with numerous other species, occurs in rocks of various lithologies. These lithologies include mudstones and wackestones (Chester Group of Illinois), gray shales (Borden Group of Indiana) and siderite concretions (Cuyahoga Formation of Ohio). The occurrence of this and other conulariid species in stratigraphic units of so vastly dissimilar lithologies and environments of deposition, and their wide geographic distribution, suggests that some species may have been planktonic or pseudoplanktonic at some point in the life cycle. The bilaterally symmetrical body plan (Babcock and Feldmann, 1984, 1986;
14470-1, holotype; view of flattened specimen and detail of minor face; locality 23. 6.2; lUPC 14470-2, paratype; locality 23. 6.3; lUPC 14470-1, same specimen as in Fig. 6.1, detail of ?minor face. 6.4-6. 5; C. desiderata Hall. 6.4; NYSM unnumbered, view of ?majorface; locality 132. 6.5; USNM 395832, detail of specimen with most of integument removed, exposing broken rods and intact spines; locality 135. Bar scales in Figs. 6.1- 6.4 represent 1 cm; bar scale in Fig. 6.5 represents 5 mm.
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Steul, 1984), however, may be an indication that some species were weakly nektonic.
Conulariids are usually referred to as solitary animals. Clusters are rare, but a few have been figured by Slater (1907, Plate 2, fig. 1), Hall (1876, Plate 28, fig. 1; 1879 Plate 24, fig. 1), Sinclair (1944, Plate 2, fig. 5) and Babcock and Feldmann( 1984, p. 17; Babcock and Feldmann, 1986, figs. 4A, 4C). In all these clusters, individual specimens are shown radiating about a central area. Apices are usually pointing inward in these aggregations (for example. Fig. 24.3). Occasionally, apices of such specimens are preserved intact (Fig. 24.1), yet in portions of a somewhat disaggregated cluster illustrated by Hall (1876, 1879), here illustrated in Figs. 4. 1 and 4.4, the apices are missing, perhaps because of post-mortem decay. The apical walls are visible in some of these specimens.
A slab exhibiting numerous P. chesterensis from the Borden Group (Mississippian) of Indiana exhibits several small, black to reddish brown, tubular structures, circular in cross section, attached to, or very near the apices of, conulariids (Figs. 24. 1, 24.3). Combined with discoveries of similar structures on conulariids from the Borden Formation (Mis- sissippian) of Kentucky (Fig. 24.2) and from the Etherington Formation (Mississippian) of British Columbia (Fig. 32.5), these structures are interpreted as attachment stalks. The association of conulariids with plant remains, perhaps algae (for example. Fig. 28.7), indicates that some conulariids may have been pseudoplanktonic and were attached to, or entwined with, planktonic algae. Alternatively, the conulariids may have attached to plant remains which had previously settled to the ocean floor.
Cluster associations involving numerous specimens, usually com- prising single species, serve to indicate that some, if not all, conulariids were gregarious, at least during some part of the life cycle. No evidence of budding or any other asexual reproductive style exists.
Regardless of whether the conulariids were planktonic, pseudoplank- tonic, benthonic, or even nektonic, attached or free swimming, the soft- part organs in the region of the aperture of the exoskeleton probably functioned for filter feeding. There is no evidence for aggressive food gathering behavior.
The style of growth in conulariids was incremental, with addition of rods and spines taking place at the aperture. Growth lines, such as those seen on mollusks and brachiopods are not present. Exoskeletal con- strictions, in exactly the same relative positions on all four faces, are possible evidences of incremental growth. Additional evidence of in- cremental growth at the aperture comes from some specimens showing healed injuries (for example. Figs. 20.1, 29.5). Along the exoskeletal
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constrictions of such examples, rods may be broken or the rod artic- ulation patterns may be disrupted. This indicates two things: 1, that the positions of apertural terminations changed and correspond to the positions of apertural constrictions; and 2, that the apertural termi- nations were easily fragmented but healing could occur during a suc- ceeding growth phase.
Development of multiple apical walls may also have been a function of an incremental growth pattern. When the internal cavity of an in- dividual reached a certain volume, an apical wall was probably secreted adaperturally of the apical terminus. Apical walls are commonly pre- served among specimens referable to the genus Conularia but are rarely found on specimens of Paraconularia. No specimens of Reticulacon- ularia have been observed with apical walls in place.
Epibionts on conulariids include orbiculoid brachiopods (Hall, 1876, 1879; Moore and Harrington, 1956a; Babcock and Feldmann, 1984, 1986; herein, Figs. 4.1, 4.4, 5.9, 13.1, 13.5, 16.4, 28.1-28.2), en- crusting bryozoans (Finks, 1955; herein. Figs. 22.4-22.5) and edrioas- teroid echinoderms (Barrande, 1867; Moore and Harrington, 1956a). Attachment of epizoans has been interpreted as either having occurred during the life of the conulariid (Finks, 1955; Moore and Harrington, 1956a) or as having occurred after the death of the conulariid (Baird, 1981; Babcock and Feldmann, 1986). In all the specimens examined in the course of this study, orbiculoids seem to have attached to the conulariids after they died and came to rest on the sediment surface. This is supported by the observation that orbiculoids are always found attached to only two faces of a conulariid (Figs. 4.1, 4.4). Presumably, these two faces were the only ones which projected above the sediment- water interface. A specimen is illustrated herein (Fig. 25.5) of a bryo- zoan encrusting a conulariid in the region of a comer groove. There is no sign of damage to the bryozoan on this specimen, indicating that encrustation by the bryozoan occurred after the death of the conulariid.
Systematic Paleontology
Summary of taxa.--A total of 69 trivial names have been applied to Devonian or Mississippian conulariids of North America prior to this paper. Of these, 54 taxa were published, and eight were described in unpublished manuscripts. Herein and in Part B, 28 species are rec- ognized as valid, of which five are new. However, primary type spec- imens of 1 8 species were not available for study; therefore, the status of these taxa was not assessed fully. In Part B, one species is removed from the Conulariida. The conulariids described below and in Part B are divided among three genera, Conularia, Paraconularia and Retie- ulaconularia, n. gen. The genus Diconularia Sinclair, 1952 is regarded
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as a junior synonym of Conularia Miller in Sowerby, 1821 and the genus Adesmoconularia Driscoll, 1963 is considered to be a junior synonym of Paraconularia Sinclair, 1 940a.
The list below summarizes all described species-level taxa of con- ulariids from North American Devonian and Mississippian rocks and indicates the status of each trivial name as currently recognized. Taxa are arranged alphabetically according to their presently recognized sta- tus. Junior synonyms and previously used combinations of each taxon are listed below each valid name.
Conularia congregata Hall, 1876
Conularia delphiensis (Maroney and Orr, 1974)
Ctenoconularia delphiensis Maroney and Orr, 1974 Conularia desiderata Hall, 1861 Conularia continens Hall, 1876 Conularia continens var. rudis Hall, 1879 Conularia elegantula Meek, 1871 Conularia milwaukeensis Cleland, 1911 Conularia congregata var. milwaukeensis Cleland, 1911 Conularia multicostata Meek and Worthen, 1865 Conularia micronema Meek, 1871 Diconularia micronema (Meek, 1871)
Mesoconularia mcfarlani Sinclair, [1948] MS Mesoconularia attica Sinclair, [1948] MS Conularia pyramidalis Hall, 1859 Conularia huntiana Hall, 1859 Conularia lata Hall, 1859 Conularia subcarbonaria Meek and Worthen, 1865 Conularia intertexta Miller, 1894 Conularia spergenensis Miller and Gurley, 1893 Conularia tuzoi Clarke, 1907 Conularia desiderata var. tuzoi Clarke, 1 907 Conularia ulsterensis Howell, 1 942 Conularia undulata Conrad, 1 84 1 Conularia cayuga Hall, 1876 Conularia crebistria Hall, 1876 Paraconularia alternistriata (Shimer, 1926)
Conularia alternistriata Shimer, 1926 Paraconularia alpenensis Babcock and Feldmann, n. sp.
Paraconularia blairi (Miller and Gurley, 1893)
Conularia blairi Miller and Gurley, 1893 Conularia sedaliensis Miller and Gurley, 1896 Paraconularia indiana Sinclair, [1948] MS Paraconularia byblis (White, 1862)
Adesmoconularia byblis (White), 1862 Conularia byblis V^hiXe, 1862 Paraconularia chagrinensis Babcock and Feldmann, n. sp.
Paraconularia chesterensis (Worthen, 1883)
Conularia chesterensis Worthen, 1883 Paraconularia missouriensis (Swallow, 1 860)
Conularia missouriensis Sw2l\\ov>/ , 1860
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Conularia gratiosa Miller and Gurley, 1893 Conularia greenei Miller and Gurley, 1 896 Paraconularia sciotoviliemis Driscoll, 1963 Paraconularia okiahomaemis Babcock and Feldmann, n. sp,
Paraconularia planicostata (Dawson, 1868)
Conularia planicostata Dawson, 1868 Paraconularia recurvatus Babcock and Feldmann, n. sp.
Paraconularia salinensis (Whiteaves, 1891)
Conularia salinensis Whiteaves, 1891 Paraconularia sorrocula (Beede, 1911)
Conularia sorrocula Beede, 1911 Paraconularia subulata (Hall, 1858)
Conularia subulata 1858 Conularia victa White, 1862 Conularia Winchell, 1865
Conularia whitei Meek and Worthen, 1865 Conularia sampsoniMillQT , 1892 Paraconularia wellsvillia Babcock and Feldmann, n. sp.
Paraconularia yochelsoni Babcock and Feldmann, n. sp.
Reticuiaconularia penouili (Clarke, 1 907)
Conularia penouili Clarke, 1 907 Conularia gaspesia Sinclair, 1 942 Reticuiaconularia sussexensis (Herpers, 1 949)
Conularia sussexensis Herpers, 1 949
Whereabouts of Type Material Unknown
Climacoconus viata Swartz and Richardson in Richardson, [1942] MS Conularia crawfordsvillensis Owen, 1862 Conularia gracilis WTnck, 1888
(Name preoccupied by C. gracile Hall, 1847; changed to C. herricki by Miller, 1892.) Conularia grandis Roemer, 1856 Conularia herricki Miller, 1892 Conularia jervisensis Shimer, 1 905
Conularia latoides Swartz and Richardson in Richardson, [1942] MS
Conularia marionensis Swallow, 1860
Conularia missouriensis var. hermansi Calvin, 1890
Conularia molaris White, 1876
Conularia novascotica Hartt, in Dawson, 1868
Conularia osagensis Swallow, 1863
Conularia pyramidalis var. parvinodis Swartz and Richardson in Richardson, [1942] MS Conularia siphunculophora Swartz and Richardson in Richardson, [1942] MS Conularia triplicata Swallow, 1860 Conularia verneuiiia Emmons, 1 846 Paraconularia welleri Sinclair, [1948] MS
Non-Conulariid
Conularia tenuicostata Branson, 1938 (Assigned, tentatively, to phylum Priapulida.)
Repositories. — Specimens are listed according to catalogue numbers with the repositories abbreviated as follows:
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AMNH AMNH CU
BMS
CM
CMNH
FMNH P
FMNH PE
FMNH UC
GSC
ISGS
lUPC
KSU
NJSM
NYSM
oc
PU
RM(MU)
UCGM
UIPC
UK
UMC
UMMP
USNM
wvu
American Museum of Natural History, New York, New York American Museum of Natural History, Columbia University Collection, New York, New York
Buffalo Museum of Science, Buffalo, New York
Carnegie Museum of Natural History, Invertebrate Paleontology Collec- tions, Pittsburgh, Pennsylvania
Cleveland Museum of Natural History, Invertebrate Paleontology Col- lections, Cleveland, Ohio
Field Museum of Natural History, Paleontology Collection, Chicago, Il- linois
Field Museum of Natural History, Invertebrate Paleontology Collection, Chicago, Illinois
Field Museum of Natural History, University of Chicago Collection from Walker Museum, Chicago, Illinois Geological Survey of Canada, Ottawa, Ontario
Illinois State Geological Survey, Illinois State Museum, Champaign-Ur- bana, Illinois
Indiana University Paleontological Collection, Bloomington, Indiana Department of Geology, Kent State University, Kent, Ohio New Jersey State Museum, Trenton, New Jersey New York State Museum and Science Service, Albany, New York Oberlin College Paleontological Collections, Oberlin, Ohio Department of Geology, Princeton University, Princeton, New Jersey Redpath Museum (McGill University), Montreal, Quebec University of Cincinnati Geological Museum, Cincinnati, Ohio University of Illinois, Department of Geology, Champaign-Urbana, Il- linois
University of Kentucky, Department of Geology, Lexington, Kentucky
University of Missouri-Columbia, Columbia, Missouri
University of Michigan, Museum of Paleontology, Ann Arbor, Michigan
United States National Museum of Natural History, Washington, D.C.
West Virginia University, Department of Geology, Morgantown, West
Virginia
Treatment of manuscript names. — One of the most influential papers regarding the systematics and morphology of conulariids is Sinclair (1948), an unpublished Ph.D. thesis. In it were proposed many new genus-level and species-level taxa. The undescribed genera identified by Sinclair were subsequently published (Sinclair, 1952); however, the majority of Sinclair’s new species have never been formally described. Moreover, many of the species removed by Sinclair from Conularia, as used in the sense of a form-genus, and placed in different genera, have not been published in their revised state by Sinclair. Nevertheless, various authors have used Sinclair’s combinations, often without ref- erence to the authority for such usage.
It is our opinion that, because of the central importance of Sinclair’s (1948) unpublished manuscript to the study of conulariid systematics, Sinclair’s unpublished species-level names and his unpublished com- binations should be included in the formal synonymies of the Devonian and Mississippian taxa discussed below. For the sake of completeness,
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we have also included in the present work the manuscript names of Swartz and Richardson in Richardson (1942). By including these un- published names in synonymy though, we do not intend to suggest that these are available names. According to Article II(d)(ii) of the Inter- national Code of Zoological Nomenclature, Third Edition (Interna- tional Commission of Zoological Nomenclature, 1985), “a previously unavailable name is not changed by its mere citation accompanied by a reference to the work in which the name was published but was not made available.” Also, in Article 1 1(e), the Code states that a, “name first published as a junior synonym is not thereby made available unless prior to 1961 it has been treated as an available name ...”
Phylum Conulariida Babcock and Feldmann, new phylum
—Animals generally possessing a four sided, steeply py- ramidal exoskeleton; bilaterally symmetrical; integument composed of calcium phosphate and protein, multilayered, moderately flexible; exo- skeletai framework composed of calcium phosphatic rods arranged transversely across each side face; adjacent rods abut or alternate at midline of each face; rods of adjacent faces articulate in a groove at junction of two faces; apical end closed either by a blunt point; one or more smooth apical walls may be present internal to the exoskeleton and aperturad the apex; apex sheathed by a ?chitinous, phosphatic, or chitinophosphatic stalk; aperture simple and open. No internal hard- part structures known; internal soft-parts comprise an elongate tube extending most of the body length, in addition to one or more globular shaped structures, all of uncertain function.
Remarks. — Approximately 40 genera of organisms have, at one time or another, been grouped among the conulariids. Of these, six genera have been excluded from the phylum to date. Based upon new infor- mation on the architecture of the conulariid skeleton (Babcock and Feldmann, 1986), it seems that even more genera have been de- scribed than are warranted by fossil evidence. Upon further study, several other genera are likely to be excluded from the phylum Con- ulariida.
Organisms which are properly included within the phylum Conu- lariida must possess a bilaterally symmetrical exoskeleton composed of calcium phosphate rods and layered calcium phosphate and protein integument. Herein, genera of conulariids are defined upon: 1, the relative spacing of rods; 2, the relative proportion of rods that abut at the midline to those that alternate; 3, the apical angles; 4, the presence or absence of nodes and spines; and 5, the spacing of nodes and spines. Species are distinguished using the characters upon which generic dis- tinctions are made as well as the following: 1, the patterns of rod articulation; and 2, the rod angles.
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Descriptions of conulariids represented in the Devonian and Mis- sissippian rocks of North America follow. After examination of spec- imens referred to the respective type species from Europe, Sinclair (1940i2) proposed North American reference species. The following diagnoses of Conularia and Paraconularia are based largely upon spec- imens referable to the North American reference species. The new genus Reticulaconularia is based upon specimens from the Devonian of east- ern North America.
Specimens of Conularia are recognizable by their closely spaced rods, by having more than 40% of the rods abutting at the midline, by having small apical angles and by having both nodes and spines which are closely spaced. Specimens referable to Paraconularia exhibit widely spaced rods, fewer than 40% of rods abutting at the midline and small apical angles. Nodes and spines may be present; if they are, they are closely spaced. Conulariids referable to Reticulaconularia, n. gen. have widely spaced rods which abut or alternate at the midline in proportions which are not well established yet. They also have large apical angles. Members of the genus Reticulaconularia are notable for the reticulate appearances of the external surfaces of the faces (Figs. 33.1-33.5, 34. 1- 34.2, 34.4), a product of nodes and spines which are widely spaced. It should be noted, however, that other conulariids, if preserved as ex- ternal molds, may exhibit patterns similar to this '‘reticulate” appear- ance (for example. Figs. 8. 5-8. 6). Such specimens are best studied with latex casts.
Key to North American Devonian and Mississippian conulariids.— The following is a key to all species of Devonian and Mississippian conulariids from North America which are currently known. Some categories are ambiguous due to the lack of complete or well preserved specimens of some species, so it is not intended for use with specimens of different ages or of other areas of the world.
1 . Faces have reticulate appearance 2
1 . Faces do not have reticulate appearance 3
2, Inflected gothic arch rod articulation present
Reticulaconularia sussexensis (Herpers)
2. Inflected gothic arch rod articulation not present
Reticulaconularia penouili (Clarke)
3. Number of rods that abut at midline less than 40% 4
3. Number of rods that abut at midline greater than or equal to 40% 19
4, Gothic arch rod articulation present ..................................... 5
4. Gothic arch rod articulation not present 9
5. Only gothic arch rod articulation present . . Paraconularia alternistriata (Shimer)
5. Gothic arch and another style of rod articulation present ................... 6
6. Gothic arch and inflected circular curve rod articulation present
Paraconularia oklahomaensis Babcock and Feldmann, n. sp.
6. Gothic arch and inflected gothic arch rod articulation present .............. 7
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7.
7.
8.
8.
9.
9.
10.
10.
11.
11.
12.
12.
13.
13.
14.
14.
15.
15.
16. 16.
17.
17.
18. 18. 19.
19.
20. 20. 21. 21. 22. 22. 23.
23.
24.
24.
25.
25.
26. 26. 27. 27.
Rods/cm fewer than or equal to 1 1 ....... Pamconularia planicostata (Dawson)
Rods/cm greater than 11
Paraconularia yochelsoni Babcock and Feldmann, n. sp.
Inflected gothic arch rod articulation present 9
Inflected gothic arch rod articulation not present 14
Only inflected gothic arch rod articulation present 10
Inflected gothic arch and another style of rod articulation present 11
Rods/cm fewer than 15
Paraconularia chagrinensis Babcock and Feldmann, n. sp.
Rods/cm greater than 15 Paraconularia sorrocula (Beede)
Rods/cm fewer than or equal to 1 2 12
Rods/cm greater than 12 13
Rods are slightly inflected at midline . . Paraconularia blairi (Miller and Gurley)
Rods are not inflected at midline Paraconularia subulata (Hall)
Number of rods that abut at midline fewer than 30%
Paraconularia salinensis (Whiteaves)
Number of rods that abut at midline greater than or equal to 30%
Paraconularia byblis (White)
Inflected circular curve rod articulation present; rods recurved near midline . . 15
Inflected circular curve rod articulation present; rods not recurved near midline
Paraconularia wellsvillia Babcock and Feldmann, n. sp.
Rods/cm fewer than or equal to 1 2 17
Rods/cm greater than 12 16
Rods/cm fewer than 18 . . Paraconularia alpenensis Babcock and Feldmann, n. sp.
Rods/cm greater than or equal to 18
Paraconularia recurvatus Babcock and Feldmann, n. sp.
Rods/cm fewer than or equal to 7 Paraconularia chesterensis (Worthen)
Rods/cm greater than 7 Paraconularia missouriensis (Swallow)
Gothic arch rod articulation present Conularia pyramidalis (Hall)
Gothic arch rod articulation not present 19
Inflected gothic arch rod articulation present 20
Inflected gothic arch rod articulation not present 23
Rods/cm fewer than or equal to 30 21
Rods/cm greater than 30 22
Rods/cm fewer than or equal to 25 Conularia milwaukeensis Cleland
Rods/cm greater than 25 Conularia multicostata Meek and Worthen
Rods/cm fewer than or equal to 39 Conularia tuzoi Clarke
Rods/cm greater than 39 Conularia ulsterensis Howell
Angulated circular curve rod articulation present 24
Angulated circular curve rod articulation not present 27
Only angulated circular curve rod articulation present 25
Angulated circular curve and another style of rod articulation present ...... 26
Rods/cm fewer than 34 Conularia subcarbonaria Meek and Worthen
Rods/cm greater than 34 Conularia elegantula Meek
Rod angle less than or equal to 1 7° Conularia congregata Hall
Rod angle greater than 17° Conularia desiderata Hall
Rods undulose Conularia undulata Conrad
Rods not undulose Conularia delphiensis (Maroney and Orr)
Genus CONULARIA Miller, in Sowerby, 1821
Type species. — Conularia quadrisulcata Sowerby, by original desig- nation; Silurian of England. Holotype is lost. North American reference
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spQciQS, fide Sinclair (1940a): Conularia niagarensis Hall, 1852 (Silu- rian). Syntypes and plastosyntypes of C. niagarensis: AMNH 31625- 31628; plastosyntype of AMNH 31625: FMNH UC 60850.
Diagnosis. — Conulariids with rods that are generally closely spaced, 9-84 rods/cm. Fewer than 60% of rods alternate at midline; more than 40% abut; two adjacent rods on a face form a single arc across the face. Apical angles small, 9-23°. Nodes, adapertural spines and adapical spines usually present and closely spaced, 1-7/mm.
CONULARIA CONGREGATA Hall, 1876 Figs. 4. 1-4.4
Conularia congregata Hall, 1876, PI. 28, fig. 1; Miller, 1877, p. 141; Bigsby, 1878, p. 418; Hall, 1879, p. 214-215, PI. 34, fig. 1, PI. 34A, figs. 9-1 1; Miller, 1889, p. 390; Harris, 1899, PI. 8, fig. 59; Clarke and Ruedemann, 1903, p. 565; Moore and Harrington, 1956^, p. F61, fig. 46.2; Palmer and Brann, 1966, fig. 59; Babcock, 1985Z?, fig. 2; Babcock and Feldmann, 1986, fig. 4A.
Conularia s.l. congregata Hall. Sinclair, 1948, p. 286.
Conularia pyramidalis Hall. Sensu Babcock and Feldmann, 1984, p. 17.
Description. — ExosktlQion up to 11 cm in length. Major apical angle 17-18°; minor apical angle 12-14°. Rod articulation exclusively of inflected circular curve style. Rods almost always abut at midline; rod angle 9-13°. 16-21 rods/cm. 6-7 nodes/mm; 6-7 adapertural spines/mm; 6-7 adapical spines/mm; spines often not easily discernible. Apical wall may be present.
Occurrences. Devonian of New York; localities 149-1 50 and
152-160. Maillieux (1933), Markovski and Nalivken (1934) and Xu and Li (1979) reported C. congregata from Devonian rocks in Belgium, the U.S.S.R. and the People’s Republic of China, respectively, but these occurrences have not been confirmed.
Types. —LQcXoXypQ and twelve paralectotypes on one slab, NYSM 3483. Remains of at least thirteen specimens are preserved on James Hall’s slab of syntypes (Figs. 4.1, 4.4). From this syntypic suite, the best preserved of Hall’s figured specimens (Fig. 4.4) is here chosen as the lectotype of C congregata. The remaining ten specimens are con- sidered paralectotypes. Counterparts of the paralectotype specimens shown in Fig. 4.1 are catalogued as FMNH unnumbered.
Remarks. — Conularia congregata Hall is most similar in morphology to C. desiderata Hall. The similarities lie in overall size, apical angle values, rod angle values and in the presence of inflected circular curve rod articulation. The differences between the two taxa are subtle. In specimens of C. congregata, few rods, generally fewer than 10%, al- ternate at the midline; also, rods show very little or no inflection toward the aperture near the midline. Among specimens referable to C. desi- derata, as many as 15% of the rods may alternate at the midline; specimens also show a strong adapertural inflection of the rods at the
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midline. Some specimens, such as that illustrated in Fig. 5.6, appear to be intermediate in morphology between C desiderata and C. con- gregata.
Conularia congregata may also be confused with C pyramidalis Hall. Conularia pyramidalis differs from both C congregata and C. desi- derata is having inflected circular curve rod articulation in the apical region and inflected gothic arch rod articulation elsewhere. There is no evidence in C pyramidalis that the rods inflect near the midline.
Material examined.— AO specimens; housed in the AMNH, BMS, FMNH, NYSM, USNM, and the private collection of Paul Zell.
CONULARIA DELPHIENSIS (Maroney and Orr, 1974)
Figs. 6. 1-6.3
Conularia sp. Kindle, 1901, p. 737, PL 123, fig. 8.
Ctenoconularia delphiensis Maroney and Orr, 1974, p. 3-6, fig. lA-F.
Description.— Exos\LQ\t\on up to 5.5 cm in length. Major apical angle and minor apical angle about 1 5°. Rod articulation inflected gothic arch style in apical region and inflected circular curve style elsewhere. Rods usually abut at midline; rod angle 10-11°. 26-42 rods/cm. 6-7 nodes/mm; adapertural spines and adapical spines not observed. Apical wall not observed.
Occurrence. Devonian of Indiana; localities 22-23.
Fpppy. —Holotype, lUPC 14470-1; five paratypes, lUPC 14470-2 through 14470-6.
Remarks.— ThQ six specimens which comprise the type suite of C delphiensis are badly preserved. Although no spines were actually ob- served on any of the type specimens, all other morphologic features are consistent with species of the genus Conularia as defined herein. A specimen figured by Kindle (1901, pi. 123, fig. 8; USNM 62210), an apparent external mold of C. delphiensis, clearly shows that nodes are present, but spines are not evident. Spines may have been present in this taxon, but have not been observed because of the poor pres- ervation of the specimens studied. A cross sectional view of an un- compressed specimen referable to this species has not been observed.
In their original description of C. delphiensis, Maroney and Orr (1974) did not indicate why they chose to include the species in the genus Ctenoconularia Sinclair, 1952. Moreover, they only compared this species to Conularia congregata. Sinclair (1952, p. 141) noted that the primary distinguishing characteristic of specimens referable to Ctenoconularia was “strikingly slender shells.” This is certainly true in the type species, Ctenoconularia obex Sinclair. Judging from Sin- clair’s published figures (1952, figs. 56 A-C), the major and minor faces subtend angles of 4° and 3®, respectively. In other respects, specimens of Ctenoconularia are very similar to specimens of Conularia. ^^Cten-
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Fig. 7. — 7. 1-7.4; Conularia elegantula Meek. 7.1; AMNH CU 282G, holotype, comer view; locality 174. 7.2; AMNH CU 282G, same specimen as in Fig. 7.1, comer major
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oconularia"" delphiensis, which has apical angles of approximately 1 5°, seems to expand from the apex too rapidly to warrant inclusion in the genus Ctenoconularia.
Conularia delphiensis is similar in general morphology, including similarity of rod articulation styles, to only one Devonian species from North America, C. milwaukeensis. Conularia delphiensis differs, how- ever, in having greater rods/cm values. Specimens here referred to C delphiensis possess 26=42 rods/cm while specimens here referred to C. milwaukeensis have fewer than 30 rods/cm.
Material examined. — ^ specimens; housed in the lUPC and the USNM.
CONULARIA DESIDERATA Hall, 1861 Figs. 3.8=3.10, 5.1-5.9, 6.4=6.5, 16.6
Conularia =? Hall, 1859, p. 480.
Conularia desiderata Hall, 1861, PI. 72A, fig. 4; Richardson, 1942, p. 30-32, PI. 4, figs.
7, 8; Babcock and Feldmann, 1986, figs. IB, IH.
Conularia continens Hall, 1876, PI. 28, figs. 4-6; Miller, 1877, p. 141; Bigsby, 1878, p. 418; Hall, 1879, p. 212-214, PI. 33, fig. 6, PI. 34, figs. 3, 4, 6, PI. 34A, fig. 6; Lesley, 1889, p. 143, fig.; Miller, 1889, p. 390; Clarke and Ruedemann, 1903, p. 565; Moore and Harrington, 1956a, fig. 25. 1; Moore and Harrington, 1956A p. F60, fig. 42. 10a- b; Tasch, 1973, fig. 5.15H, Tasch, 1980, fig. 5.15H.
Conularia continens var. rudis Hall, 1879, p. 215-216, PL 34A, figs. 7-8; Miller, 1889, p. 390; Clarke and Ruedemann, 1903, p. 565; Grabau, 1906, p. 331.
Conularia s.l. continens Hall. Sinclair, 1948, p. 286,
Par aconularia desiderata {Hail). Sinclair, 1948, p. 185.
Conularia s.l. rudis Hall. Sinclair, 1948, p. 284.
Description. —ETLOsk&lQton up to 10 cm in length. Major apical angle 14-27®; minor apical angle 14-18°. Rod articulation inflected circular curve in early stages to angulated circular curve in later stages. Rods almost always abut at midline; rod angle 7-17°. About 41 rods/cm in apical region; 14-27 rods/cm elsewhere. 3-4 nodes/mm; 3-4 adapertural spines/mm; 3-4 adapical spines/mm. Apical wall not observed.
Occurrences. — and Middle Devonian rocks of New York and
Pennsylvania; localities 121, 124=126, 132-135, 142, 144, 232, 235- 236, and 239. Conularia continens, here referred to C. desiderata, has been reported from Devonian rocks in Ohio (Claypole, 1903) and in Indochina (Patte, 1 926), but the specimens upon which these references were based were not studied. Woodward (1943) identified C continens
face. 7.3; AMNH CU 282G, same specimen as in Fig. 7.1, detail of major face. 7.4; CMNH 44584, ?major face of flattened specimen; locality 175. 7. 5-7. 9; C. milwaukeensis Cleland. 7.5; USNM 85988, holotype; locality 255. 7.6; USNM 78212; detail of apical region. Note that no apical wall is present; locality 253. 7.7; MPM 20252, complete specimen; locality 253. 7.8; MPM 20252, same specimen as in Fig. 7.7, detail of major face. 7.9; MPM 22974; locality 253. Bar scales represent 1 cm.
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Fig. %.-~Conularia pyramidalis Hall. 8.1; AMNH 33017, lectotype; comer view of flat- tened specimen, preserved in calcareous shale; locality 117. 8.2; NYSM 3488, holotype of C. huntiana Hall, comer view, preserved in calcareous shale; locality 1 1 8. 8.3; AMNH
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var. rudis (=C delphiensis) in the Devonian of West Virginia, but Woodward’s specimens were unavailable for study.
Types. — Holotype and plastoholotype of C delphiensis Hall, AMNH 2697; two syntypes of C continens Hall, NYSM 3485, NYSM 3486; holotype of C continens var. rudis Hall, part and counterpart, NYSM 3487.
Remarks. — Conularia desiderata Hall is similar in appearance only to C. pyramidalis Hall. Both species have rod angles in the range of 14-27°. Also, specimens of both taxa possess rods which appear to be inflected at the midline. Features that are present in C. desiderata, but not present in C pyramidalis include inflected circular curve to an- gulated circular curve rod articulation style and rod angles of 14-27°.
The syntypes of C. continens Hall differ morphologically from the holotype of C desiderata Hall only in being flattened. The holotype of C continens var. rudis differs from the holotype of C desiderata in being larger and better preserved. The type specimens of each taxon have inflected gothic arch rod articulation in the region close to the apex and inflected circular curve rod articulation elsewhere. The major apical angle of the holotype of C desiderata, measures 23°; the minor apical angle cannot be measured with certainty. The syntypes of C continens have major apical angles of 16° and 19°. The major apical angle of the holotype of C. continens var. rudis is 27°. All of the type specimens have between 14 and 27 rods/cm and possess spines which are difficult to observe in most specimens. Conularia continens Hall, and C continens var. rudis Hall are, therefore, here considered junior synonyms of C desiderata Hall.
Material examined. specimens; housed in the AMNH, BMS, GSC, NJSM, NYSM, USNM and the private collections of Larry De- cina and Paul Zell.
CONULARIA ELEGANTULA Meek, 1871 Figs. 3.4, 7. 1-7.4
Conularia elegantula Meek, 1871, p. 85-86; Meek, 1873, p. 228-229, PI. 23, fig. 4;
Miller, 1877, p. 141; Bigsby, 1878, p. 78; Whitfield, 1882, p. 242; Miller, 1889, p.
390; Babcock and Feldmann, 1986, fig. 2K.
Conularia s.l. elegantula Meek, Sinclair, 1948, p. 283.
33017, same specimen as in Fig. 8.1, detail of a minor face. 8.4; NYSM 3488, same specimen as in Fig. 8.2, detail of a minor face. 8.5; NYSM 3490, holotype of C. lata Hall, detail of major face; locality 122. 8.6; NYSM 3490, same specimen as in Fig. 8.5, entire specimen, preserved as an external mold in fine-grained sandstone. 8.7; AMNH 33018; paralectotype, preserved in three dimensions, major face; locality 1 17. 8.8; AMNH 33018, same specimen as in Fig. 8,7, comer view. 8.9; AMNH 33018, same specimen as in Fig. 8.7, minor face. Bar scales represent 1 cm.
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Description.— Exoskoidon up to 7 cm in length. Major apical angle 20-30°; minor apical angle about 1 7°. Rod articulation uniformly of angulated circular curve style. 6 nodes/mm on rods. 6 adapertural spines/mm; 6 adapical spines/mm. Rods usually abut at midline; approximately 40% alternate left superior on major and minor faces; rod angle 3-13°. 32-39 rods/cm. Apical wall not observed.
Occurrences. — Middle Devonian of Ohio; localities 174-176.
7>/?^.-Holotype, AMNH CU 282G.
Remarks. Conularia elegantula Meek is similar to C pyramidalis Hall in the values for rod angles, the number of nodes/mm on the rods and in the number of spines/mm. Conularia elegantula can be distin- guished from C pyramidalis, and, indeed, all other species of Conularia from the Devonian or Mississippian of North America by its rod ar- ticulation architecture, which seems to be uniformly of angulated cir- cular curve style. Partial specimens of C. desiderata exhibiting rods from the apertural region can also be distinguished from specimens of C elegantula by the curvature of the rods as they approach the comer angles. The rods of specimens of C elegantula are noticeably more inflected in the apertural direction than are rods belonging to specimens of C. desiderata.
Material examined. —ThtCQ specimens; housed in the AMNH and the CMNH.
CONULARIA MIL WA UKEENSIS Cleland, 1911
Figs. 7.5-7.9
Conularia congregata var. milwaukeensis Cleland, 191 1, p. 130, pi. 26, figs. 4-7; Teller, 1911, p. 251.
Mesoconularia milwaukeensis Sinclair, 1948, p. 119.
Conularia congregata milwaukeensis Cleland. Munthe, 1980, p. 6.
Description. — Exoskddon up to 5 cm in length. Major apical angle 12-15°; minor apical angle 11-14°. Rod articulation uniformly of inflected gothic arch style. Rods abut at midline; rod angle 5-18°. 18-24 rods/cm. 5-7 nodes/mm; 5-7 adapertural spines/ mm; 5-7 adapical spines/mm. Apical walls not observed.
Occurrence. —Middle Devonian of Wisconsin; localities 253-255.
7>/?c5. — Holotype, USNM 85988; five paratypes, USNM 78212. Two paratypes, listed by Sinclair (1948) as MPM 244-245, could not be found.
Remarks. — Conularia milwaukeensis may be distinguished from similar appearing species such as C. elegantula Meek and C. desiderata Hall by having only inflected gothic arch style rod articulation and by attaining lengths of up to 5 cm, apparently without the addition of apical walls. Conularia elegantula has narrow apical angles like C. milwaukeensis, but its rod articulation style is exclusively angulated circular curve. Likewise, C. desiderata has narrow apical angles, but
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its rod articulation patterns include inflected circular curve style in the apical region and angulated circular curve style elsewhere.
None of the examined specimens of C milwaukeensis preserves in- tegument over the entire exoskeleton. Because of this, rods have ob- viously been moved slightly from their original positions, making the measurement of rod angles difficult. Also, because of the general lack of preserved integument in most specimens, there is no evidence that apical walls were developed in this species. Apical walls may have been a feature of this species, but simply have not been preserved in the samples studied.
Material examined, — 2>1 specimens; housed in the GSC, MPM, and the USNM.
CONULARIA MULTICOSTATA Meek and Worthen, 1865 Figs. 9.1-9.5, 9.8, 10.1, 10.4, 12.1-12.5
Conularia multicostata Meek and Worthen, 1865, p. 252-253; Bigsby, 1878, p. 316;
Miller, 1889, p. 390; Weller, 1898, p. 190; Babcock and Feldmann, 1986, figs. lA, IF.
Conularia micronema Meek, 1871, p. 84; Meek, 1875, p. 316, PI. 18, figs, la-d; Miller, 1877, p. 141; Bigsby, 1878, p. 316; Herrick, 1888<3. p. 94-95, PI. 2, figs. 18-19; Herrick, 1888Z?, p. 49, PL 8, figs. 4-4a; Lesley, 1889, p. xv; Miller, 1889, p. 390, fig. 643; Herrick, 1893, PI. 19, figs. 4-4a; Weller, 1898, p. 190; Grabau and Shimer, 1910, p. 13, figs. 1227c-f.
Mesoconularia multicostata (Meek and Worthen). Sinclair, 1948, p, 125. Mesoconularia micronema (Meek), Sinclair, 1948, p. 124.
Mesoconularia mcfarlani Sinclair, [1948], p. 126-128, PI. 16, figs. 3-5.
Mesoconularia attica Sinclair, [1948], p. 125-126, PI. 9, fig. 2, PI. 17, figs. 9-11. Diconularia micronema (Meek). Sinclair, 1952, p. 138-139; Moore and Harrington, 1956Z), p. F61, fig. 47.2.
Non Conularia trentonensis multicosta Ruedemann, 1912, p. 115-116; Ruedemann, 1930, p. 36; Goldring, 1935, p. 63.
Description.— Exosk&leion up to 25 cm in length. Major apical angle 20-24®; minor apical angle 1 8-22®. Rod articulation of inflected gothic arch style in apical and most other regions and of angulated circular curve style in apertural region of large specimens; rods exhibiting inflected circular curve style articulation are mildly recurved near the midline but they are angulated in the apertural direction at the midline. Rods abut at midline; rod angle 9-17°. 26-60 rods/cm. Nodes prominent; 2-3 nodes/mm; 2-3 ad- apertural spines/mm; 2-3 adapical spines/mm. Apical wall not observed.
Occurrences. — Mississippian of Indiana, Kentucky and Ohio;
localities 25, 71=73, 79-80, 193, 197-200, 205, 209, 214, 217-218,
223, 225, 227 and 228.
7>/7e'5'. — Holotype of Conularia multicostata Meek and Worthen is lost; plastoholotype, with small fragments of fossil adhering, USNM 50157. Holotype of C. micronema Meek is apparently lost (Sinclair, 1948, p. 124); neotype, AMNH 6713. Specimen intended by Sinclair (1948) to be holotype of Mesoconularia mcfarlani, UK 6089. Three
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specimens which Sinclair (1948) intended to designate as types of M attica, CM 34533-34534, GSC 87204.
Remarks. — Conularia multicostata Meek and Worthen is most sim- ilar in morphology to C subcarbonaria Meek and Worthen. Like C subcarbonaria, C. multicostata has prominent nodes and can have more than 30 rods/cm. However, C. multicostata can be distinguished from this and all other species of Conularia by the combination of its very closely spaced rods, 26-60/cm, and its inflected gothic arch rod artic- ulation in most places except the apertural region of large specimens. More importantly, though, the rods of specimens referred to C sub- carbonaria appear to be undulose. This is especially true for large individuals (for example, Figs. 11.2-1 1.3).
In the apertural region of specimens of C. multicostata which are large for the species, a unique form of angulated circular curve rod articulation is present. This rod articulation pattern involves slight recurvature beginning about 72 to V3 of the distance between the midline and the comer angle; at the midline, the rods are slightly angulated. Approximately 70% of specimens of C multicostata are found to have well developed exoskeletal constrictions (for example. Figs. 9. 3-9. 5, 9.8, 10.1). These cannot be used as a species-level or genus-level taxo- nomic criterion, however, since well preserved exoskeletal constrictions have been observed in specimens belonging to nearly every taxon re- ported herein and in Part B. This species is notable because it shows good examples of this structure more frequently than any other known species of conulariid from the Devonian or Mississippian of North America.
According to Meek and Worthen (1865, p. 253), C. multicostata is unique among conulariids in that it possesses rods so closely spaced that “it is only under a good magnifier that the very minute crenulations can be seen.” Later, Meek (1871, p. 84) described C micronema as a
Fig. 9. -—9. 1-9.5; Conularia multicostata Meek and Worthen. 9.1; USNM 50157, plas- toholotype. Dark areas on photograph indicate areas where integument of the original specimen still adheres; locality 203. 9.2; UK 6089, specimen intended by Sinclair (1948) to be the holotype of Mesoconularia mcfarlani Sinclair, preserved in siderite concretion; probably minor face; locality 73.9.3; AMNH 6713, specimen chosen by Sinclair (19421?) as neotype of C micronema Meek, major face; locality 225. 9.4; AMNH 6713, same specimen as in Fig. 9.3, comer view. 9.5; AMNH 6713, same specimen as in Fig. 9.3, minor face. 9. 6-9. 7; C. subcarbonaria Meek and Worthen. 9.6; FMNH UC 6610, ho- lotype of C intertexta Miller; detailed view of exoskeleton, locality 28. 9.7; FMNH UC 6610, same specimen as in Fig. 9.6, view of entire specimen. 9.8; C multicostata Meek and Worthen, AMNH 6713, same specimen as in Fig. 9.3, detailed view of minor face. Bar scales represent 1 cm.
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Fig. 10,— 10J--10.3; Conuiaria muiticostata Meek and Worthen, 10.1; USNM 50128, comer view of specimen with pronounced exoskeletal constrictions; locality 225, 10,2-
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conulariid distinct from all others because it possessed rods so closely spaced that “it requires the aid of a magnifier to see them distinctly.” In all morphological respects, the two species are remarkably similar, assuming that AMNH 6713 is “typical” of the species C micronema as suggested by Sinclair (1948, p. 142). Conularia micronema Meek is here considered a junior synonym of C multicostata Meek and Wor- then.
Two manuscript species, Mesoconularia mcfarlani Sinclair and M. attica Sinclair, are here placed in synonymy with C multicostata for the reason that they have very closely spaced rods and nodes. Values for spacing of the rods and nodes are consistent with other specimens referred to C. multicostata (see Appendix B in part B). The specimen which Sinclair intended to designate the holotype of M. mcfarlani (Fig. 9.2) possesses an angulated circular curve style of rod articulation and is very similar in morphology to specimens from the Mississippian of Ohio which have been referred herein to C multicostata. The same can be said for the three specimens (CM 34533-34534, GSC 87204) referred to as M. attica in Sinclair’s (1948) unpublished thesis.
Conularia micronema was used by Sinclair (1 952, p. 138) as the type species of the genus Diconularia. Sinclair (1952, p. 138-139) noted that Diconularia was a probable form-genus which differs from Con- ularia in routinely having closely spaced rods and an “accentuation of the pustules.” Conularia, by contrast, was judged by Sinclair to exhibit these features only in gerontic specimens. The genus Diconularia is here included as a junior subjective synonym of Conularia because there does not appear to be any consistent pattern of differences between “Z). ” micronema and species included by Sinclair in the genus Conu- laria. Examples supporting this argument are given below.
Conularia niagarensis Hall, the North American reference species for the genus Conularia (Sinclair, 1940^z), has large, well-pronounced nodes and rod spacing values in the range of about 12-24/cm. These characters closely resemble those of the “typical” specimen of “D.” micronema, AMNH 6713 (see Appendix B in Part B). The number of rods/cm exhibited in AMNH 6713, 28-32, is well within the limits of the genus Conularia as recognized herein. Species referable to Conu-
10.3; C. subcarbonaria Meek and Worthen. 10.2; UIPC 10680, holotype, minor face, preserved in limestone; locality 13. 10.3; UIPC 10680, same specimen as in Fig. 10.2, detail of minor face. 10.4; C. multicostata Miller and Gurley, USNM 50647, apical region of specimen showing apical wall; locality 228. 10.5; C. subcarbonaria Meek and Worthen, UIPC 10680, same specimen as in Fig. 10.2, major face. Bar scales represent 1 cm.
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Fig. ll. — Conularia subcarbonaria Meek and Worthen. 11.1; FMNH UC 6289, badly weathered holotype of C. spergenensis Miller and Gurley, preserved in limestone; locality
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laria from the Devonian and Mississippian of North America exhibit 9 to 84 rods/cm. Furthermore, no species of Conularia examined from Devonian or Mississippian age rocks of North America are known to consistently possess rods which are more closely spaced in large, pre- sumably gerontic, individuals than in “average-sized” individuals.
Material examined. — 61 specimens; housed in the AMNH, CM, CMNH, FMNH, GSC, and the USNM and the private collection of Ron Fisher.
CONULARIA PYRAMIDALIS Hall, 1859 Figs. 3.3, 8.1-8.9, 16.7
Conularia pyramidalis Hall, 1859, p. 347-348; Hall, 1861, PI. 72A, figs, la-c; Miller, 1877, p. 141; Miller, 1889, p. 390; Whitfield and Hovey, 1899, p. 170-171; Rich- ardson, 1942, p. 23-26, PI. 3, figs. 4-9; Sinclair, 1948, p. 106; Babcock and Feld- mann, 1986, fig. 2E.
Conularia huntiana Hall, 1859, p. 348; Hall, 1861, PI. 72A, figs. 2a“b; Clarke and Ruedemann, 1903, p. 566; Grabau, 1906, p. 151, fig. 65; Grabau and Shimer, 1910, p. 13, fig. 1224; Sinclair, 1948, p. 106, PI. 9, fig. 1.
Conularia lata Hall, 1859, p. 479-480, PI. 70A, fig. 3, PI. 91, fig. 1; Bigsby, 1878, p. 78;
Miller, 1889, p. 390; Sinclair, 1948, p. 104.
Conularia huntana (sic) Hall. Miller, 1877, p, 141; Miller, 1889, p. 390.
Description.— Exoskelcton up to 20 cm in length. Major apical angle 17-23°; minor apical angle 16-17°. Rod articulation gothic arch style in apical region and inflected gothic arch elsewhere. Rods abut at midline; rod angle 5-22°. 7-17 rods/cm. 1-4 nodes/ mm; 1-4 adapertural spines/mm; 1-4 adapical spines/mm. Apical wall not observed.
Occurrences.— Lower Devonian of New Jersey and New York; lo- calities 114, 116-120 and 122.
Types.— LeeXoXype, designated herein from James Hall’s syntypic suite of four specimens, AMNH 33018, smaller of two specimens bear- ing this number (Figs. 3.3, 8.1, 8.3); three paratypes, AMNH 33018, larger specimen, and AMNH 33019, two specimens. Holotype of C huntiana Hall, NYSM 3488; holotype of C. lata, NYSM 3490, plas- toholotype, GSC unnumbered.
Remarks. — Conularia pyramidalis Hall is similar in size and apical angles to C. desiderata Hall. Distinction between the two taxa is made on the basis of differences in rod angles, 9-14® for C. pyramidalis versus 13-17® for C. desiderata. Additionally, when complete enough speci-
44. 11.2; FMNH UC 18494, larger of two specimens, flattened specimen with apical wall preserved, preserved in calcareous shale; locality 38. 1 1.3; FMNH UC 18494, same specimen as in Fig. 11.2, detail of major face. Note apparent convergence of rods at exoskeletal constriction. 11.4; UIPC 10680, same specimen as in Fig. 10.2, detail in region of comer groove; locality. 1 1.5; FMNH UC 6289, same specimen as in Figure 11.1, detail of exoskeleton. Bar scales represent 1 cm.
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mens are studied, distinction between these two species can be made on the basis of rod articulation style, gothic arch to inflected gothic arch for C pyramidalis, compared to gothic arch to inflected circular curve for C. desiderata. Differences in rod articulation styles imme- diately serve to help distinguish C pyramidaiis Hall from C undulata Conrad, though both may attain lengths greater than 1 5 cm. Conularia pyramidaiis Hall is readily distinguished from C. elegantuia Meek by having significantly fewer nodes and spines per unit space than are present in C. pyramidaiis. The feature which immediately serves to distinguish C. pyramidaiis from C multicostata Meek and Worthen is the greater spacing between rods, 9- 14/cm in C pyramidaiis and up to 32/cm in C. multicostata. Conularia multicostata and C subcar- bonaria Meek and Worthen have more prominent nodes on the rods than does C pyramidaiis.
Conularia huntiana Hall was distinguished from C pyramidaiis Hall (Hall, 1859, p. 348) by its greater length, its smaller apical angles, the greater convexity of its faces and differences in the appearance of ridges and spines. The apical angles of the lectotype of C pyramidaiis are 1 8® and 16°, and in the holotype of C. huntiana, the apical angles as mea- sured are 13° and 11°. However, the holotype of C. huntiana is not compressed to the extent that the lectotype of C pyramidaiis is. More- over, the lectotype of C. pyramidaiis is a smaller specimen and pre- sumably represents an earlier growth interval than does the holotype of C huntiana. Therefore, a smaller set of apical angles is expected in C. huntiana. Differences in convexity of the faces between the two taxa is likely a result of differences in collapse of the exoskeletons after death of the animals and/or differences in compression of the exoskeletons. Differences in the appearance of the ridges and spines seem to be functions of preservation: most of the lectotype of C pyramidaiis is preserved as an internal mold, whereas the holotype of C huntiana retains much integument. Where integument is present on the lectotype of C pyramidaiis, it is identical to that on the holotype of C. huntiana (compare Figs. 8.3 and 8.4). In both specimens, most of the rods are broken out, leaving a “double ridge” arrangement of the integument,
Fig. 12. — Conularia multicostata Meek and Worthen. 12.1; GSC 87204, enlargement of a specimen intended by Sinclair (1948) to be a paratype of Mesoconularia attka; locality 193. 12.2; CMNH 4684, external mold of a flattened specimen preserved in a siderite concretion; locality 198. 12.3; CM 34533, specimen intended by Sinclair (1948) to be the holotype of M attka; locality 193. Note that the apical wall is present. 12.4; FMNH UC 540 14A, specimen figured by Herrick (1888a) as C. micronema; locality 228. 12,5; GSC 87204, same specimen as in Fig. 12.1, view of a collapsed specimen preserved in a siderite concretion; locality 193, Bar scales represent 1 cm.
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marking the former positions of the margins of the rods within the integument.
Conularia lata Hall is included as a junior subjective synonym of C pyramidalis largely because of qualitative similarities including near- ly identical rod articulation styles. Also, values for apical angles and rod spacing are similar in the lectotype of C pyramidalis and the sole specimen referred to C. lata by Hall, NYSM 3490. Major and minor apical angles for the lectotype of C. pyramidalis are 18° and 17°, re- spectively. For the holotype of C. lata, they are 23° and 17°. Rods/cm values for the lectotype of C. pyramidalis vary between 9 and 14 while those of the holotype of C lata vary between 7 and 10. Values for rod angles and node spacing on the rods for both specimens are also well within the expected normal distribution for a species.
Material examined. ~A\ specimens; housed in the AMNH, CM, CMNH, FMNH, GSC, NJSM, NYSM, USNM, and the private col- lection of Paul Zell.
CONULARIA SUBCARBONARIA Meek and Worthen, 1865 Figs. 9.6-9.7, 10.2-10.3, 10.5, 11.1-11.5
Conularia subcarbonaria Meek and Worthen, 1865, p. 253; Meek and Worthen, 1873, p. 520-522, PI. 19, figs. 4a-c; Miller, 1877, p. 141; Bigsby, 1878, p. 316; Miller, 1889, p, 390; Keyes, 1894, p. 218; Weller, 1898, p. 191-192.
Conularia intertexta Miller {nomen nudum), \%91b, p. 692.
Conularia intertexta Miller, 1894, p. 317, PI. 10, fig. 4; Weller, 1898, p. 190.
Conularia spergenensis Miller and Gurley, 1893, p. 74-75, PL 8, fig. 2; Miller, 1897, p. 765; Weller, 1898, p. 191.
Mesoconularia subcarbonaria (Meek and Worthen). Sinclair, 1948, p. 123. Mesoconularia intertexta (Miller). Sinclair, 1948, p. 123.
Diconularia micronema (Meek). Sensu Lane, 1973, p. 92-93, PI. 8, figs. 2-3.
Description.— ExoskeXQlon up to 35 cm in length. Major apical angle 10-21°; minor apical angle 9-19°. Rods are undulose; rod articulation uniformly of angulated circular curve style. Rods abut at midline; rod angle 4-6°. 1 7-3 1 rods/cm. Nodes prominent; 3- 5 nodes/mm on rods; no spines present. Apical wall may be present.
Occurrences. — Upper Devonian-Lower Mississippian of Illinois, In- diana, Iowa and Missouri; localities 8-9, 13, 20, 24, 27-28, 37-38, 42, 44^46, 48, 51, 61, 64, 67, and 101.
Holotype, UIPC 10680, plastoholotypes USNM 50158, FMNH UC unnumbered and GSC unnumbered. Holotype of C. in- tertexta, FMNH UC 6610, plastoholotypes, USNM 68130 and GSC unnumbered; holotype of C. spergenensis, FMNH UC 6289.
Remarks. — In the possession of prominent nodes along the rods, C. subcarbonaria Meek and Worthen resembles the Mississippian species, C. multicostata Meek and Worthen, and the Devonian species, C. ulsterensis Howell. These taxa are readily distinguished, however, on the basis of rod articulation style: C subcarbonaria possesses angulated
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circular curve rod articulation in which the rods are undulose, partic- ularly in large individuals. On the other hand, C muiticostata has both angulated circular curve and inflected gothic arch rod articulation and C ulsterensis possesses only inflected gothic arch rod articulation. Ad- ditionally, C ulsterensis is unique among the species of Conularia studied in having a rod spacing of 45-84 rods/cm in adult specimens.
Like the Devonian species, C undulata Conrad, C subcarbonaria possesses undulose rods. Conularia subcarbonaria can be distinguished from C undulata in having more prominent nodes along the rods and by having rods which nearly always abut at the midline. The rods of C undulata alternate at the midline in approximately 1 0% of the cases studied.
Conularia intertexta Miller is considered synonymous with C sub- carbonaria because of general qualitative similarities between the types of the two species and because of similarities in the value for number of rods/cm (see Appendix B in Part B). However, the holotype specimen (Figs. 9. 6-9. 7) of C. intertexta is very badly preserved, maHng any species-level assignment somewhat doubtful. One characteristic of the specimen is an apparently undulose set of rods. It is primarily because of its undulose rods that C intertexta is here included in synonymy with C subcarbonaria.
The holotype of C. spergenensis Miller and Gurley (Figs. 11.1, 1 1.5) is, like the holotype of C intertexta, badly preserved. However, like the holotype of the latter taxon, C spergenensis bears qualitative and quantitative similarities to C subcarbonaria, most notably, undulose rods. It too is therefore considered to be a junior subjective synonym of C subcarbonaria.
Material examined. —46 specimens; housed in the AMNH, FMNH, GSC, ISGS, lUPC, and the USNM.
CONULARIA rC/ZOJ Clarke, 1907 Fig. 15.3
Conularia desiderata var. tuzoi Clarke, 1907, p. 181, fig.; Clarke, 1908, p. 144, PL 11, fig. 13; Dresser and Denis, 1944, p. 326.
Conularia tuzoi Clarke. Sinclair, 1948, p. 105,
Description. — based only upon holotype. Exoskeleton 1 1.3 cm in length. Major apical angle approximately 10®; minor apical angle not observed. Rod articulation style unknown in vicinity of apex and of inflected gothic arch style elsewhere; rods are broadly inflected. Rods abut at midline; rod angle 9-10®. Nodes and spines not observed. Apical wall not observed.
Type —Holotype, NYSM 9404.
Occurrence. — CowQT Mississippian of Quebec; locality 244.
Remarks. — The holotype of C tuzoi (Clarke) is badly preserved and no additional specimens are known. A cross sectional view is not pre-
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Fig. \3. — Conularia undulata Conrad. 13,1; AMNH 41093, neotype, preserved as an external mold in siltstone. Note rounded marks produced by orbiculoid brachiopods
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served on the holotype. This taxon appears to differ from other species of Conuiaria included in this paper in the possession of broadly rounded inflections where the rods are of inflected gothic arch style.
Material examined, — 1 specimen, NYSM 9404.
CONULARIA ULSTERENSISllowGll, 1942 Figs. 15.4, 16.1-=16.3, 16.5
Conuiaria ulsterensis Howell, 1942, p. 91, figs. 10-11; Goldring, 1943, p. 208. Mesoconularia ulsterensis (Howell). Sinclair, 1948, p. 117.
D^5m>£ioii.““Exoskdeton up to 3 cm in length. Major apical angle 14-19°; minor apical angle 10-15°. Rod articulation uniformly of inflected gothic arch style. Rods usually abut at midline; rod angle 1 1-15°. 45-84 rods/cm. Nodes prominent; 6-8 nodes /mm; 6-8 adapertural spines/mm; 6-8 adapical spines/mm. Apical wall not observed.
Typex— Holotype, PU 42071; two paratypes, PU 42072-42073.
Occurrences. — 'Lowqt Devonian of New York and Pennsylvania; lo- calities 115, 231, 234 and 236. Specimens possibly referable to this taxon have also been found in the Lower Devonian of Quebec; locality 242.
Remarks. — This species is distinct in its possession of up to 84 rods/ cm in adult specimens, the largest number recorded in the genus Con- uiaria. It can also be distinguished by its prominent, closely spaced nodes. In this form, nodes are spaced as closely as 7/mm. Some spec- imens of C undulata Conrad may have as many as 7 nodes/mm, but C undulata has neither inflected gothic arch rod articulation through- out the exoskeleton nor prominent nodes.
The specimen of C. ulsterensis Howell illustrated in Fig. 16.2 is preserved as an internal mold. It is unusual in that it clearly shows that a longitudinal invagination existed along the integument internal to the midline.
Material examined.--! specimens; housed in the CM, NJSM, and the NYSM.
CONULARIA UNDULATA Conrad, 1841
Figs. 13.D13.5, 14.1-14.5, 15.1-15.2, 16.4
Conuiaria undulata Conrad, 1841, p. 57; Hall, 1861, p. 62-63; Bigsby, 1878, p. 62-63; Hall, 1876, PL 29, figs. 1-7; Hall, 1879, p. 208-209, PI. 33, figs. 1-5, 7; PL 34A, figs. 1-4; Miller, 1889, p. 390; Whitfield and Hovey, 1901, p. 326“”327; Grabau,
that were previously attached to the conulariid exoskeleton; locality 145. 13.2; NYSM 3493, preserved in siltstone, major face; locality 145. 13.3; NYSM 3494, preserved in siltstone, major face; locality 145. 13.4; AMNH 41093, same specimen as in Fig. 13.1, detail of major face. 13.5; AMNH 41093, same specimen as in Fig. 13.4, detail in region of comer groove. Bar scales represent 1 cm.
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Fig. 14. — Conularia undulata. 14.1; AMNH 5439, specimen showing relation of apical wall to the remainder of the exoskeleton. Note flange for connection at adapertural end
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1899, p. 284-285, fig. 219; Grabau and Shimer, 1910, p. 13, fig. 1225-1226; Moore and Harrington, \956b, p. F60, fig. 25.2; Babcock and Feldmann, 1986, fig. 21. Conularia cayuga Hall, 1876, PL 28, figs. 2-3; Hall, 1879, p. 211-212, PI. 34, figs. 2, 5; Miller, 1889, p. 390.
Conularia crebistria Hall, 1876, PI. 29, figs. 8, 9.
Conularia crebistriata {sic) Hall, 1879, p. 210-211, PI. 33, figs. 8-9, PL 34A, fig. 5; Miller, 1889, p. 390; Moore and Harrington, 1956A p. F60, fig. 42.9.
Mesoconularia undulata (Conrad). Sinclair, 1948, p. 118.
Conularia s.l. cayuga Hall. Sinclair, 1948, p. 285.
Ctenoconularia crebistriata {sic) (Hall). Sinclair, 1948, p. 241, PL 6, figs. 10-11. Ctenoconularia crebistria (Hall). Sinclair, 1952, p. 142.
Conularia sp. cf. C. undulata Conrad. Kasznica, 1986, p. 14-15, fig. 2.
Description.— Exos\ie\Qion up to 15 cm in length. Major apical angle 10-18°; minor apical angle 8-15°. Rod articulation uniformly of inflected circular curve style; rods are undulose in the apertural Vi. Rods usually abut at midline; rod angle 10-18° in apical region and 4-20° elsewhere. 17-32 rods/cm. 4 nodes/mm; 4 adapertural spines/mm; 4 adapical spines/mm. Apical wall may be present.
Neotype, AMNH 41093, plastoneotype, FMNH UC 694; James Hall’s figured specimens, NYSM 3493, 3494, AMNH 5439. Holotype of C. crebistria, AMNH 5440, plastoholotype, FMNH UC 679; holotype of C. cayuga, NYSM 3482, plastoholotype, FMNH UC
685.
Occurrences. Devonian of Maine, Maryland, New York, Ontario, Pennsylvania, and Quebec; localities 90-91, 115, 128-129, 136-138, 140-141, 143, 145-148, 230, 236-238, and 241. Ulrich (1892) has indicated that C undulata is present in the Devonian of Bolivia and Reed (1904) has cited this taxon in the Devonian of South Africa. These identifications are erroneous, and are described briefly below. They will also be described in greater detail elsewhere. Cordini (1955, p. 275, fig. 81) referred and figured some fossils found in Antarctica as C. cf. C. undulata, but these have been subsequently identified as plant remains (Dalziel et al., 1981).
Remarks.— Contdi&s, suite of syntypes is lost. However, judging from his description of C. undulata (Conrad, 1841, p. 57) it is clear that the species is based upon specimens now referable to either C. undulata or C. pyramidalis Hall. Hall’s early figures and description of C. un- dulata (Hall, 1876, Plate 29, figs. 1-7, explanation of Plate 29; 1879, p. 208-209; Plate 33, figs. 1-5, 7, Plate 34A, figs. 1-4) have served as bases for all subsequent studies on the species. Therefore, it is appro-
of apical wall; locality 145. 14,2; AMNH 5440, holotype of C. crebistria Hall, preserved in calcareous shale, minor face; locality 136. 14.3; AMNH 5440, same specimen as in Fig, 14.2, detail of minor face. 14.4; NYSM 3482; holotype of C. cayuga Hall, preserved in calcareous shale; locality 136. 14.5; NYSM 3482, same specimen as in Fig. 14.4, detailed view of minor face. Bar scales represent 1 cm.
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Fig, 15.-15.1-15.2; Conularia cf. C undulata Conrad. 15.1; NYSM 9410, detail of exoskeleton; locality 241, 15.2; NYSM 9408, flattened and tectonically distorted spec- imen; locality 90. 1 5.3; C tuzoi Clarke, NYSM 9404, holotype; locality 244. 1 5.4; NYSM
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priate to select a neotype from among HalFs extant specimens. The specimen chosen, AMNH 41093, is the subject of one of the best known of Hairs figures of the taxon.
Two Devonian species are here considered junior subjective syn- onyms of C undulata Conrad: C cayuga Hall and C crebistria Hall. Conularia crebistria, according to Hall (1879, p. 210) differed from C undulata in being “more slender in its mode of growth” and in having more closely spaced rods. In terms of the arrangement of nodes and spines, the two were judged to be “precisely similar.” The holotype of C crebistria, AMNH 5440, is a badly preserved specimen exhibiting one face (Figs. 13.2-13.3). The apical angle cannot be determined be- cause the specimen has been distorted and the comer grooves are not preserved on the surface of the slab. Like specimens of C. undulata, this specimen has inflected circular curve rod articulation. The rods are undulose in the vicinity of the aperture. The holotype of C. crebistria has rod spacing values ranging from 24 to 30 rods/cm and node spacing values of about 4 nodes/mm. No other values can be determined with confidence from this specimen. The equivalent values for the lectotype of C. undulata are 20-27 rods/cm and 6 nodes/mm. There seems to be no significant difference in the features exhibited in AMNH 5440 from other specimens here referred to C. undulata.
The holotype of C cayuga, NYSM 3482, is a flattened specimen preserved as an external mold (Figs. 13.4-13.5). An apical wall is present. Hall (1879, p. 211-212) indicated that, in general, this spec- imen is “not dissimilar to C undulata. ” However, subtle differences, including “stronger” rods, wider spaced rods except in the apertural region and the presence of “striae” between adjacent rods (=spines) were used as key characters which served to distinguish this taxon from C. undulata. Spines, of course, are present in C. undulata, just as they are in the holotype of C. cayuga. The rods of NYSM 3482 have a spacing of 17-22/cm, well within the expected range of values for individuals of C undulata. Other quantitative determinants, given in Appendix B in Part B, substantiate this conclusion. The rods are un- dulose except near the apical wall and are articulated in angulated circular curve style, similar to the neotype of C undulata. The rods do not appear to be better pronounced than those shown in specimens referred to C. undulata which are preserved as external molds, including the neotype (Fig. 1 3.4). Thus, the holotype of C. cayuga is here referred to Conrad’s species, C undulata.
9411, C. ulsterensis Howell, two specimens, one preserving an apical wall; locality 242. Bar scales represent 1 cm.
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Fig. 16.— 16.1“16.3; Conularia ulsterensis Howell. 16.1; PU 420715 holotype; locality 115.1 6.2; PU 42072, paratype; internal mold showing longitudinal ridges at the midlines; locality 115. 16.3; NJSM 12843, external mold; locality 236. 16.4; CM 34520, C un-
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Sinclair (1948, p. 118) and Sinclair and Richardson (1954, p. 105) stated that the species C grandis Roemer is synonymous with C. un- dulata. However, this cannot be substantiated here because the spec- imen upon which Roemer’s description is based (Roemer, 1856, p. 436, Plate 3, figs. 21a-b) was not available for study. It is likely that the holotype of C. grandis is lost.
Conularia undulata is similar to C subcarbonaria Meek and Wor- then in the possession of an undulose mode of angulated circular curve style rod articulation. It differs from the latter form in having nodes on the rods which are not prominent. Moreover, the rods of C sub- carbonaria nearly always abut at the midline. In C undulata, as many as 10% of the rods may alternate at the midline.
Two specimens from the Devonian of the Malvinokaffric Realm have been misidentified as C. undulata. The first (Ulrich, 1892, p. 31-33, PI. 3, figs. 6a-b), which was collected in Bolivia, is referable to C albertensis Reed, judging from Ulrich’s well-executed figure. Conularia albertensis differs from C. undulata in having gothic arch rod articu- lation in the apical region and inflected gothic arch rod articulation elsewhere, whereas C undulata has only inflected circular curve rod articulation. The second specimen from the Malvinokaffric Realm which was misidentified as C undulata was described and figured by Reed (1904, p. 248-249, PL 31, figs. 1-la). It was collected from the Bok- keveld beds of South Africa. A latex mold (UCGM 34720) of the specimen has been examined. It is referable to C quichua Ulrich. Conularia quichua and C. undulata both have undulose rods, but C quichua has rods articulated in gothic arch fashion in the apical region and in angulated circular curve style elsewhere.
Material examined.— 43 specimens housed in the AMNH, CM, FMNH, GSC, NYSM, NJSM, USNM, and the private collections of Gordon Baird, Robert Tinsley and Paul Zell.
CONULARIA sp.
Fig. 16.8
Conularia cf. huntiana Hall. Merriam, 1973, p. 35, PL 12, figs. 18-20.
Occurrence.— Upper Devonian of Nevada; Locality 108.
Remarks.— Remains of at least six conulariid specimens from the Devonian of Nevada were figured and described by Merriam (1973,
dulata Conrad, portion of specimen with orbiculoid brachiopod attached; locality 91. 16.5; C. ulsterensisYiov<iQ\\, CM 34528. 16.6; C. desiderataUdlX, USNM 395827, juvenile, with apex intact; locality 142. 16.7; C. cf. C. pyramidalis Hall, GSC 2598; locality 241. 16.8; C sp., USNM 159536; locality 108. Bar scales represent 1 cm.
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p. 35, Plate 12, figs. 18-20). Merriam identified these specimens as C. cf. huntiana. The specimens (USNM 159536), one of which is figured herein (Fig. 16.8), are poorly preserved and cannot be assigned to a species at present.
The specimens in question possess an inflected circular curve rod articulation. The rods are slightly deflected adaperturally at the midline. This style of rod articulation is similar to that observed in specimens of C. desiderata Hall.
Material examined. — 6 specimens; housed in the USNM.
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Babcock, L. E., and R. M. Feldmann. 1984. Mysterious fossils. Earth Science, 37(3): 16-17.
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Barrande, J. 1867. Systeme silurien du centre de la Boheme. lere Partie. Tome 3.
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Bigsby, j. j. 1878. Thesaurus Devonico-Carboniferous. The flora and fauna of the Devonian and Carboniferous periods. . . London, 447 pp.
Boucek, B. 1939. Conularida. Pp. A1 13-A131, in Handbuch der Palaozoologie, Vol. 2A (O. H. Schindewolf, ed.), Berlin.
Boucek, B., and F. Ulrich. 1929. Etude sur la coquille du genre Conularia Miller. Statniho geologickeho ustavu Ceskoslovenske Republicky, Vestnik, Roc., 5(2/3): 1-25.
Branisa, L. 1965. Los fosiles guias de Bolivia. Servicio Geologico de Bolivia, Boletin
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Branisa, L., and J. Vanek. 1973. Metaconularia cahuanotensis sp. n. from Bolivian Lower Devonian. Vestnik Ustfedniho ustavu geologickeho, 48:95-96.
Branson, E. B. 1938. Stratigraphy and paleontology of the Lower Mississippian of Missouri. Part 1. University of Missouri Studies, 13(3), 205 pp.
1944. The geology of Missouri. University of Missouri Studies, 19(3), 535 pp.
Calvin, S. 1890. Note on a specimen of Conularia missouriensis Swallow, with cren- ulated costae. American Geologist, 5:207-208.
Caster, K. E. 1957. Problematica. In Treatise on marine ecology and paleoecology. Volume 2, Paleoecology (H. S. Ladd, ed.). Geological Society of America, Memoir, 67:1025-1042.
Clarke, J. M. 1907. Some new Devonic fossils. New York State Museum, Bulletin, 107 (Geology 12) (New York State Education Department, Bulletin, 401:153-291.
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1908. Early Devonic history of New York and eastern North America. New
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VOLUME 55 31 DECEMBER 1986 ARTICLE 16
DEVONIAN AND MISSISSIPPIAN CONULARIIDS OF NORTH AMERICA.
PART B. PARACONULARIA, RETICULACONULARIA,
NEW GENUS, AND ORGANISMS REJECTED FROM CONULARIIDA
Loren E. Babcock* ^
Rodney M. Feldmann*
Research Associate, Section of Invertebrate Fossils
Abstract
Descriptions of the species assigned to Pamconularia Sinclair, 1940 and Reticulacon- ularia Babcock and Feldmann, new genus, as well as organisms rejected from the Con- ulariida, are treated in Part B of this two-part work on the Devonian and Mississippian conulariids of North America. Fifteen species of Pamconularia are considered valid, of which five are new. The new taxa are P. alpenensis, P. chagrinensis, P. oklahomaensis, P. wellsvillia, and P. yochelsoni. Adesmoconularia Driscoll, 1963 is considered a junior synonym of Paraconularia. Two species are referable to Reticulaconularia Babcock and Feldmann, new genus; Conularia penouili is selected as the type species.
Introduction
This paper is the second, and final, part of “Devonian and Missis- sippian Conulariids of North America.” This work contains descrip-
‘ Address: Department of Geology, Kent State University, Kent, Ohio 44242.
2 Present address: Department of Geology, University of Kansas, Lawrence, Kansas 66045.
Submitted 2 June 1986.
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tions of species referable to the genera Paraconularia and Reticula- conularia n. gen., as well as specimens described in the literature as conulariids but which are here rejected from the phylum. Locality descriptions and measurements of selected specimens are included as appendices A and B, respectively, herein. Figures are numbered con- secutively in both Parts A and B in order to avoid cross-reference confusion.
Genus PARACONULARIA Sinclair, 1940
Type species. — Conularia inequicostata Koninck, 1883, designated by Sinclair (1940); Carboniferous of Belgium. Holotype: Musee Royal d’Histoire Naturelle de Belgique, Brussels, Belgium. North American reference species, Sinclair (1940): Conularia blairi Miller and Gur- ley, 1893 (Mississippian). Lectotype of C blairi: UCGM 3985.
Diagnosis. —Corml^xiids with rods that are generally widely spaced, 4-35 rods/cm. More than 60% of rods alternate at midline; fewer than 40% abut. Apical angles small, 9-28°. Nodes, adapertural spines and adapical spines may or may not be present; if present, they are usually widely spaced, 2-6/mm.
PARACONULARIA ALPENENSIS Babcock and Feldmann, new species Figs. 17.1-17.3
Description. — T)Q^cnpX\on based only upon holotype. Exoskeleton 3.6 cm in length. Major apical angle 21°; minor apical angle 16°. Rod articulation inflected circular curve style; rods are slightly recurved near midline in apertural region. Rods abut or alternate at midline; alternation pattern either right superior or left superior on major face, usually left superior on minor face; rod angle 9-10°. 14 rods/cm. Nodes not observed; spines absent. Apical wall not observed.
Occurrence.— Middle Devonian of Michigan; locality 92.
7>/7e -Holotype, GSC 85060.
Remarks.— taxon is similar to P. chesterensis (Worthen), P. missouriensis (Swallow) and P. recurvatus Babcock and Feldmann, n. sp. in the possession of rods that are recurved near the midline. Par- aconularia alpenensis, however, exhibits rods that are not recurved in the apical region, at least not in the holotype. None of the other three taxa possess this characteristic.
It is not known whether P. alpenensis possessed nodes on the rods. The holotype, and only known specimen, is weathered and lacks the external surfaces of all the rods which are present. No spines are present. The uniqueness of the rod articulation patterns are sufficient to distin- guish this taxon from all other described taxa.
Material examined. — \ specimen, GSC 85060.
1986 Babcock and Feldmann— and Reticulaconularia
413
Etymology of trivial —Named for the Alpena Limestone, in
which the holotype was found.
PARACONULARIA ALTERNI STRIATA (Shimer, 1926)
Figs. 17.5-17.6
Conularia alternistriata Shimer, 1926, p. 84, PL 4, figs. 1 la-b.
Paraconularia alternistriata (Shimer). Sinclair, 1948, p. 190.
Description.— YOQsctiption based only upon holotype. Length 1.9 cm. Major apical angle 1 1®; minor apical angle 10®. Rod articulation inflected gothic arch style; rods are almost imperceptibly inflected near the comer angles. Rods always alternate at midline; alternation pattern usually right superior on major face and usually left superior on minor face; rod angle 9-10°. 28 rods/cm (extrapolated). Nodes and spines absent. Apical wall not observed.
Occwrrpwc^. — Mississippian of Alberta; locality 4.
Type. — Holotype, GSC 5111.
Remarks. —Paraconularia alternistriata (Shimer) is similar in mor- phology to specimens of P. yochelsoni Babcock and Feldmann, n. sp. Both are of similar size, less than 3.5 cm in maximum length and both exhibit similar forms of rod articulation style. The rod articulation present on the holotype of P. alternistriata is here judged to be a form of inflected gothic arch style. The rods in this specimen are inflected so little, though, that the articulation could easily be confused for a gothic arch style articulation pattern. This may simply be a function of the small size of the holotype; a sample close to the aperture of a larger specimen may yield a rod articulation pattern more distinctly of an inflected gothic arch style. Paraconularia yochelsoni possesses rods which are clearly articulated in an inflected gothic arch style close to the apex and trending towards an inflected circular curve style near the aperture. Paraconularia alternistriata is further distinguished from P. yochelsoni in having a smaller apical angle, 10-11®, as compared to 1 5-20° in P. yochelsoni and, finally, in having greater rod spacing, 28 rods/cm as compared to 13-18 rods/cm.
The holotype, and only known specimen, of P. alternistriata exhibits longitudinal folds in the integument between adjacent rods suggesting that spines may have been present in this taxon. The folds are best developed near the comer angles. Their occurrence seems to be erratic and the spacing between adjacent folds is inconsistent. In all likelihood, these folds do not represent integument folded over spines but simply folds resulting from a contraction of the integument about the rods and, perhaps, some shearing of the exoskeleton due to compression. This phenomenon is relatively common among specimens of Para- conularia, having also been observed in P. subulata (Fig. 3.2) and P. missouriensis (Fig. 25.3).
Material examined. — 1 specimen, GSC 5111.
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1986 Babcock and Feldmann— PyiiMCOivt/i^i?//4 and Reticulaconularia
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PARACONULARIA BLAIRI (Miller and Gurley, 1893)
Figs. 18.1--18.5, 22.1=^22.3
Conularia biairi Miller and Gurley, 1893, p. 73-74, pi. 7, figs. 14-15; Miller, 1897, p. 765; Weller, 1898, p. 189; Chappars, 1936, p. 16; Branson, 1938, p. 110, PI. 14, figs. 7-8; Branson, 1944, p. 216.
Conularia sedaliensis Miller and Gurley, 1896, p. 28, PL 3, figs. 4-5; Miller, 1897, p.
765; Weller, 1898, p. 191; Chappars, 1936, p. 16.
Conularia {Conularia) sedaliensis Miller and Gurley. Boucek, 1939, p. A 121. Conularia (Paraconularia) biairi (Miller and Gurley). Sinclair, 1940, p. 74. Paraconularia biairi (Miller and Gurley). Sinclair, 1948, p. 197; Moore and Harrington, 1956, p. F65, fig. 50.2.
Paraconularia sedaliensis Miller and Gurley. Sinclair, 1948, p. 201.
Paraconularia indiana Sinclair, [1948], p. 195, PL 18, figs. 1-2.
Paraconularia cf. newberryi (Winchell). Sensu Sinclair, 1948, PL 13, figs. 1-3. Paraconularia missouriensis (Swallow). Sensu Babcock and Feldmann, 1984, p. 16-17.
Description. -~Exo%kQ\QXon up to 20 cm in length. Major apical angle 1 1-23°; minor apical angle 10-20°. Rod articulation inflected gothic arch style in apical region and inflected circular curve with a slight adapertural inflection at the midline elsewhere. Rods almost always alternate at midline; alternation pattern usually right superior on major and minor faces; rod angle 8-19°. 6-13 rods/cm. 2-3 nodes/mm; 2-3 adapertural spines/ mm; adapical spines absent. Apical wall not observed.
Types. —ThvQQ syntypes of C. biairi, UCGM 3984-3986, of which UCGM 3986 (Fig. 1 8.5) is here designated the lectotype; UCGM 3984- 3985 are here designated the lectotype; UCGM 3984-3985 are here considered paralectotypes. Four syntypes of C. sedaliensis, preserved in five pieces, UCGM 1393, 1399; specimen intended by Sinclair (1948) to be holotype of P. indiana, AMNH 25056.
Occurrences.— Towqt Mississippian of Illinois, Indiana, Iowa, and Missouri; localities 11, 32, 35, 61, 94-98, 101, and 105. Laudon and Bowsher (1941) reported this taxon in the Mississippian of New Mex- ico, but their material was not available for study.
Remarks.— Paraconularia biairi (Miller and Gurley) is unique among species of Paraconularia in having rods that exhibit inflected circular
Fig. 17.— 17.1; Paraconularia alpenensis Babcock and Feldmann, n. sp., GSC 85060, holotype, minor face of specimen preserved in micrite; locality 92. 17.2; GSC 85060, same specimen as in Fig. 17.1, comer view. 17.3; GSC 85060, same specimen as in Fig. 17.1, major face. 17.4; USNM 173926, Hyolithes sp., cmshed specimen of a hyolithid; locality 240. 17.5-17.6; P. alternistriata Shimer. 17.5; GSC 5111, holotype, major face; locality 4. 17.6; GSC 5111, same specimen as in Fig. 17.5, minor face. 17.7-17.8; P. chesterensis (Swallow). 17.7; GSC 85061, a collapsed specimen preserved in siltstone; locality 27. 17.8; GSC 85061, enlargement of same specimen as in Fig. 17,7. Note inconspicuous spines on the rods. Bar scales in Figs. 17.1-17.4 and 17.7-17.8 represent 1 cm; bar scales in Figs. 17,5 and 17.6 represent 5 mm.
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Fig. IH.—Paraconularia blairi (Miller and Gurley). 18.1; UMC 4270, detailed view of well preserved ?major face; locality 96, 18.2; UCGM 3985, syntype of Conularia se-
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curve style rod articulation and which are slightly inflected at the mid- line. However, distinction between this species and P. subulata is often difficult, especially in small specimens that preserve only inflected goth- ic arch style rod articulation. Distinction between the two species can be made on these criteria: 1 , P. blairi possesses distinct nodes on the rods, P. subulata usually does not; 2, P. blairi has a slight adapertural inflection of the rods near the midline except in the apical region, P. subulata does not; and 3, P. blairi possesses 6-13 rods/cm whereas P. subulata has 20-35 rods/cm.
Two species are here considered synonymous with P. blairi: P. se- daliensis (Miller and Gurley) and P. indiana Sinclair, MS. The type specimens of both are well enough preserved to compare all qualitative and quantitative features of taxonomic interest. Values obtained by measuring these specimens are given in Appendix B. In all respects, P. sedaliensis and P. indiana are indistinguishable from the lectotype and paralectotypes of P. blairi.
Material examined. — 34 specimens; housed in the AMNH, FMNH, UCGM, UMC, and the USNM.
PARACONULARIA BYBLIS (White, 1862)
Figs. 3.7, 19.1-19.6, 23.2, 31.4
Conularia byblis White, 1862, p. 22; Miller, 1877, p. 141; Bigsby, 1878, p. 78; Herrick, 1888^2, p. 95; Miller, 1889, p. 390; Weller, 1898, p. 189; Weller, 1900^, p. 118- 1 19, PL 7, fig. 7; Weller, 1900Z?, p. 73; Grabau and Shimer, 1910, p. 14.
Conularia byblis White. Winchell, 1870, p. 257.
Conularia biblis (sic) White. Bigsby, 1878, p. 316.
Paraconularia byblis (V^hiXo). Sinclair, 1948, p. 200-201; Babcock and Feldmann, 1986, figs. IE, 2B.
Adesmoconularia byblis (White). Driscoll, 1963, p. 40-41, PI. 3, figs. 1-7; Tasch, 1973, fig. 5.14 Ga-b; Tasch, 1980, fig. 5.14 Ga-b.
Conularia! sp. Driscoll, 1963, p. 41, PL 3, fig. 8.
Description. —Exoskeleton up to 7 cm in length. Major apical angle 18-26®; minor apical angle 10-19®. Rod articulation inflected gothic arch style in apical region and inflected circular curve style elsewhere; rods are strongly inflected adaperturally at mid- line; rod angle 12-18°. Rods generally abut at midline; 12-29 rods/cm. 1-2 nodes/mm; spines seem to be absent. Apical wall may be present.
Occurrences. — Lower Mississippian of Indiana, Iowa, Kentucky and Ohio; localities 29, 36, 39, 43, 50, 60, 62, 66, 71-72, 76, 77, 78, 81,
daliensis Miller and Gurley; locality 98. 18.3; UCGM 3985, counterpart of specimen in Fig. 18.2. 18.4; UCGM 3984, paralectotype, ?minor face; locality 98. 18.5; UCGM 3986, lectotype, a flattened specimen preserved in micrite; locality 98. Bar scales represent 1 cm.
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185, 195, 210, and 223-224. A specimen referred with question to this species, UMMP 26735, is from Tennessee; locality 250.
—Holotype, UMMP 2167.
Remarks. —Paraconularia byblis (White) is distinguished from all other species of Paraconularia by the combination of closely spaced rods, 12-29/cm, the lack of spines and by rod articulation involving an inflected gothic arch style in the apical region and an inflected circular curve style elsewhere. There does not seem to be a species present in the Devonian or Mississippian rocks of North America with which this taxon could be easily confused if well preserved specimens were available for study.
Driscoll (1963), designated C. byblis White as the type species of a new genus, Adesmoconularia. Adesmoconularia, by Driscoll’s defini- tion, is distinguished from Calloconularia Sinclair by a larger size and a lack of “swelling” of the interridge areas near the comer angles in Adesmoconularia. Adesmoconularia was deemed by Driscoll unlike Paraconularia Sinclair in the lack of nodes and by the presence of an apical wall in Adesmoconularia. To date, no other species have been referred to the genus Adesmoconularia.
Examination of the holotype of Calloconularia strimplei Sinclair (FMNH PE 1 42), the type species of the genus Calloconularia, reveals that there is no expansion of the interridge areas near the comer angles. Examination of the holotype of Conularia byblis White, type species of the genus Adesmoconularia Driscoll, shows that nodes are present on the rods, but they are very small and inconspicuous. The holotype appears to have been considerably weathered, rendering the nodes inconspicuous in most places on the specimen. Apical walls are present, but rare, in specimens of Paraconularia. According to Driscoll’s di- agnosis, then, size is the only criterion which distinguishes Adesmo- conularia from Calloconularia', there is no distinction between Ades- moconularia and Paraconularia. Therefore, Adesmoconularia Driscoll, 1963 is here considered a junior synonym of Paraconularia Sinclair, 1940.
Fig. \9.— Paraconularia byblis (White). 19.1; UMMP 2167, holotype, a weathered spec- imen preserved in micritic limestone, comer view; locality 62, 19.2; CMNH 4492, small specimen, preserved in shale and compressed along the faces and at the aperture; locality 219. 19.3; CMNH 2295, external mold preserving apical region; locality 195. 19.4; UMMP 2167, same specimen as in Fig. 19.1, minor face. 19.5; UMMP 2167, same specimen as in Fig. 19.1, detail of minor face. 19.6; CMNH 4691, ?major face of a specimen preserved in shale; locality 185. Bar scales represent 1 cm.
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Fig. 20.— Paraconularia chagrinensis Babcock and Feldmann, n. sp. 20,1; CMNH 6717, paratype, preserved in a dark gray phosphatic concretion. Note apparent healed wound;
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Material examined. — S'i specimens; housed in the CM, CMNH, GSC, FMNH, UMMP, and the USNM.
PARACONULARIA CHAGRINENSIS Babcock and Feldmann, new species Figs. 20.1-20.6, 21.1, 21.2
Description. ~~Exosk.Q\Qton up to 9 cm in length. Major apical angle about 28®; minor apical angle 20-21°. Rod articulation exclusively of inflected gothic arch style. Rods usually alternate at midline; rods, if they alternate, usually alternate left superior on both major and minor faces; rod angle 9-12®. 16-20 rods/cm. 3-4 nodes/mm; nodes appear to be subtle in apical region and prominent in apertural region; 3-4 adapertural spines/ mm; 3-4 adapical spines/mm. No apical wall observed.
Occurrences.— Devonian of Ohio; localities 178-184.
rj;/7^.y.-Holotype, CMNH 6633; 12 paratypes, CMNH 1247, 1272, 1427, 1622, 1674, 1788, 1818, 4030, 4292, 6717, 6807-6808.
Remarks.— Among species of Paraconularia, only P. chagrinensis Babcock and Feldmann, n. sp. possesses the combination of wide apical angles, 20-28°, adapertural and adapical spines, as well as rod articu- lation which is exclusively of inflected gothic arch style. More striking, however, is the pattern of nodes on the rods. Paraconularia chagri- nensis is the only conulariid observed which appears to have nodes which increase in size aperturally. Nodes are inconspicuous in the apical region, but are prominent in the apertural region. The increase in size of the nodes is not well shown in the holotype owing to the poor preservation of the apertural region of this individual. The pattern is well documented, however, in CMNH 6717 (Fig. 20.6).
When fragments of exoskeleton from the apertural region are found alone, as is the case with the specimen illustrated in Figs. 20.3-20.4, they are easily mistaken for species of Conularia such as C subcar- bonaria or C. multicostata. This dilemma can be resolved only when more complete material is found. Most conulariids from the small, presumably phosphatic, nodules found in the Upper Devonian Chagrin Shale of northeastern Ohio are preserved as fragmentary specimens, which renders generic identification difficult. To date, only P. chagri- nensis has been identified from this unit.
One paratype, CMNH 6717, is noteworthy not only for demonstrat- ing the unique pattern of the nodes, but also for exhibiting an apparent
locality 179. 20.2; CMNH 1622, paratype; locality 184. 20.3; CMNH 1818, paratype; locality 181. 20.4; CMNH 1818, same specimen as in Fig. 20.3, detail showing nodes and spines. 20.5; CMNH 6633, holotype; locality 1 80. 20.6; CMNH 67 1 7, same specimen as in Fig. 20.1, ?minor face. Note increase in size of nodes adaperturally. Bar scales represent 1 cm.
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healed injury on one face (Fig. 20,1). On this specimen, rods in the apertural region have been truncated and their broken ends rounded slightly. Several rods have filled much of the region from where exo- skeleton has been removed. These rods are oriented at a high angle to the rods which comprise the remainder of the exoskeleton and are complete with a midline distinct from the original midline on this face. A small gap is left between the most adapical portion of the injury and the most adapical rods which have filled the void. This region is filled with integument that lacks embedded rods.
Material examined. --13 specimens; housed in the CMNH.
Etymology of trivial name. — Named for the Chagrin Shale, currently the only known occurrence of this taxon.
PARACONULARIA CHESTEREN SI S {V^orthQn, 1883)
Figs 17.7-17.8, 22.4-22.7, 23.1-23.4,
23.7, 24.1, 24.3, 32.5
Conularia chesterensis Woithen, 1883, p. 325; Miller, 1889, p. 390, Worthen, 1890, p.
134, PI. 1 1, fig. 9a-b; Miller, 1897, p. 765; Weller, 1898, p. 189; Kent, 1982, p. 27. Pamconularia chesterensis Worthen. Sinclair, 1948, p. 201-202; Babcock and Feldmann, 1986, fig. 4C.
Paraconularia newberryi (Winchell). Sensu Driscoll, 1963, PI. 2, figs. 6-9. Paraconularia crawfordsvillensis (Owen). Sensu Lane, 1973, p. 93-95, PI. 8, fig. 1, PI. 9, figs. 1-2.
Description. — Exoskeleton up to 20 cm in length. Major apical angle 14-21®; minor apical angle 1 0-18°. Rod articulation exclusively of inflected circular curve style, recurved near the midline. Rods alternate or abut at midline; rods, if they alternate, are not preferentially right superior or left superior on either the major or minor face; rod angle 8-12°. 8-20 rods/cm. 4-5 nodes/mm; adapertural spines appear to be absent in apical region, but small spines are sometimes present, 4-5/mm, in apertural region; adapical spines absent. Apical wall not observed.
Occurrences. — Upper Mississippian of Alabama, Kentucky, Illinois, Indiana, Iowa, Missouri, Nevada, Tennessee; localities 1, 10-12, 16- 21, 33, 35-36, 38-41, 47, 49-59, 70, 84-89, 104, 1 10, and 251. Spec- imens referred questionably to this species have also been found in British Columbia and Utah; localities 7 and 352.
Types. -nololypc, ISGS 2489.
Remarks. —Paraconularia chesterensis (Worthen) is similar to P. al~ penensis Babcock and Feldmann, n. sp., P. missouriensis (Swallow) and P. recurvatus Babcock and Feldmann, n. sp. in having rods that are recurved near the midline. Of these, P. alpenensis does not exhibit rods that are recurved in the apical region, and both P. missouriensis and P. recurvatus exhibit rods that are strongly recurved. The rods of P. chesterensis tend to be slightly recurved. A rod pair in this taxon often approximates the outline of a truncated pyramid (Figs. 22.4, 22.7).
In some cases, values for apical angles, rods/cm and rod angles may
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be similar for specimens of P. chesterensis and P. missouriensis. If rod articulation is also similar, distinction between the two may be made on the basis of the spacing between nodes. Paraconularia chesterensis possesses 4-5 nodes/mm and P. missouriensis possesses only 2-3 nodes/ mm.
Specimens referable to P. chesterensis which have been observed with well-preserved spines are few in number. Moreover, it seems that spines are only produced in the apertural regions of those individuals that have them. When present, the spines are usually inconspicuous and seem to be directed only in the apertural direction (for example, Fig. 17.8). Many specimens referable to P. chesterensis (for example, Fig. 23.2), however, seem to have small ridges developed in the integ- ument between ridges. These ridges resemble interridge crests. Such structures may indicate that adapertural, and perhaps even adapical, spines are produced in areas other than the apertural region in this taxon. No specimens exhibiting this have been observed to date. Thus, the observations that only adapertural spines are present in P. ches- terensis, and when present, that they occur only in the apertural region, may be erroneous and owing to a lack of evidence to the contrary.
Material examined. — ?> 5 5 specimens; housed in the FMNH, GSC, ISGS, lUPC, and the USNM.
PARACONULARIA MISSOURIENSIS 1860)
Figs. 21.3, 25.1, 25.2-25.5, 26.1-26.2, 32.1
Conularia missouriensis Swallow, 1860, p. 657; Miller, 1877, p. 141; Bigsby, 1878, p. 316; Miller, 1889, p. 390; Keyes, 1894, PI. 35, fig. la-b; Miller, 1897, p. 765; Weller, 1898, p. 190; Grabau and Shimer, 1910, p. 14; Branson, 1944, p. 246.
Conularia missouriensis Swallow?. Meek and Worthen, 1873, p. 541-542, PL 22, fig. 5;
White, 1880, p. 513, PL 6, fig. 4; Walcott, 1884, p. 264, PL 23, fig. 4. Paraconularia (Swallow). Sinclair, 1948, p. 198-199.
Conularia gratiosa Miller and Gurley, 1893, p. 74, PL 8, fig. 1; Miller, 1897, p. 765; Weller, 1898, p. 190.
Conularia greenei Miller and Gurley, 1896, p. 27-28, PL 3, fig. 3; Miller, 1897, p. 765;
Weller, 1898, p. 190, Cumings, 1906, p. 1367, PL 24, fig. 14.
Paraconularia greenei (Miller and Gurley). Sinclair, 1948, p. 194.
Paraconularia gratiosa (Miller and Gurley). Sinclair, 1948, p. 198.
Paraconularia sciotovillensis OtiscoW, 1963, p. 37-40, PL 1, figs. 9-12; Tasch, 1973, fig.
5.16, Table 5.2; Tasch, 1980, fig. 5.16, Table 5.2.
Ctenoconularial greenei (Miller and Gurley). Moore and Harrington, 1956, p. F65, fig. 51.4.
Conularia sp. Leary, 1985, PL 3, fig. 4.
Paraconularia cf. P. missouriensis (Swallow). Babcock, 1985a, p. 66-70, fig. lA-B.
Description.— up to 22 cm in length. Major apical angle 14-22°; minor apical angle 10-18°. Rod articulation inflected circular curve style, strongly recurved near the midline. Rods usually alternate at midline; if rods alternate, pattern is usually left superior; rod angle 6-17°. 4-10 rods/cm. 2-3 nodes/mm; 2-3 adapertural spines/ mm; adapical spines absent. Apical wall not observed.
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Fig. 2 1 . — 2 1 . 1 ; Paraconularia chagrinensis Babcock and Feldmann, n. sp., CMNH 1788, two small paratype specimens, presumably attached to the same object (obscured) and
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Occurrences.— l^owQt Mississippian of Alberta, Illinois, Indiana, Iowa, Kentucky, Missouri, and Ohio; localities 2, 5, 14-15, 31, 39, 69, 83, 94, 100, 103, 223 and 226-227. A specimen referred with question to this species has been found in Alberta; locality 5.
Plaster cast of presumed holotype, FMNH UC 6639. Ho- lotype of C. gratiosa, FMNH UC 6627, plastoholotype, USNM 67893; holotype of C greenei, FMNH UC 6628, plastoholotype USNM 67880; holotype of P. sciotovillensis, UMMP 26740.
—Considerable confusion has existed over the definition of P. missouriensis (Swallow). This confusion of nomenclature is related to at least two problems: 1, an ambiguous original definition of the species, a definition which may have incorporated characters now iden- tified as belonging to at least two species; and 2, a loss of Swallow’s original specimens. Swallow’s type material was found in the “Car- boniferous Limestone” of Cooper County, Missouri. This locality has yielded at least two conulariid species; herein, they are identified as P. missouriensis and P. blairi. It is possible, from Swallow’s description (Swallow, 1860, p. 657), that specimens belonging to both forms were used in the formulation of the original definition of P. missouriensis.
Some early authors, most notably Meek and Worthen (1873) and Keyes (1894), used Swallow’s indication that the faces of P. missou- riensis were “marked by flexuous, high, sharp plications” as the primary determinative characteristic of the species. This concept of the species is followed herein. A plaster cast of a specimen, marked “holotype?” of P. missouriensis (FMNH UC 6639) is presumed to represent a cast of the holotype of this species.
Paraconularia missouriensis is similar to P. chesterensis (Worthen) in having an inflected circular curve style of rod articulation, with the rods being reflexed near the midline. The degree of reflexure, however, is greater in P. missouriensis. Paraconularia missouriensis can also be distinguished from P. chesterensis by having a greater number of nodes/ mm on the rods, 4-5 nodes/mm as compared to 2-3 nodes/mm.
Other species of Paraconularia which have recurved rods include P. alpenensis Babcock and Feldmann, new species and P. recurvatus Bab-
preserved in a phosphatic concretion. Arrow indicates a stalk; locality 1 84. 2 1 .2; CMNH 4294, partially disarticulated paratype; locality 183. 21.3; P. missouriensis (Swallow), FMNHUC 1 125, view of major face; locality 14. 21.4; P. subulata(¥i?A\), USNM 395829, preserved in a very dark gray, organic-rich shale. Note that no integument is present and that rods are disarticulated. This specimen indicates that rods and integument are separate components of the conulariid exoskeleton. Two specimens of ^^Linguld’" {=Bar- roisellal) are visible in this photograph; locality 189. Bar scales in Figs. 21.2-21.4 rep- resent 1 cm; bar scale in Fig. 21.1 represents 5 mm.
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cock and Feldmann, new species. Neither of these taxa have rod spacing values less than 14 rods/mm or adapical spines on the rods and are therefore easily distinguished from P. missouriensis. Moreover, P. ah pensis possesses non-recurved rods in the apical region, unlike P. mis- souriensis.
The species Conularia gratiosa Miller and Gurley, C. greenei Miller and Gurley and P. sciotovillensis Driscoll are here included as junior synonyms of C missouriensis because they all exhibit: 1 , similar values for apical angles; 2, similar values for rod angles; 3, same rod articu- lation style, including right superior rods if the rods alternate at the midline; 4, similar values for rod spacing; 5, similar values for nodes/ mm; and 6, presence of adapical spines. Comparative values are given in Appendix B.
Material examined. — 30 specimens; housed in the AMNH, FMNH, GSC, ISGS, and the USNM.
PARACONULARIA OKLAHOMAENSIS
Babcock and Feldmann, new species Fig. 27.5
Description. — Ty^scn^iion based only upon holotype. Exoskeleton 5.4 cm in length. Major apical angle 19°; minor apical angle 17°. Rod articulation of gothic arch style in apical end and of inflected circular curve style elsewhere. Rods usually abut at midline; if they alternate, pattern is usually right superior on major face and left superior on minor face; rod angle 12-13°. 24 rods/cm in apical region, 12 rods/cm elsewhere. Nodes and spines absent. Apical wall not observed.
Occurrence.— UppQT Mississippian of Oklahoma; locality 229.
--Holotype, USNM 409811.
Remarks.— Par aconularia oklahomaensis Babcock and Feldmann, n. sp. differs from all other described species of the genus in the com- bination of rod articulation pattern, frequent rod abuttment, spacing of rods, with 24 rods/cm in the apical region and 1 2 rods/cm elsewhere and the lack of nodes and spines on the rods.
Fig. 22. — 22.1--22.3; Paraconularia blairi (Miller and Gurley). 22.1; AMNH 25056, specimen intended by Sinclair (1948) to be holotype of P. indiana Sinclair, major face; locality 32. 22.2; AMNH 25056, same specimen as in Fig. 22,1, comer view. 22.3; AMNH 25056, same specimen as in Fig. 22.3, minor face. Note overturned apertural termination. 22.4--22.7; P. chesterensis (Worthen). 22.4; lUPC 17414, bryozoan=en- cmsted specimen; locality 1. 22.5; lUPC 17415, comer region of collapsed specimen that has been encmsted by bryozoans subsequent to collapse; locality unknown, 22.6; FMNH UC 23023; locality 38. 22.7; lUPC 11316; locality 57. Bar scales represent 1 cm.
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Fig. 23.-23.1-23.4; Paraconularia chesterensis (Worthen). 23.1; ISGS 2489, holotype, preserved in micritic to sparry limestone, minor face; locality 10. 23.2; ISGS 2489, same specimen as in Fig. 23.1, comer view. 23.3; ISGS 2489, same specimen as in Fig. 23.1,
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This taxon is easily confused with P. subulata (Hall), which also lacks nodes and spines on the rods and has inflected gothic arch rod artic- ulation in the apical region and inflected circular curve rod articulation elsewhere. The rods of P. oklahomaensis, however, are greatly inflected in the vicinity of the comer angles in the apical region and almost imperceptibly inflected elsewhere. In specimens of P. subulata, the pattern of relative inflexure of the rods is reversed. Paraconularia okla- homaensis is further distinguished from P. subulata in having 40-50% of the rods abutting at the midline whereas specimens referred to P. subulata seldom have more than 10% of the rods abutting.
Material examined. — 1 specimens, USNM 40981 1.
Etymology of trivial name. — Named for the State of Oklahoma.
PARACONULARIA PLANICOSTATA (Dawson, 1868)
Figs. 3.1, 27.1-27.4, 21 .6-21
Conularia planicostata Dawson, 1868, p. 307-308, fig. 117; Dawson, 1878, p. 307-308, fig. 117; Bigsby, 1878, p. 316; Dawson, 1883, p. 416; Lesley, 1889, p. 145, fig.; Beede, 1911, p. 174, 186; Bell, 1929, p. 98-100, PI. 32, figs. 1-2; Bamber and Copeland, 1976, PI. 15, fig. 3.
Conularia planocostata (sic) Dawson. Miller, 1877, p. 141; Miller, 1889, p. 390; Weller, 1898, p. 191.
Conularia quadrisulcata Miller in Sowerby. Sensu Dawson, 1889, p. 87, fig.
Conularia sorrocula Beede. Sensu Bell, 1929, p. 100, PL 32, figs. 3-3a.
Conularia cf. tenuis Slater. Sensu Bell, 1929, p. 100, PL 32, figs. 4-5.
Paraconularia planicostata (Dawson). Sinclair, 1948, p. 199-200; Babcock and Feld- mann, 1984, p. 16-1 7; Babcock and Feldmann, 1986, fig. 2G.
Connularia (sic) planicostata Dawson. Alison and Carroll, 1972, p. 17.
Description.— FxosV.Q\QXon up to 8 cm in length. Major apical angle 21-25°; minor apical angle 18-22°. Rod articulation inflected gothic arch style in apical region and gothic arch style elsewhere. Rods abut or alternate at midline; if they alternate, rod pattern is usually right superior; rod angle 1 1-16°. 12-20 rods/cm. Nodes and spines absent. Apical wall not observed.
Occurrences. —Lowqx to Upper Mississippian of Nova Scotia and Quebec; localities 163-173 and 248.
— Holotype, RM(MU) 2749, plastoholotype, GSC unnum- bered.
Remarks.— Paraconularia planicostata (Dawson) is distinguished
major face. 23.4; FMNH UC 25175; small specimen preserving apical region; locality 12. 23.5-23.6; P. missouriensis (Swallow). 23.5; FMNH UC 1125; specimen preserved in micrite showing darkened areas in the integument along the midline and surrounding the ridges. Darkened areas of integument in the vicinity of the midline have been in- terpreted by numerous authors as remains of original color markings. Specimen not coated with ammonium chloride; locality 14. 23.6; USNM 14425, original of Walcott (1884, PL 23, fig. 4), locality 110. 23.7; P. chesterensis (Worthen), ISGS 2489, same specimen as in Fig. 23.1, detail of minor face. Bar scales represent 1 cm.
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Fig. 24. — 24.1; Pamconularia chesterensis (Worthen), USNM 50150, portion of large cluster of individuals preserved in siltstone and showing incomplete remains of stalks
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from other members of the genus by the combination of: 1 , its small size, generally less than 7.5 cm in length; 2, its inflected gothic arch and gothic arch rod articulation styles; 3, its very narrow apical angles, 8-14®; 4, its widely spaced rods, 12-20 rods/cm; and 5, its lack of nodes and spines. Paraconularia planicostata is particularly notable, and eas- ily distinguished from all other taxa described herein because it pos- sesses gothic arch rod articulation up to 7.5 cm from the hypothetical apex in the adapertural direction.
Dawson (1868, p. 308), in describing the species Conularia plani- costata, compared the taxon to an apparent manuscript species, C novascotica Hartt. Dawson considered this taxon, also from the Mis- sissippian of Nova Scotia, to be a variety of C planicostata. The in- tended holotype specimen of C novascotica is lost, but based upon Hartt’s scant description {in Dawson, 1868), it is likely to be an example of P. planicostata (Dawson).
Material examined. — 30 specimens; housed in the CM, GSC, NYSM, RM(MU), and the USNM.
PARA CONULARIA RECUR VA TUS
Babcock and Feldmann, new species Figs. 32.3, 32.6
Description. — up to 8 cm in length. Major apical angle about 16°; minor apical angle about 1 5°. Rod articulation exclusively of inflected circular curve style, greatly recurved near midline in apical region and slightly recurved near midline elsewhere. Rods abut or alternate at midline; if they alternate, pattern is usually left superior; rod angle 8-12°, 18-28 rods/cm. 2-3 nodes/mm; spines absent. Apical wall not observed.
Occurrence.— UpptY Devonian of Nevada; locality 109.
—Holotype, part and counterpart, USNM 409806. Three paratypes, USNM 409807-409809, all present on the same slab as the holotype. The paratype labelled as USNM 409808 is preserved as part and counterpart.
Remarks. —Paraconularia recurvatus Babcock and Feldmann, n. sp. is unique among members of this genus in having rods which are both closely spaced and which are recurved near the midline. Three other species of Paraconularia examined in this study have recurved rods, namely, P. alpenensis Babcock and Feldmann, n. sp., P. chesterensis
(arrows), attached to possible plant matter. Specimen not coated with ammonium chlo- ride; locality 27. 24.2; Paraconularia byblis (White), USNM 409800, specimen preserved in siderite concretion and showing a stalk (arrow); locality 7 1 . Specimen not coated with ammonium chloride. 24.3; P. chesterensis (Worthen), USNM 50150, same specimen as in Fig. 24.1, view showing the complete aggregation of conulariids as exposed at the surface of the slab. Bar scales represent 1 cm.
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(Worthen) and P. missouriensis (Swallow). Paraconularia recurvatus differs from all of these forms in rod spacing. Specimens in the type lot of P. recurvatus have 18“28 rods/cm while the holotype of P. al- penensis has 14 rods/cm, specimens of P. chesterensis have 8™25 rods/ cm and specimens of P. missouriensis have 6-10 rods/cm.
In its overall appearance, it seems as though P. recurvatus could be mistakenly included in the genus Conularia. However, the lack of spines indicates that this species should be included in the genus Para- conularia. All quantitative data (Appendix B) further support this con- clusion.
Material examined.— A specimens; housed in the USNM.
PARACONULARIA (Whiteaves, 1891)
Figs. 28.3, 28.5»28.6
Conularia salinensis Whiteaves, 1891, p. 244, PL 32, figs. 9-9a.
Conularia s.l. salinensis Whiteaves. Sinclair, 1948, p. 287.
Description. — I>Qscnption based only upon holotype. Exoskeleton 3 cm in length. Major apical angle approximately 24®; minor apical angle 21®. Rod articulation inflected gothic arch style in apical region and inflected circular curve style elsewhere. Rods usually alternate at midline; if they alternate, pattern is usually right superior on major and minor faces; rod angle 1 3® in apical region, 8® elsewhere. 24 rods/cm. 3-4 nodes/mm; 3-4 prominent adapertural spines/mm; adapical spines absent. Apical wall not observed.
Occurrence.— Mhsisuppmn of Alberta; locality 3.
-Holotype, GSC 4292.
Remarks.— This taxon can be distinguished from other species of Paraconularia by the combination of: 1 , inflected gothic arch and in- flected circular curve styles of rod articulation; 2, apical angles of 8- 13°; 3, rod spacing of 24 rods/cm; 4, node spacing of 3-4 nodes/mm; and 5, prominent adapertural spines. Paraconularia salinensis (Whit- eaves) does not seem to be easily confused with any other conulariid species described to date from the Devonian or Mississippian rocks of North America.
Material examined. — 1 specimen, GSC 4292.
Fig. 25.— Paraconularia missouriensis (Swallow). 25.1; FMNH UC 6639, plaster cast of presumed holotype specimen; locality 100. 25.2; ISGS 2619; oblique view of specimen with three intumed apertural terminations. The fourth apertural termination is broken off, but there is no indication of infolding. 25.3; UMMP 26740, holotype of P. scioto- villensis Driscoll, same specimen as in Fig. 26.1, detail of major face; locality 226. 25.4; FMNH UC 6628, holotype of Conularia greenei Miller and Gurley; minor face of a specimen preserved in micrite. 25.5; FMNH UC 6628, same specimen as in Fig. 25.4, comer view. Bar scales represent 1 cm.
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Fig. 26.— Pamconularia missouriensis (Swallow). 26.1; UMMP 26740, holotype of P. sciotovillensis Driscoll, minor face of a somewhat distorted individual preserved in a
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PARACONULARIA SORROCULA (Beede, 1911)
Figs. 28.1-28.2
Conularia sorrocula Beede, 191 1, p. 184, 186, 2 figs.
Paraconularia sorrocula Beede. Sinclair, 1948, p. 199.
Description.— Exoskeleton up to 3 cm in length. Major apical angle 19-24°; minor apical angle 1 7-22°. Rod articulation exclusively inflected gothic arch style with slight adapertural inflection at midline. Rods usually alternate at midline, less frequently they abut; rods, if they alternate, pattern is usually left superior on both major and minor faces; rod angle 11-14°. 18-20 rods/cm. Nodes elongate and appear to be continuous structures with adapertural spines; 5-6 nodes/mm; 5-6 adapertural spines/cm. Adapical spines absent. Apical wall not observed.
Occurrence. —Mississippisin of Quebec; locality 247.
Typp. — Holotype, part and counterpart, NYSM 9414.
Remarks. —Paraconularia sorrocula (Beede) is unique among North American Devonian or Mississippian examples of Paraconularia in the possession of only inflected gothic arch rod articulation. It is also the only conulariid species reported herein which has the nodes merged with the adapertural spines without a significant change at the junction of the two structures (Fig. 28.2). The nodes are not round or oblate in outline as in other species of Paraconularia, but are elongate.
The holotype of P. sorrocula is curved in the apertural region. This feature may have been present on the specimen in life, although this cannot be confirmed owing to the crushed nature of the fossil. Material examined. — specimens; housed in the NYSM.
PARACONULARIA SUBULATA (Hall, 1858)
Figs. 3.2, 3.5-3.6, 21.4, 29.1-29.10,
30.1-30.8, 31.1-31.5, 33.4
Conularia subulata Hall, 1858, p. 32; Miller, 1877, p, 141; Bigsby, 1878, p. 316; Whit- field, 1882, p. 91, PI. 8, fig. 3; Hall, 1883, p. 372-373, PI. 31, fig. 3; Miller, 1889, p. 390; Lesley, 1889, p. 146, fig.; Lesley, 1895, p. 1690, fig.; Weller, 1898, p. 192; Whitfield and Hovey, 1901, p. 406-407; Cumings, 1906, p. 1366, PL 25, fig. 3. Conularia victa White, 1862, p. 22-23; Miller, 1877, p. 141; Bigsby, 1878, p. 316; Miller, 1889, p. 390; Herrick, 1893, PI. 19, fig. 3; Weller, 1898, p. 192.
Conularia newberryi Winchell, 1865, p. 130; Winchell, 1870, p. 258; Meek, 1875, p. 316-317, PL 18, fig. 2a-b; Miller, 1877, p. 141; Bigsby, 1878, p. 316; Hall, 1879, PL 34A, fig. 12; Herrick, 1888^, p. 93-94, PL 6, figs. 13, 17, PL 8, fig. 9; Herrick,
siderite concretion; locality 226. 26.2; UMMP 26740, same specimen as in Fig. 26.1, major face. 26.3; AMNH 28692, minor face of a specimen preserved in sparry limestone; locality 30. 26.4; FMNH UC 6627, holotype of Conularia gratiosa Miller and Gurley, preserved in micritic limestone, comer view; locality 30. 25.5; FMNH UC 6627, same specimen as in Fig. 26.4, detailed view of a minor face. Bar scales represent 1 cm.
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1888Z?, PI. 8, fig. 5, PI. 10, figs. 27-28; Lesley, 1889, p. xv; Miller, 1889, p. 390; Herrick, 1893, PL 19, fig. 5; Miller, 1897, p. 765; Weller, 1898, p. 191; Clarke and Ruedemann, 1903, p. 566; Grabau and Shimer, 1910, p. 13, figs. 1227a-b; Tasch, 1973, fig. 5.16, Table 5.2; Tasch, 1980, fig. 5.16, Table 5.2.
Conularia whitei Meek and Worthen, 1865, p. 253-254; Bigsby, 1878, p. 316. Conularia newberryi WinchelYI Herrick, 1887, p. 146-147, PI. 14, fig, 14.
Conularia victa White? Herrick, 1888Z), p. 47-48, PL 8, fig. 3.
Conularia whitii (sic) Miller, 1889, p. 390; Weller, 1898, p. 192.
Conularia sampsoni Miller, 1892<2, p. 690-691, PL 14, figs. 11-12; Miller, 1892^, p. 692; Holm, 1893, p. 125; Weller, 1898, p. 191; Branson, 1938, p. 110-111, PL 14, fig. 9; Branson, 1944, p. 216.
Paraconularia subulata (Hall). Sinclair, 1948, p. 198; Babcock and Feldmann, 1986, figs. IC-D, IG, 2F, 3A-C.
Paraconularia victa (White). Sinclair, 1948, p. 200.
Paraconularia newberryi (Winchell). Sinclair, 1948, p. 191-192; Driscoll, 1963, p. 34- 37, PL 1, figs. 1-5, PL 2, figs. 1-4.
Paraconularia whitei (Meek and Worthen). Sinclair, 1948, p. 192.
Paraconularia sampsoni Miller. Sinclair, 1948, p. 197.
Paraconularia sp. Feldmann, Coogan and Heimlich, 1977, fig. 2.50A.
""Conularia"" sp. Thompson, 1982, fig. 357.
Paraconularia cf. P. missouriensis {SwbWow). Sensu Babcock, 1985^z, figs. la-b. Paraconularia missouriensis (Swallow). Sensu Babcock, 1985^2, fig, 2.
Paraconularia cf. P. subulata. Babcock and Feldmann, 1986, fig. 2H.
Description. --ExoskQlQXovi up to 17 cm in length. Major apical angle 17-22®; minor apical angle 12-18°. Rod articulation inflected gothic arch style in apertural region and inflected circular curve style elsewhere. Rods usually alternate at midline; if they alternate, pattern is usually right superior on major face and usually left superior on minor face; rod angle 15-18®. 20-35 rods/cm. Nodes absent or present; if present, they are incon- spicuous, 2-3/mm; spines absent. Apical wall present.
Occurrences. --1.0SNQV Mississippian of Illinois, Indiana, Kentucky, Montana, Ohio; localities 16, 26, 29, 38, 39, 72, 75, 82, 106-107, 185- 192, 194, 196-203, 205-208, 212-217, 220-222.
— Lectotype, designated herein from James Hall’s suite of three syn types of C. subulata, AMNH 32403, smaller of two specimens bearing this number; two paratypes, AMNH 32403, larger of two spec- imens bearing this number, and AMNH 32404. Holotype of C victa,
Fig. 27.-27.1-27.4; Paraconularia planicostata (Dawson). 27,1; RM(MU) 2749, ho- lotype; major face, locality 164. 27.2; RM(MU) 2749, same specimen as in Fig. 27.1, comer view. Note exoskeletal constrictions. 27.3; RM(MU) 2749, same specimen as in Fig. 27.1, minor face. 27.4; RM(MU) 2749, same specimen as in Fig. 27.1, detail of major face. 27.5; P. oklahomaensis Babcock and Feldmann, n. sp., USNM 409801, holotype, a flattened individual. 27.6-27.8; P. planicostata (Dawson). 27.6; CM 22667, major face; locality 168. 27.7; CM 22667, same specimen as in Fig. 27.6, minor face. 27.8; CM 22667, same specimen as in Fig. 27.6, comer view. Bar scales in Figs. 27.1- 27.5 represent 1 cm; bar scales in Figs. 27-6-27.8 represent 5 mm.
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Fig. 28. —28. 1-28.2; Paraconularia sorrocula (Beede). 28.1; NYSM 9414, slab showing two specimens, holotype to the right. Note orbiculoid brachiopods attached to, and
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UMMP 2178, plastoholotype, GSC unnumbered; holotype of C new- berryi, UMMP 245; holotype of C whitei, UIPC 10866; holotype of C sampsoni, FMNH UC 6961, plastoholotype, USNM 68156.
Remarks.— Par aconularia subulata (Hall) is most similar in mor- phology to P. oklahomaensis Babcock and Feldmann, n. sp. Both taxa have inflected gothic arch rod articulation in the apical region and inflected circular curve rod articulation elsewhere. Also, specimens referable to both taxa may lack nodes and spines. Except for rod spacing values, the quantitative measures are also very similar. Paraconularia subulata has a rod spacing of about 20-35 rods/cm whereas the ho- lotype of P. oklahomaensis has a value of 1 2-24 rods/cm. Paraconu- laria subulata differs most substantially from P. oklahomaensis in hav- ing very little infleetion of the rods in the vicinity of the apex and in having a strong inflection of the rods elsewhere.
In addition to P. oklahomaensis, P. blairi (Miller and Gurley) bears close similarity to P. subulata. Both taxa have inflected gothie arch rod articulation adapically and inflected circular curve rod articulation ad- aperturally. Apical angles and rod angles are nearly equal in the two forms. Paraconularia subulata differs from P. blairi in its lack of nodes on the rods or in having inconspicuous nodes, in the lack of any ad- apertural inflection of the rods near the midline and in the possession of a greater number of rods/cm, 20-35 as compared to 6-8.
The species P. victa (White), P. newberryi (Winchell), P. whitei and P. sampsoni (Miller) are all included as junior synonyms of P. subulata because they are indistinguishable from the lectotype and paralecto- types of P. subulata. The type specimens of all these species bear subtle nodes on the rods; all have apical angles in the range of 12-22®; all have inflected gothic arch styles of rod articulation; all have 20-35 rods/cm; and all have rod angles of 1 5-18®. The holotype of P. sampsoni possibly could be construed as a juvenile of P. blairi, but the lack of rods which are slightly inflected at the midline makes assignment of this specimen to P. subulata more reasonable.
Paraconularia subulata is one of the most abundant conulariids in
located near, the conulariids; locality 247. 28.2; NYSM 9414, same specimen as in Fig. 28.1, detail of holotype. 28.3; P. salinensis (Whiteaves), GSC 4292, holotype, minor face; locality 3. 28.4; P. sp., CM 34531, a collapsed and poorly preserved specimen in tan and dark red colored dolostone; locality 249. 28.5-2^6; P. salinensis (Whiteaves). 28.5; GSC 4292, same specimen as in Fig. 28.3, comer view. 28.6; GSC 4292, same specimen as in Fig. 28.3, detail of minor face. 28.7; P. yochelsoni Babcock and Feldmann, n. sp., external molds of two specimens attached to plant matter, holotype, UMMP 45499, to the right, paratype, UMMP 65509, to the left; locality 93. Specimen is not coated with ammonium chloride. Bar scales represent 1 cm.
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Fig. 29.— Par aconularia subulata (Hall). 29. 1; AMNH 32404 (smaller of two specimens), lectotype, minor face; locality 16. 29.2; UMMP 245, holotype of Conularia newberryi
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the Lower Mississippian of the North American midcontinent. It is often confused with other taxa, especially P. missouriensis, in museum collections. The reason for this confusion is not clear. Some specimens of this taxon, which superficially appear very similar to specimens of P. biairi, may have been confused with P. missouriensis because of ambiguity in Swallow’s original description of the latter. It is likely that Swallow’s description was based upon specimens now referable to both P. missouriensis and P. biairi.
Conulariids collected from the Bear Gulch Limestone of Montana (CM 34507--34527 and 35000) and from the Cameron Creek Shale of Montana (USNM 118731) are here assigned to P. subulata with little reservation. The specimens differ from the type series of P. subulata only in the uniform lack of nodes on the rods. However, samples of many specimens referable to P. subulata from Illinois, Ohio, and else- where indicate that nodes are frequently lacking in this taxon. Even when nodes are present on such specimens, they are subtle.
Material examined. — I A9 specimens; housed in the AMNH, CM, CMNH, FMNH, GSC, OC, UMMP, USNM, and the private collection of Ron Fisher.
PARACONULARIA WELLSVILLIA Babcock and Feldmann, new species Figs. 33.3, 33.6-^33.8
Paraconularia sp. Babcock and Feldmann, 1986, fig. 2J.
Description. —Exosk&lQlon up to 13 cm in length. Major apical angle 14-18°; minor apical angle 12-15°. Rod articulation gothic arch style in apical region and inflected gothic arch style elsewhere; rods almost always alternate at the midline; if they alternate, pattern is usually left superior on major face and usually right superior on minor face; rod angle 26-31°. 4-5 rods/cm; 2-3 nodes/mm; 2-3 adapertural spines/mm; adapical spines appear not to be present. Apical wall not observed.
Occurrence. Devonian of New York; localities 161“162,
Holotype, CM 35001; 12 paratypes, CM 34538-34550. Remarks. —Paraconularia wellsvillia Babcock and Feldmann, n. sp.
Winchell, major face; locality 206. 29.3; UMMP 245, same specimen as in Fig. 29.2, minor face. 29.4; AMNH 32404 (smaller of two specimens), same specimen as in Fig. 29.1, major face. The pitted material attached to the specimen is glue. 29.5; CMNH 5988, comer view of large, partially compressed specimen. Note healed injury near top of minor face; locality 198. 29.6; UMMP 2178, holotype of C. victa White, ?minor face; locality 63. 29.7; UMMP 2178, same specimen as in Fig. 29.6, comer view. 29.8; FMNH UC 6961, holotype of C. sampsoni Miller, minor face; locality 94. 29.9; FMNH UC 6961, same specimen as in Fig. 29.8, comer view. 29.10; FMNH UC 6961, same spec- imen as in Fig. 29.8, major face. Bar scale in Fig. 29.5 represents 1 cm; bar scales in Figs. 29.1-29.4 and 29.6-29.10 represent 5 mm.
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is distinguished from all other species of Paraconularia known from the Devonian and Mississippian rocks of North America in having gothic arch rod articulation in the apical region and inflected gothic arch rod articulation elsewhere. Superficially, this taxon resembles P. yochelsoni in the general pattern of rod articulation near the aperture. P. wellsvillia, however, possesses nodes and adapertural spines whereas P. yochelsoni does not.
Of the 13 specimens examined and referred to P. wellsvillia, only the holotype, CM 35001, shows well preserved nodes and spines (Figs. 33.7-33.8). Others, such as the specimen illustrated in Fig. 33.6, seem to lack these structures. After examination of the holotype and 12 paratypes, it seems that two factors affect these profound preservational differences: 1, degree to which the integument is fit around the rods, nodes, and spines; and 2, type of lithology in which the specimen is preserved. The Wellsville Formation, from which all specimens in the type suite were collected, varies from a fine grained silty sandstone to a micaceous siltstone to a micaceous shale. Preservation of a conulariid tends to be better in a fine grained matrix.
The holotype of P. wellsvillia shows a wrinkling of the integument only partially related to the pattern of nodes and spines (Fig. 33.8). This wrinkling is also attributed, in part, to a tight fitting of the integ- ument about the framework of the exoskeleton and slight displacement of the framework.
Material examined. — 13 specimens; housed in the CM.
Fig. 30. —Paraconularia subulata (Hall). 30.1; USNM 409802, minor face of specimen preserved in phosphatic concretion; locality 72. 30.2-30,3; USNM 395828, right and left halves of a specimen preserved in a phosphatic concretion and showing internal soft- parts; locality 72. 30.4; USNM 409802, same specimen as in Fig. 30.1, view from apical end showing rounded cross section of soft-parts (arrow). 30.5; USNM 409802, same specimen as in Fig. 30.1, x-ray photograph of specimen preserved in a phosphatic con- cretion. The photograph was obtained using a Hewlett-Packard Faxitron Series x-ray unit located in the Department of Anthropology, Kent State University. The specimen is in the same orientation as in Fig. 30. 1 . Presumed internal soft-parts appear as a single, elongate tube. 30.6; USNM 409803, specimen preserved in a phosphatic concretion with some of the exoskeleton broken away, revealing remains of limonite-coated internal soft- parts. 30.7; USNM 409804, two specimens preserved in same orientation in siltstone block. Sole marks on reverse side of slab parallel the orientation of the conulariids and indicate that these specimens have been current aligned; locality 220. 30.8; UIPC 10866, holotype of Conularia whitei Meek and Worthen preserved in siderite; locality 20 1 . Bar scales in Figs. 30.1-30.3 and 30.5-30.8 represent 1 cm; bar scale in Fig. 30.4 represents 5 mm. Specimens in Figs. 30.2-30.4 and 30.6 have not been coated with ammonium chloride.
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Etymology of trivial we. —Named for the Wellsville Formation, from which the holotype specimen was collected.
PARACONULARIA YOCHELSONI
Babcock and Feldmann, new species Figs. 28.7, 33.1-33.2, 33.5
Paraconularia newberryi (Winchell). Sensu Driscoll, 1963, p. 34-40, PL 1, figs. 6-8.
Description. —Exoskeleton up to 3.5 cm in length. Major apical angle 17-20®; minor apical angle approximately 18°. Rod articulation gothic arch style in earliest stages, inflected gothic arch style in later stages. Rods usually alternate at midline; if they alternate, pattern is usually right superior on both major and minor faces; rod angle 1 5- 20°. 13-18 rods/cm. Nodes appear to be absent; spines absent. Apical wall present.
Occurrence. —Lower Mississippian of Michigan; locality 93.
Types. — Holotype, UMMP 45499; two paratypes, UMMP 65509 on the same slab as UMMP 45499, and UMMP 45500.
Remarks. —Paraconularia yochelsoni Babcock and Feldmann, n. sp. is only similar in morphology to P. alternistriata (Shimer). Both species seem to be less than 3.5 cm in maximum length, lack nodes on the rods and have rod articulation patterns which appear to be similar. Paraconularia yochelsoni, however, has larger apical angles, 1 5-20° as compared to 10-1 1°, and fewer rods/cm, 13-15 as compared to 28.
The holotype, UMMP 45499, and paratype, UMMP 65509, speci- mens of P. yochelsoni, are preserved as three dimensional specimens lacking the integument except along the midline. The midline may have been thickened in this taxon.
The holotype and paratype of P. yochelsoni are located on the same slab as a large portion of black, carbonaceous matter composed largely of densely packed, filamentous strands (Fig. 28.7). This material prob- ably represents plant matter of some sort, perhaps a planktonic alga. The two conulariids appear to be attached to the presumed plant matter by stalks extending from their apices; only small traces of the stalks remain in place. The two conulariids on this slab are radiating away from the center of the dark mass.
Material examined. — 3 specimens; housed in the UMMP.
Etymology of trivial ^aw^.— Named for Ellis L. Yochelson, a dis- tinguished student of problematic fossils.
Fig. 31.— Paraconularia subulata (Hall). 31.1; CM 34524, preserved in micrite; locality 106. 31.2; CM 35000, preserved in micrite; locality 106. 31.3; CM 34521, preserved in micrite, locality 106. 31.4; NYSM 3491, preserved in siderite; locality 203. Original of Hall’s ^Eonularia newberryE (1879, PI. 34A, fig. 12). 31.5; USNM 118731; locality 107. Bar scales represent 1 cm.
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Fig. 32. — 32.1; Paraconularia missouriensis (Swallow)?, GSC 85062; locality 5. 32.2; P. sp., USNM 409805, fragment of specimen lacking integument preserved in soft blue-
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PARACONULARIA sp.
Fig. 28.4
Occurrence.— Upper Devonian-Lower Mississippian transition of South Dakota; locality 27.4.
Figured specimen. — CM 3453 1 .
Remarks.— A single specimen of conulariid was collected from the Englewood Formation at Deadwood, South Dakota. It can be reliably identified only to the genus level.
The specimen is badly crushed and incomplete, making it possible to perform few qualitative or quantitative observations. The specimen in question possesses 7 rods/cm and lacks nodes and spines; it is there- fore referred to the genus Paraconularia. The specimen seems to have a gothic arch style of rod articulation in the apical region and an in- flected circular curve style elsewhere.
Species of Paraconularia examined in this work which may possess 7 rods/cm include P. blairi (Miller and Gurley) and P. missouriensis (Swallow); P. chesterensis (Worthen) can have as few as 8 rods/cm and P. wellsvillia Babcock and Feldmann, n. sp. has 4-5 rods/cm. Of these, CM 34531 appears to be most similar to P. blairi or P. subulata (Hall) in terms of rod articulation.
Material examined. — I specimen, CM 34531.
RETICULACONULARIA Babcock and Feldmann, new genus
— Conulariids with rods that are widely spaced, 12-39/ cm. 30-80% of rods alternate at midline; 20-70% abut. Apical angles large, 22-59°. Nodes and adapertural spines present and widely spaced; adapical spines not known.
Type species. — Conularia penouili Clarke, 1907; Lower Devonian of Quebec. Holotype, NYSM 9412.
Remarks. — Species referable to Reticulaconularia differ from all oth- er conulariids in having very large apical angles, 22-59° in the speci- mens measured in this study. The wide spacing between adjacent rods and between nodes, as well as between adapertural spines is also unique to species of this genus. In specimens retaining the external surface of the integument, this pattern of rods, nodes, and spines gives the exo-
gray calcareous shale; locality 177. 32.3; P. recurvatus Babcock and Feldmann, n. sp., USNM 409806. Holotype to right; a paratype (USNM 409807) is indicated by arrow; locality 109. 32.4; P. byblis (White)?, UMMP 26735, a poorly preserved, collapsed specimen; locality 250. Ori^nal of Winchell (1871, p. 257). 32.5; P. chesterensis (Wor- then)?, GSC 49383, a juvenile specimen with stalk preserved. Specimen not coated with ammonium chloride; locality 7. 32.6; P. recurvatus Bdhcock and Feldmann, n.sp., USNM 409806, same specimen as in Fig. 32.3, detail of holotype. Bar scales represent 1 cm.
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skeleton a somewhat reticulate pattern. The genus is named for this
characteristic.
At present, three species of conulariids are referred with certainty to Reticulaconularia: C. penouili from the Lower Devonian of Quebec, C. sussexensis Herpers from the Lower Devonian of New Jersey, and C baini Ulrich from the Devonian of Bolivia. This third species occurs outside the geographic limits of this paper; we and others will redescribe the taxon in a paper on the Devonian conulariids of Bolivia.
RETICULACONULARIA PiE^A^OU/L/ (Clarke, 1907)
Figs. 34.3-34.5; 35.3
Conularia penouili Clarke, 1907, p. 180--181, 2 figs.; Clarke, 1908, p. 144, PL 11, figs. 10-=11.
Conularia gaspesia Sinclair, 1942, p. 15 8-- 160, fig.
Conularia s.l. penouili Clarke. Sinclair, 1948, p. 283.
Description. —ExoskQlQton a curved pyramid, expanding slowly and non-uniformly from the apex. Exoskeleton up to 6.7 cm in length. Major apical angle approximately 59°; minor apical angle approximately 22-30°. Rod articulation of inflected circular curve style in apical ‘A, angulated circular curve style in middle Vs, and inflected circular curve style in apertural '/a. Rods usually alternate at midline; if they alternate, pattern is usually left superior on both major and minor faces; rod angle 0-8°. 12-21 rods/cm. 1-2 nodes/ mm. 1-2 adapertural spines/cm; adapical spines probably not present. Interridge furrows broadly rounded into longitudinally oblong pits. Apical wall not observed.
Occurrence. — Lowqy Devonian of Quebec; localities 228 and 246. Type'y. —Holotype, NYSM 9412; topotype of Conularia gaspesia,
GSC 87242.
Remarks. —Reticulaconularia penouili (Clarke) is distinguished from R. sussexensis (Herpers) by the following features. First, R. penouili has 12-21 rods/cm and R. sussexensis has 11-14 rods/cm. Greater rod spacing in R. penouili, combined with wide spacing between the nodes and between the adapical spines has produced large, oblong, hollowed
Fig. 33,-33. 1-33.2, Paraconularia yochelsoni Babcock and Feldmann, n. sp. 33, 1 ; UMMP 45499, holotype; comer view; locality 93. 33.2; UMMP 45499, same specimen as in Fig. 33.1, major face. 33.3; P. wellsvillia Babcock and Feldmann, n. sp., CM 34502, paratype, preserved in siltstone; ?minor face; locality 161. 33.4; P. subulata (Hall), KSU 1 172, cross section showing weakly bilateral, four-sided nature of the exoskeleton. Spec- imen not coated with ammonium chloride; locality 216. 33.5; P. yochelsoni Babcock and Feldmann, n, sp, UMMP 45500, paratype; locality 93. 33.6-33.8; P. wellsvillia Babcock and Feldmann, n. sp. 33.6; CM 34503, detail of paratype preserved as an internal mold in siltstone and not exhibiting interrod ridges or interrod furrows; locality 162. 33.7; CM 35001, holotype, a collapsed specimen preserved in silty shale; locality 161. 33.8; CM 3500 1 , same specimen as in Fig. 33.7, detail of a minor face. Bar scales represent 1 cm.
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Fig, 34.-34. 1-34.2; Reticulaconulariasussexensisi^QrpQT^). 34. 1; NJSM 10806, smaller of two specimens, major face of small specimen preserved in somewhat metamorphosed
1986 Babcock and Yeldmann—Paraconularia and Reticulaconularia
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out interridge furrows on the exterior surface of the exoskeleton. This feature is not exhibited, to this extent, on any other known species of conulariid.
Secondly, the values obtained for rod angles are consistently smaller for R. penouili than they are for R. sussexensis, 0-8° compared to 1 1- 14°. Reticulaconularia sussexensis exhibits a greater number of rods which abut at the midline than does R. penouili. Up to 70% of rods abut in specimens of R. sussexensis while 20-30% of rods abut in the holotype of R. penouili. Unlike R. sussexensis, R. penouili shows an alternation between a circular curve style of rod articulation and an angulated circular curve style. Reticulaconularia sussexensis exhibits only an inflected gothic arch style of rod articulation. Finally, judging from the available sample, specimens of R. sussexensis seem to have a smaller maximum length than R. penouili. The maximum recorded hypothetical length of a specimen of R. sussexensis is about 2.5 cm. The hypothetical length of the holotype of R. penouili is 6.7 cm.
The holotype of Conularia gaspesia Sinclair, which was said to have been deposited in the RM(MU) (Sinclair, 1942, p. 160) apparently never was deposited in that museum (Ingrid Birker, written commu- nication, 1985), and is now presumed to be lost. However, we have found, in the Sinclair collection housed in the GSC, a specimen labelled as C. gaspesia (GSC 87242; Fig. 35.3). The handwriting on a label accompanying the specimen is unmistakably that of G. Winston Sin- clair. This specimen is of further value because a label glued to the specimen indicates that it was collected from Lower Devonian Grande Greve Limestone on the Gaspe Peninsula; thus, the specimen is a topotype.
This topotype specimen of C. gaspesia, GSC 87242, exhibits one well preserved face and has all of the salient morphological character- istics that the holotype of R. penouili posseses. Among the character- istics shown by the topotype of C. gaspesia are large, oblong, hollowed out interridge furrows, just as are present in the holotype of R. penouili. A good photograph of the holotype of C. gaspesia (Sinclair, 1 940, fig.) shows these same features. Measurements taken on the topotype spec- imen are given in Appendix B. The only way in which GSC 87242 differs from the holotype of R. penouili, NYSM 9412, is that it is not
siltstone; locality 1 12, 34,2; NJSM 10806, same specimen as in Fig, 34.1, comer view. Note prominent exoskeletal constriction. 34. 3-34.5; R. penouili (Clarke). 34.3; NYSM 9412, holotype, detail of minor face; locality 246. 34.4; I^SM 9412, same specimen as in Fig. 34,3, minor face of specimen preserved in micrite. 34.5; NYSM 9412, same specimen as in Fig. 34,3, major face. Bar scales represent 1 cm.
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Fig. 35.-35.1-35.2. Reticulaconularia smsexensis 35.1; NJSM 10750, exter-
nal mold of apical region of very small paratype specimen; locality 112. 35.2; NJSM 10751, largest of three specimens, paratype, preserved as an external mold; locality 1 12. 35.3; R. penouiii (Clarke), GSC 87242, ?minor face of topotype of Conularia gaspesia
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curved in the apical region. However, NYSM is a somewhat crushed or collapsed individual. One final interesting point is that both NYSM 9412 and GSC unassigned were collected from the Grande Greve Lime- stone on the Gaspe Peninsula. Based upon this topotype specimen of C gaspesia, this taxon is here placed in synonymy with R. penouili.
Material examined.— 2 specimens; housed in the GSC and the NYSM.
RETICULACONULARIA SUSSEXENSIS {ULqvpqxs, 1949)
Figs. 34.1-34.2, 35.1-35.2, 35.4
Conularia sussexensis Herpers, 1949, p, 1-7, PL 1, 2.
Conularia gaspesia Sinclair. Sensu Herpers, 1950, p. 619.
Description. —FxoskeXeXon up to 2.5 cm in length. Major apical angle 24-28°; minor apical angle 22-25°. Rod articulation uniformly of inflected gothic arch style. Rods usually abut at midline; if they alternate, they usually occur left superior on major face and right superior on minor face; rod angle 11-14°. Approximately 39 rods/cm (extrapolated) in apical region; 14-18 rods/cm elsewhere. 2 nodes/mm; 2 adapertural rods/mm; adapical spines absent. Apical wall not observed.
Holotype, NJSM 10749; four paratypes,NJSM 10750, NJSM 10751 (three specimens).
Occurrences. — Lowqt Devonian of New Jersey; localities 111, 112, 113.
Remarks.— ThQ present samples of R. sussexensis (Herpers) differ from the holotype of R. penouili (Clarke) in that the former: 1, is of smaller size; 2, has no curvature to the exoskeleton; 3, has smaller apical angles, especially on the major face; 4, has a greater number of rods/cm; 5, has larger rod angles; 6, shows greater than 30% of the rods abutting at the midline; and 7, shows only an inflected gothic arch style of rod articulation. These differences are discussed more fully in the remarks accompanying the description of R. penouili, above.
Material examined. — \ 0 specimens; housed in the NJSM.
Organisms Previously Assigned to Conulariida,
Here Rejected from the Phylum
Phylum Mollusca Class Hyolitha HYOLITHES sp.
Fig. 17.4
Conularia sp. Ellison, 1965, p. 48-49, PL 4, fig. 1.
Hyolithes sp. Babcock, 1985^, p. 14-16, fig. 1.
Sinclair, preserved in micritic limestone; locality 242. 35.4; R. sussexensis (Herpers), NJSM 10749, ?major face of holotype, preserved in somewhat metamorphosed siltstone; locality 112. Bar scales represent 1 cm.
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Fig. 36. — 36.1-36.3; ^'‘Conularia" tenuicostata Branson, here interpreted to be a ?pri- apulid worm. 36.1; UMC 4271, holotype; locality 99. 36,2; UMC 4271, detail of same
1986 Babcock and Feldmann—P^/?^ and Reticulaconularia
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Occurrence.— Middle Devonian of Pennsylvania; locality 240.
Figured sped men. — U SNM 173928.
Remarks. —Ellison’s figured specimen (1965, plate 4, fig. 1) is a small conical shell expanding slowly and uniformly from a bluntly rounded apex. The specimen possesses thin, closely spaced, raised lines, con- centric about the apex. Crushing has produced a long, irregular line down the middle of the shell. A ligula, or an apertural extension of the shell on the dorsal side, is present, clearly indicating that the specimen is a hyolith, not a conulariid.
Phylum Priapulida?
Figs. 36.1-36.3
Conularia tenuicostata Branson, 1938, p. 1 1 1, PL 14, figs. 5-6; Branson, 1944, p, 216. Mesoconularia tenuicostata (Branson). Sinclair, 1948, p. 128.
Occurrence. — Fowtr Mississippian of Missouri; locality 99.
Type. -Holotype, UMC 4271.
Remarks.— The holotype, and only known specimen of Conularia tenuicostata Branson, possesses tiny nodes or pustules arranged in closely spaced rows which appear to run essentially perpendicular to the long axis of the fossil. The rows of nodes or pustules are not supported by calcium phosphate rods. Additionally, the specimen is flattened and micrite replaced. This type of preservation is unlike that expected of an animal composed of calcium phosphate, such as a conulariid. Rath- er, the preservation is similar in appearance to the preservation of objects having a tough cuticle, such as Plectodiscus discoideus (Rauff), a chondrophorine cnidarian from the Hunsriick Slate (Devonian) of West Germany (Yochelson et aL, 1983).
Branson’s specimen possesses indiscrete ringlike segments delimited by thin, latitudinally arranged crests 0.5 to 0.7 mm apart. These seg- ments are each covered with closely spaced, latitudinally arranged rows of minute papillae which are strikingly similar to the cuticle of living priapulid worms such as Priapulus and Tubiluchus. However, there is not enough of the holotype preserved to determine whether the animal possessed spines, a common feature of living priapulids. Thus, this fossil is referred to the phylum Priapulida with reservation.
The specimen in question preserves only a small portion of cuticle,
specimen as in Fig. 36.1, showing ridges delimiting annular segments. 36.3; detail of same specimen as in Fig. 36.1, showing surface structure. 36.4-36.5; Oracanthus sp. 36.4; USNM 409810; locality 6. 36.5; USNM 409810; detail of specimen in Fig. 36.5, interpreted to be remains of fish spine. Bar scales in Figs. 36.1 and 36.4-36.5 represent 1 cm; bar scales in Figs. 36.2 and 36.3 represent 1 mm.
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7.5 mm long and 8.1 mm wide. It is broken at its upper and lower margins, presumably between adjacent segments. This mode of pres- ervation supports the interpretation that the specimen possessed a multielement covering, capable of readily fragmenting or tearing.
Phylum Chordata Class Vertebrata Order Pisces ORACANTHUS sp.
Figs. 36.4-36.5
Conularia newberryi Winchell? Sensu McKee and Gutschick in McKee and Gutschick, 1969, p. 125-172.
Occurrence.— Mississippian of Arizona; locality 5.
Figured specimen. — External mold preserved in dolostone and latex mold, USNM 409810.
Remarks.— This figured specimen mimics a conulariid in having nodose structures arranged in rows, crossing the surface transversely. The rows, however, are discontinuous, and form chevron-shaped pat- terns in some places. The rows seem to be composed of semidiscrete pits arranged in side-by-side fashion. As the specimen is an external mold, the “pits” would correspond to nodes. These is no evidence on the specimen of either a midline or a comer groove. The specimen is undulated near the left margin, however.
This specimen is here regarded as the spine of a gyracanthid shark. Michael E. Williams, of the Cleveland Museum of Natural History, has viewed this specimen and has noted that small portions of dentine adhere to it in places, confirming that it is a vertebrate fossil. It is his suggestion that this specimen be referred to the genus Or acanthus.
Acknowledgments
Parts A and B of this work are based upon Babcock’s M.S. thesis, completed at Kent State University. The study was suggested by Ellis L. Yochelson, U.S. National Museum of Natural History, Washington, D.C. Alan H. Coogan and Barry B. Miller read various drafts of this paper. Others who have assisted in this study through collecting specimens, loaning specimens, providing reference materials or aiding with electron microprobe or x-ray analyses include the following: Donald Baird, Princeton University; Gorden C. Baird, SUNY College at Fredonia; Roger L. Battin, American Museum of Natural His- tory; Gordon L. Bell, Jr., Red Mountain Museum; Ingrid Birker, Redpath Museum (McGill University); Daniel B. Blake, University of Illinois at Urbana-Champaign; Thomas H. Bolton, Geological Survey of Canada; Arthur J. Boucot, Oregon State University; Ernest H. Carlson, Kent State University; Robert L. Carroll, Redpath Museum (McGill University); John L, Carter, Carnegie Museum of Natural History; Mitchell J. Ciccarone, Canton, Ohio; Frederick J. Collier, United States National Museum of Natural History; G. Arthur Cooper, United States National Museum of Natural History; Murray J. Cope- land, Geological Survey of Canada; Roger J. Cuffey, Pennsylvania State University; Larry Decina, Drexel Hill, Pennsylvania; Ding Baoliang, Nanjing Institute of Geology and
1986 Babcock and Felumann —Paraconularia and Reticulaconularia
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Mineral Resources, Nanjing, People’s Republic of China; Ruth L. Elder, Oberlin College; Niles Eldredge, American Museum of Natural History; Frank R. Ettensohn, University of Kentucky; David F. Factor, Hiram, Ohio; Howard R. Feldman, American Museum of Natural History; Daniel C. Fisher, University of Michigan; Ron Fisher, Homer, Ohio; Raymond C. Gutschick, University of Notre Dame; Kurt F. Hallin, Milwaukee Public Museum; Joseph T. Hannibal, Cleveland Museum of Natural History; Alan Stanley Horowitz, Indiana University; Thomas W. Kammer, West Virginia University; Gerald J. Kloc, University of Rochester; Albert D. Kollar, Carnegie Museum of Natural History; Ed Landing, New York State Geological Survey; Ralph L. Langenheim, University of Illinois at Urbana-Champaign; Richard S. Laub, Buffalo Museum of Science; Richard L. Leary, Illinois State Museum; Pierre J. Lesperance, Universite de Montreal; Richard Lindemann, Skidmore College; Richard Lund, Adelphi University; Royal H. Mapes, Ohio University; Dagmar Merino, Yacimientos Petroliferos Fiscales Bolivianos, La Paz, Bolivia; Jonathon Mortin, University College of Swansea; Matthew H. Nitecki, Field Museum of Natural History; James C. Ohman, Kent State University; William A. Oliver, Jr., United States Geological Survey, Washington, D.C.; David C. Parris, New Jersey State Museum; the late Eugene S. Richardson, Jr., Field Museum of Natural History; Gabrielo Rodrigo, Museo Nacional de Historia Natural, La Paz, Bolivia; Edgar Roeser, Cleveland Museum of Natural History; Colin T. Scrutton, University of Newcastle- upon-Tyne; Robert Segedi, Cleveland Museum of Natural History; Peter F, Sheehan, Milwaukee Public Museum; Thomas M. Stanley, St. Joe Mining, Deadwood, South Dakota; James T. Stitt, University of Missouri-Columbia; Mario Suarez-Riglos, Yaci- mientos Petroliferos Fiscales Bolivianos, Santa Cruz, Bolivia; Robert Walker, Kent State University; Steven C. Ward, Kent State University; Lawrence A. Wiedman, Monmouth College; Michael E. Williams, Cleveland Museum of Natural History; Margaret T. Wil- son, Kent State University; and Paul Zell, State College, Pennsylvania.
We would especially like to thank Margaret Wilson for her unflagging support and encouragement of the research and for her review of the final manuscripts; without her, these papers would have been impossible. Ellis L. Yochelson and Richard H. Lindemann reviewed the manuscripts for the Annals of the Carnegie Museum. This study supported in part by American Association of Petroleum Geologists Grant-in-Aid no. 582-12-01 and by a Grant-in-Aid of Research from Sigma Xi, The Scientific Research Society, both to Babcock. A portion of the cost of publication has been provided by the Office of Research and Sponsored Programs, Kent State University. Part A is contribution 313 and Part B is contribution 314 of the Department of Geology, Kent State University, Kent, Ohio 44242.
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— . 1 900Z). The succession of fossil faunas in the Kinderhook beds at Burlington,
Iowa. Iowa Geological Survey, 10:59-79.
White, C. A. 1862. Description of new species of fossils from the Devonian and Carboniferous rocks of the Mississippi Valley. Boston Society of Natural History, Proceedings, 9:8-33.
. 1880. Fossils of the Indiana rocks. Indiana Department of Statistics and Ge- ology, 2nd Annual Report, pp. 471-552.
Whiteaves, J. F. 1891. The fossils of the Devonian rocks of the Mackenzie River Basin. Geological and Natural History Survey of Canada, Contributions to Canadian Palaeontology, 1:197-253.
Whitfield, R. P. 1882. On the fauna of the Lower Carboniferous limestones of Spergen Hill, Ind., with a revision of its fossils hitherto published, and illustrations of the species from the original type series. American Museum of Natural History, Bulletin, l(3):39-97.
Whitfield, R. P., and E. O. Hovey. 1901. Catalogue of the types and figured specimens in the paleontological collection of the Geological Department, American Museum of Natural History. Part IV. Carboniferous to Pleistocene, inclusive. American Mu- seum of Natural History, Bulletin, 1 1:357-500.
WiNCHELL, A. 1865. Descriptions of new species of fossils from the Marshall Group of Michigan, and its supposed equivalent, in other states; with notes on some fossils of the same age previously described. Academy of Natural Sciences of Philadelphia, Proceedings for 1865, pp. 109-133.
1986 Babcock and Felumann—Paraconularia and Reticulaconularia
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1870. Notices and descriptions of fossils, from the Marshall Group of the
western states, with notes on fossils from other fossils. American Philosophical Society, Proceedings, 11:245-260.
WoRTHEN, A. H. 1883. Description of some new species of fossil shells from the Lower Carboniferous limestones and Coal Measures of Illinois. Geological Survey of Illi- nois, 7:323-326.
1890. Description of fossil invertebrates. Geological Survey of Illinois, 8:69-
154.
Yochelson, E. L., G. D. Stanley, and W. Sturmer. 1983. Plectodiscus discoideus (Rauff): a redescription of a chondrophorine from the Early Devonian Hunsriick Slate, West Germany. Palaontologie Zeitschrift, 57:39-68.
Appendix A— Locality Index
Collector, year of collection if known, description of locality, stratigraphic assignment and conulariid taxa present
Alabama
1 A. S. Horowitz. “Keyes site,” Skyline, Alabama. Formation unknown; probably Chesterian Series. Paraconularia chesterensis.
Alberta
2 Anonymous. Near Banff, Alberta. Formation unknown and series unknown; Mis- sissippian Subsystem. Paraconularia missouriensisl
3 R. G. McConnell, 1890. Athabasca River, La Saline, Alberta. Formation unknown; Chautauquan Series. Paraconularia salinensis.
4 H. W. Shimer, pre-1926. Lake Minnewanka, Alberta. Formation and series un- known; Mississippian Subsystem. Paraconularia alternistriata.
5 F. Beales. Upper part of Job Creek, western Alberta. Upper Rundle Formation, 345 m from the base of the formation; Chesterian Series. Paraconularia missouriensisl
Arizona
6 R. C. Gutschick and P.C.H., 1954. Top of mesa on point between Rock and Blye Canyons on 7BarV Ranch, WF Cattle Company, south of Peach Springs and Cher- okee Point, Arizona. Chert in Member 2 of Redwall Limestone, about 53 m above base of Redwall Formation; ?Osagean Series. No conulariids collected; Oracanthus spine.
British Columbia
7 D. Scott, 1962. Spur on northeast comer of Mt Hosmer, 14.5 km northeast of Femee and 14.5 km southwest of Natal, British Columbia. Lower Etherington Member of the Rocky Mountain Formation of the Rundell Group; Chesterian Series. Paraconularia chesterensisl
Illinois
8 Anonymous. Kinderhook, Pike County, Illinois. Kinderhook Group; Kinderhook- ian Series. Conularia subcarbonaria.
9 W. F. E. Gurley; Anonymous. Hamilton, Illinois. Keokuk Limestone; Osagean Series. Conularia subcarbonaria.
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10 Anonymous. Chester, Randolph County, Illinois. Chester Limestone; Chesterian
Series. Paraconularia chesterensis.
1 1 Sloss. Pike County, Illinois. Burlington Limestone; Osagean Series. Paraconularia
chesterensis, P. blairi.
12 S. Weller, 1912. About 2.4 km south of Marigold, Illinois. “Lower Okaw, Marigold Oolite” (=Burlington Limestone?); Osagean Series. Paraconularia chesterensis.
13 W. F. E. Gurley; Anonymous. Hamilton, Illinois. Keokuk Limestone; Osagean Series. Conularia subcarbonaria.
14 W. F. E. Gurley. Madison County, Illinois. St. Louis Limestone; Meramecian series. Paraconularia missouriensis.
1 5 Anonymous. Warsaw, Madison County, Illinois. St. Louis Limestone; Meramecian Series. Paraconularia missouriensis.
16 Anonymous. Alton, Illinois. St. Louis Formation; Meramecian Series. Paraconu- laria subulata, P. chesterensis.
17 W. F. Gurley. Madison County, Illinois. St. Louis Limestone; Meramecian Series. Paraconularia chesterensis.
1 8 S. Weller, 1912. About 3.2 km east of Waterloo, Illinois, Illinois. Renault Limestone; Chesterian Series. Paraconularia chesterensis.
19 S. Weller, 1918. About 4 km southeast of Vienna, Illinois. Glen Dean Limestone; Chesterian Series. Paraconularia chesterensis.
20 A. S. Horowitz, 1966. Debris from slope below quarry in Mississippi River bluffs above Illinois State Highway 3 Bypass, SW'A, SW‘/4 Sec.29, TVS, R6W, Chester, Randolph County, Illinois, Chester 7.5' Quadrangle. Menard Limestone; Chesterian Series. Paraconularia chesterensis, Conularia cf. C. subcarbonaria.
21 L. F. Rauchfrise; Anonymous. Pope County, Illinois. Chester Group; Chesterian Series. Paraconularia chesterensis.
Indiana
22 E. M. Kindle. Delphi, Indiana. Sellersburg Formation; Chautauquan Series. Con- ularia delphiensis.
23 D. G. Maroney and R. W. Orr, pre-1974. Delphi Limestone Company Quarry, north side of U.S. Highway 421, northwest edge of Delphi, SWV4, SW‘/4 Sec. 19, T25N R2W, Carroll County, Indiana, Delphi 7.5' Quadrangle. 0-10 cm thick phos- phatic pebble bed at base of New Albany Shale; Chautauquan Series. Conularia delphiensis.
24 D. E. Hattin? Probably Indiana. Probably Harrodsburg Formation; Osagean Series.
Conularia subcarbonaria.
25 D. E. Hattin. Hattin location S-776. Indiana. Borden Group; Meramecian Series. Conularia multicostata.
26 Anonymous. Curiosity Hollow, near Martinsville, Indiana. New Providence For- mation; Osagean Series. Paraconularia cf. P. subulata.
27 Anonymous. Crawfordsville, Indiana. “Keokuk Group” (=Borden Group); Osagean Series. Conularia subcarbonaria, Paraconularia chesterensis.
28 W. F. E. Gurley. West Point, Indiana. “Keokuk Group” (=Borden Group); Osagean Series. Conularia subcarbonaria.
29 G. K. Greene; Washburn. New Albany, Indiana. “Knobstone Group” (=Borden Group); Osagean Series. Paraconularia byblis, P. subulata.
30 Anonymous. Spergen Hill, Indiana. St. Louis Limestone; Meramecian Series. Par- aconularia missouriensis.
31 W. F, Gurley; Anonymous. Edwardsville, Indiana. “Keokuk Group” (=Borden Formation); Osagean Series. Paraconularia missouriensis.
32 Klippart. New Providence, Indiana. Carwood Member, Borden Formation; Osagean
Series. Paraconularia blairi.
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33 E. O. Ulrich. Crawfordsville, Indiana. Borden Formation (“Keokuk Group”); Osa-
gean Series. Paraconuiaria chesterensis.
34 Anonymous. Near Providence, Indiana. Borden Formation; Osagean Series, Par- aconuiaria blairi.
35 C. Rominger? Crawfordsville, Montgomery County, Indiana. Near middle of Bor- den Group; Osagean Series. Paraconuiaria chesterensis.
36 S. Makvrat. Bed of Gnaw Bone Creek just south of Indiana Highway 46, just east of Gnaw Bone, Indiana, Nashville 7.5' Quadrangle. New Providence Shale according to collector; more likely Carwood or Locust Point Formation of Borden Group according to A, S. Horowitz (written communication, 1984); ?Osagean Series. Par- aconuiaria cf. P. byblis, P. chesterensis.
37 R. Fields and J. Harris, 1981. Section on old Indiana Highway 37, SE'A, NW*/}, NW*/4 Sec.21, T9N, RIW, Monroe County, Indiana, Bloomington 7.5' Quadrangle. Ramp Creek Member of Harrodsburg Limestone, 3-7 m above contact with Borden Group; ?Osagean Series. Conularia subcarbonaria.
38 J. Hall?; Washburn; Anonymous. Crawfordsville, Indiana. “Keokuk Group” (=Bor- den Group); Osagean Series. Conularia subcarbonaria, Paraconuiaria chesterensis, P. subulata.
39 W. F. E. Gurley; G. Robb, pre-1923. New Albany, Indiana. “Knob or Knobstone Shale” (=Borden Group?); Osagean Series. Conularia multicostata, Paraconuiaria byblis, P. chesterensis, P. missouriensis, P. subulata.
40 W. F. E. Gurley. West Point, Indiana. “Keokuk Group (=Borden Formation)”; Osagean Series. Paraconuiaria chesterensis.
41 R. L. Anstey et al., 1968. Bed of Indian Creek on O. C. Bennett or Ben Wilson property, approximately .8 km north of Indiana Highway 234, NE'A, NW ‘A, SE'A Sec, 8, T17N, R5W, Montgomery County, Indiana, ?Edwardsville Formation of Borden Group; Osagean Series. Paraconuiaria cf. P. chesterensis.
42 C. Rominger? Crawfordsville, Montgomery County, Indiana. Near middle of Bor- den Group; Osagean Series. Conularia subcarbonaria.
43 G. Campbell. Floyds Knob Hill, Highway 150, near center of NE'A 28-25-6E {sic), 3.2 km northwest of New Albany, Indiana. Kenwood Formation; ?Osagean Series. Paraconuiaria byblis.
44 W. F. E. Gurley. Spergen Hill, Indiana. Probably St. Louis Limestone, though possibly Salem Limestone; Meramecian Series. Conularia subcarbonaria.
45 W. F. E. Gurley. Spergen Hill, Indiana. St. Louis Limestone or Salem Limestone; Meramecian Series. Conularia subcarbonaria.
46 Anonymous. Salem, Indiana. Salem Limestone; Meramecian Series. Conularia sub- carbonaria.
47 A. S. Horowitz, N. G. Lane et al. Outcrop along west branch of Mosquito Creek, 0.48 km west and 0.24 km north of southeast comer of Sec. 25, T5S, R5E, ap- proximately 4.5 km east of Laconia, Harrison County, Indiana; Laconia 7.5' Quad- rangle, Somerset Shale; Chesterian Series. Paraconuiaria chesterensis.
48 J. J. Galloway, 1949. Galloway location L40C, ravine beginning at railroad, 1,2 km northwest of Harrodsburg and mnning northwest 0.8 km to old Indiana Highway 7, SEVa, SWV4 Sec. 20, T7N RIW, Monroe County, Indiana, Clear Creek 7.5' Quadrangle. Lower part of Harrodsburg Formation; Meramecian Series. Conularia cf. C. subcarbonaria.
49 J. J, Galloway, 1949. Galloway location 1.70A, old quarry, 4 km northwest of Dolan, Monroe County, Indiana. Site is probably an abandoned Quarry 2,4 km northwest of Dolan shown on Modesto 7.5' Quadrangle topographic map; WV2, SW*/4, NWV4 Sec. 34, T ION RIW, approximately 0.4 km east of old Indiana High- way 37 (A.S. Horowitz, written communication, 1984). Lower part of Harrodsburg Formation; Meramecian Series. Paraconuiaria chesterensis.
50 D, E. Hattin, 1965. Roadcut on State Highway 46, east of Gnaw Bone, approxi-
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mately NV2, NV2 Sec.25, T9N, R4E, Brown County, Indiana, Nashville 7.5' Quad- rangle. Carwood Formation of Borden Group; Chesterian Series, Paraconularia cf. P. byblis, P. chesterensis.
5 1 Anonymous. Spergen Hill, Indiana. Spergen Limestone; Meramecian Series. Con- ularia subcarbonaria, Paraconularia chesterensis.
52 Anonymous. Spergen Hill, Indiana? Spergen Hill Limestone?; Meramecian Series? Paraconularia chesterensis.
53 Anonymous. Hendricks County, Indiana. Formation unknown; probably Chesterian Series. Paraconularia chesterensis.
54 W. F. E. Gurley. Evansville, Indiana. “Chester Group”; Chesterian Series. Para- conularia chesterensis.
55 A. S. Brockley and T. G. Perry, 1954. Debris from Mulzer Brothers Quarry, SWy4, NE‘/4 Sec. 3, T2S, R2W, Crawford County, Indiana, Taswell 7.5' Quadrangle. “Glen Dean Limestone” (=Lower Tar Springs Formation); Chesterian Series. Paraconu- laria chesterensis.
56 A. S. Horowitz. Spoil heaps from Mulzer Brother Quarry, north and south of county road, approximately 1.5 km south of junction of Indiana Highways 145 and 164 and approximately 4 km north of Eckerty and .4 km east of Indiana Highway 145, SW'/4, SE‘/4 Sec. 10, T2S, R2W, Crawford County, Indiana, Taswell 7.5' Quadrangle. Glen Dean Limestone; Chesterian Series. Paraconularia chesterensis.
57 A. S. Horowitz et al., 1956-1982. Railroad cuts on west side of Baltimore and Ohio Railroad spur leading to National Gypsum Company quarry, near Shoals, SW*/*, NE‘/4 Sec. 28, T3N R3W, Martin County, Indiana, Huron 7.5' Quadrangle. “Gol- conda Formation” (=Indian Springs Member of the Big Clifty Formation); Ches- terian Series. Paraconularia chesterensis.
58 A. C. Brookley and T. G. Perry, 1954. Debris from abandoned Lutgring Quarry, 9.6 km east of Branch ville, NW'A, SW*/4 Sec. 18, TIS, RIW, Perry County, Indiana, Branchville 7.5' Quadrangle. “Glen Dean Limestone” (=Lower Tar Springs For- mation); Chesterian Series, Paraconularia chesterensis.
59 Haines. Washington County, Indiana. Formation unknown; probably Chesterian Series. Paraconularia chesterensis.
60 J. Below, 1965. Dam site 4. 8-6.4 km north off Route 46, 4.8 km east of Gnaw Bone, Indiana. Borden Group; Osagian Series? Paraconularia byblis.
Iowa
61 W. F. Gurley. Le Grand, Iowa. Formation unknown; Kinderhookian Series. Con- ularia subcarbonaria, Paraconularia blairi.
62 C. A. White? Burlington, Iowa. English River Sandstone of the Kinderhook Group; Kinderhookian Series. Paraconularia byblis.
63 C. A. White? Burlington, Iowa. “Upper Division of the Burlington Limestone”; probably Osagian Series. Paraconularia subulata.
64 Anonymous. Iowa City, Iowa. Cedar Valley Limestone; Chautauquan Series. Con- ularia subcarbonaria.
65 Anonymous. Probably Iowa City area, Iowa. Probably Cedar Valley Limestone; Chautauquan Series. Conularia subcarbonaria.
66 Anonymous. Burlington, Iowa. Burlington Limestone; Osagean Series. Paraconu- laria byblis.
67 Fenton. Southwest of Waverly, Iowa. Cedar Valley Limestone, lower part; Osagean Series. Conularia subcarbonaria.
68 Anonymous. Burlington, Iowa. Burlington Limestone; Osagean Series. Conularia subcarbonaria.
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69 Anonymous. Keokuk, Iowa. Keokuk Formation; Osagean Series. Paraconularia
missouriensis.
70 S. Weller? Keokuk, Iowa. Keokuk Limestone, “bed 1 1”; Osagean Series. Paracon- ularia chesterensis.
Kentucky
71 C. E. Mason, 1984. Outcrops along Interstate 64, 8.2 km east of the junction with Kentucky Route 32, near Morehead, Rowan County, Kentucky. Float from lower few meters of Nancy Member of Borden Formation; Osagean Series. Conularia multicostata, Paraconularia byblis.
72 C. E. Mason, T. M. Stanley, and L. E. Babcock, 1984. Nancy Member of Borden Formation; phosphate pebble bed about 1 m above top of “dysaerobic fauna”; Osagean Series. Spillway to Cave Run Lake, Daniel Boone National Forest, Bath County, Kentucky, Salt Lake 7.5' Quadrangle, Conularia multicostata, Paraconu- laria byblis, P. subulata.
IZ Anonymous. Natural Bridge, Kentucky. Borden Formation, probably Nancy Mem- ber {fide F. R. Ettensohn, written communication, 1985); Osagean Series. Conularia multicostata.
74 Anonymous. Marion County, Kentucky. Borden Formation?; Osagean Series? Par- aconularia byblis.
75 Anonymous. Lebanon, Kentucky. “Waverly Formation” (=Borden Group?); prob- ably Osagean Series. Paraconularia subulata.
76 Anonymous. About 2.5 km east of Lebanon, Kentucky. New Providence Formation; Osagean Series. Conularia subcarbonaria, Paraconularia byblis.
11 T. W. Kammer. Kammer location 10885, St. Francis, Kentucky, Nancy Member of the Borden Formation; Osagean Series. Paraconularia byblis.
78 T. W. Kammer. Kenwood Hill, Louisville, Kentucky. New Providence Shale Mem- ber of the Borden Formation; Osagean Series. Paraconularia byblis.
79 Anonymous, Kentucky? Waverly Group equivalent?; Osagean Series? Conularia multicostata.
80 U. P. James. Boyle or Marion County, Kentucky. “Waverly Group” (=Borden Group?); probably Osagean Series. Conularia multicostata.
8 1 Anonymous. Knob just south of Louisville, Kentucky. New Providence Formation; ?Osagean Series. Paraconularia byblis.
82 G. Robb, pre-1923. Marion County, Kentucky. “Keokuk Formation, Knob Shale” (=Borden Formation?); Osagean Series. Paraconularia subulata.
83 Anonymous. Elizabethtown, Kentucky. St. Louis Limestone; Meramecian Series. Paraconularia missouriensis.
84 A. S. Horowitz. Pond north of Kentucky Highway 1576, about 2 miles east of Morrilla, Jackson County, Kentucky. Pennington Formation; Chesterian Series. Paraconularia chesterensis.
85 A. S. Horowitz. Near Colesburg, Hardin County, Kentucky. Somerset Shale Member of Salem Limestone; Meramecian Series. Paraconularia chesterensis.
86 S. Weller, 1920. 5.2 km south of lola, Kentucky (GK 12). Glen Dean Limestone; Chesterian Series. Paraconularia chesterensis.
87 Anonymous. About 1 .5 km west of Mongomery Switch, Caldwell County, Kentucky. Claystone bed of upper Chester Formation; Chesterian Series. Paraconularia ches- terensis.
88 A. S. Horowitz, 1966, etc. Pond above road leading to Pearson Farm glade (road not on topographic map), near junction of road with Kentucky Highway 1576, approximately 3.0 km east ofjunction of Kentucky Highway 1 576 and U.S. Highway
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421, approximately 3 km east of Morrill, Jackson County, Kentucky, Big Hill 7.5' Quadrangle. Lower part of Pennington Formation, just above top of Bangor Lime- stone; Chesterian Series. Paraconularia chesterensis.
89 A. S. Horowitz, 1969. Roy Norton Farm, glades on north, west and south slopes of tributaries on north side of Broad Run, SO-lOO m south of Kentucky Highway 434, approximately 2.6 km west-southwest of Colesburg, Hardin County, Kentucky, Colesburg 7.5' Quadrangle. Somerset Shale Member of the Salem Limestone; Ches- terian Series. Paraconularia chesterensis.
Maine
90 Anonymous. Presque Isle stream. Chapman Plantation, Maine. Chapman Sand- stone; series unknown, Devonian System. Conularia cf C undulata.
Maryland
91 F. M. Swartz. West Maryland Railroad tracks, Corrigansville, Maryland. Upper part of the Shriver Chert; Ulsterian Series. Conularia undulata.
Michigan
92 Anonymous. Alpena, Michigan. Alpena Limestone; Brian Series. Paraconularia alpenensis.
93 E. W. Hard. U.S. Gypsum Company Quarry, Sec. 27, T27N, R7E, near Alabaster, Michigan. Michigan Formation, gray gypsiferous limestone bed 1.3 m below 5.3 m thick bed of mottled white gypsum; Osagean Series. Paraconularia yochelsoni.
Missouri
94 Sampson; Faber. Sedalia, Missouri, Chouteau Limestone; Kinderhookian Series. Paraconularia blairi, P. missouriensis.
95 J. S. Williams, 1930? Easley, Missouri. Chouteau Limestone; Kinderhookian Series. Paraconularia blairi.
96 E. B. Branson. Providence, Missouri. Chouteau Limestone; Kinderhookian Series. Paraconularia blairi.
97 E. B. Branson, 1930. Providence, Missouri. Couteau Limestone; Kinderhookian Series. Paraconularia blairi.
98 Anonymous. Pettis County, Missouri. Chouteau Limestone; Kinderhookian Series. Paraconularia blairi.
99 E. B. Branson. Browns, Missouri. Chouteau Limestone; Kinderhookian Series. No conulariids identified; ?priapulid worm, ^‘‘Conularia"'’ tenuicostata.
100 W. F. E. Gurley. Boonville, Missouri. Keokuk Limestone; Osagean Series. Para- conularia missouriensis.
101 Anonymous. Carthage, Missouri. Formation unknown; Meramecian Series. Con- ularia subcarbonaria.
102 Van Home. Foot of La Beaume Street, St. Louis, Missouri. St. Louis Limestone; Meramecian Series. Paraconularia blairi, P. chesterensis.
103 Anonymous, Kansas City, Missouri. “Coal Measures”; probably Osagean or Mer- amecian Series. Paraconularia missouriensis.
1 04 Anonymous. Carthage, Missouri. Keokuk Limestone; Osagean-Meramecian Series, Paraconularia chesterensis.
105 Anonymous. Little Rock, St. Genevieve County, Missouri. St. Louis Limestone;
Meramecian Series. Paraconularia blairi.
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Montana
106 R. Lund et al., 1978-1984. Potter’s Creek Dome, approximately 50 km southeast of Lewistown, Fergus County, Montana. Bear Gulch Limestone of Heath Formation; Chesterian Series. Paraconularia subulata.
107 W, H, Easton? Delpine, Meagher County, Montana. Cameron Creek Shale of the Big Snowy Group; Chesterian Series. Paraconularia subulata.
Nevada
108 Merriam. Simpson Park Range, Nevada. Rabbit Hill Limestone; Ulsterian Series. Conularia sp.
109 A. J. Boucot, 1984. West face of Red Hill, Eureka County, Nevada. “Fish bed” of the Denay Limestone; Senecan Series. Conularia recurvatus.
110 C. D. Walcott. Eureka District, Nevada. Formation and series unknown; probably Chesterian Series. Paraconularia chesterensis.
New Jersey
111 D. Parris and K. Cruikshank, 1980. On Weider Road, near County Road 521, Montague Township, Sussex County, New Jersey. Esopus Formation; Ulsterian Series. Reticulaconularia sussexensis.
1 12 H. Herpers. Montague, Sussex County, New Jersey. Esopus Formation; Ulsterian Series. Reticulaconularia sussexensis.
113 H. Herpers, 1948. Millville, Montague Township, Sussex County, New Jersey. Esopus Formation; Ulsterian Series. Reticulaconularia sussexensis.
1 14 D. Parris, K. Cruikshank et al., 1 984. 1.7 km southwest of Wallpack Centre, roadcut across from Batteli’s Campground, Wallpack Township, Sussex County, New Jersey. Port Ewen Formation; Ulsterian Series. Conularia pyramidalis.
New York
115 R. M. Fulle. First Esopus outcrop south-southeast of Hurley, on left fork of road, about 1 .6 km from Hurley, Ulster County, New York. Esopus Formation; Ulsterian
Series. Conularia ulsterensis.
116 Anonymous. Schoharie, New York. New Scotland Limestone; Ulsterian Series. Conularia pyramidalis.
1 1 7 Anonymous. Clarkesville, Schoharie County, New York. “Lower Helderberg Group” (=New Scotland Limestone); Ulsterian Series. Conularia pyramidalis.
1 18 Anonymous. Probably Clarksville area, Schoharie County, New York. Helderberg Group?, possibly New Scotland Limestone; Ulsterian Series. Conularia pyramidalis.
1 1 9 Anonymous. NYSM locality 2969, near Helderberg, New York. Manlius Limestone; Ulsterian Series. Conularia pyramidalis.
120 Anonymous. Countryman’s Hill, New Salem, New York. Coeymans Limestone or New Scotland Limestone; Ulsterian Series. Conularia pyramidalis.
121 Anonymous. Knox, Albany County, New York. Oriskany Sandstone; Ulsterian Series. Conularia desiderata.
122 Anonymous. Schoharie, New York. Oriskany Sandstone; Ulsterian Series. Conu- laria pyramidalis.
123 F. M. Swartz. Clarksville, New York. “Lower Helderberg Group,” probably New Scotland Limestone; Ulsterian Series. Conularia pyramidalis.
124 Anonymous. NYSM location 2, about 2 km south of Bridgewater, New York. Marcellus Shale; Erian Series, Conularia desiderata.
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125 Anonymous. Bridgewater, New York. Marcellus Shale; Erian Series. Conularia desiderata.
1 26. Anonymous. Probably from near Bridgewater, New York. Probably from Marcellus Shale; Erian Series. Conularia desiderata.
127 Anonymous. Vicinity of Hamilton, New York. Lower part of Hamilton Group; Erian Series. Conularia desiderata.
128 Anonymous. Morrisville, New York. Hamilton Group, probably Solsville Member of the Marcellus Formation; Erian Series. Conularia undulata.
129 Anonymous. About 1.2 km northwest of Solsville, Madison County, New York. Hamilton Group; Erian Series. Conularia undulata.
130 P. Zell, 1982, Swamp Road quarry, near Morrisville, New York, Morrisville IS Quadrangle. Marcellus Formation; Erian Series. Conularia pyramidalis.
131 L. E. Babcock, J. T. Hannibal, and R. M. Feldmann, 1984. Borrow pit on east side of Swamp Road, 4,2 km north of Morrisville, New York, Morrisville 7.5' Quad- rangle. Solsville Member of the Marcellus Formation; Erian Series. Conularia pyr- amidalis.
132 Anonymous. Schoharie County, New York. Hamilton Group, possibly Schoharie Formation; Erian Series. Conularia desiderata.
133 M. Kopf. 4 km east of Alexander, New York. Centerfield Limestone Member, Ludlowville Formation; Erian Series. Conularia desiderata.
134 1. H. Reimann. Spring Creek, Alden, New York. Ledyard Shale Member of Lud- lowville Formation; Erian Series. Conularia desiderata.
135 G. J. Kloc, 1983. Lake Erie shore, south of the Wanakah Water Plant, Wanakah, Erie County, New York, N42°44'50" W78°54'13". Nautilus Bed of the Wanakah Shale Member of the Ludlowville Formation; Erian Series. Conularia desiderata.
136 Anonymous. Genesee Valley, New York. Hamilton Group; Erian Series. Conularia undulata.
1 37 Anonymous. Norton’s Landing, Cayuga Lake, New York. Hamilton Group, possibly King Ferry Shale Member of the Ludlowville Formation; Erian Series. Conularia undulata.
138 Anonymous. NYSM location 428, Shurger’s Glen, near Norton’s Landing, Cayuga Lake, New York. Hamilton Group; Erian Series. Conularia undulata.
139 Anonymous. NYSM location 437, Shurger’s Glen, near Norton’s Landing, Cayuga Lake, New York. Hamilton Group; Erian Series. Conularia sp.
140 G. C. Baird, ca. 1980; L. E. Babcock, 1983. Banks and bed of Bamum Creek, below high falls 0.4 km west (upstream) from New York Route 89 overpass, Sheldrake Quadrangle, New York. Bamum Creek Bed of the King Ferry Shale Member of the Ludlowville Formation, approximately 1 0 m above top of the Pleurodictyum zone; Erian Series. Conularia undulata.
141 G. C. Baird, ca. 1980. Bed of Sheldrake Creek, below high falls 0.48 km northeast (downstream) from New York 89 overpass, Sheldrake Quadrangle, Seneca County, New York. 2 m below Bamum Creek Bed, in the King Ferry Shale Member of Ludlowville Formation; Erian Series. Conularia undulata.
142 G. A. Cooper, ca. 1930. Hamilton, New York, Cooper location 8Qa. Upper part of Pompey Formation; Erian Series. Conularia desiderata.
143 Anonymous. NYSM location 558, Norwich, Chenango County, New York. Ham- ilton Group; Erian Series. Conularia undulata.
144 Anonymous. NYSM location 611, Schoharie County, New York. Hamilton Group; Erian Series. Conularia desiderata.
1 45 Anonymous. Near Cazenovia, New York. Hamilton Group; Erian Series. Conularia undulata.
1 46 Anonymous. Cazenovia, New York. Hamilton Group, possibly Moscow Formation; Erian Series. Conularia desiderata, C. undulata.
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469
147 P, Zell, 1983. Thompson Hill Road quarry, near Earlville, New York, Earlville 1.5' Quadrangle. Moscow Formation; Erian Series. Conularia undulata.
148 R. M. Linsley; L. E. Babcock, J. T. Hannibal, and R. M. Feldmann, 1984. “Earlville trilobite quarry,” off Morris Road, near Morrisville, New York, Earlville 7.5' Quad- rangle. Upper part of Moscow Formation; Erian Series. Conularia undulata.
149 Anonymous. Folsomdale, New York. Rhinestreet Shale; Senecan Series. Conularia congregatal
150 Anonymous. Ithaca, New York. Ithaca Formation; Senecan Series. Conularia con- gregata.
151 Anonymous. NYSM location 347, Ithaca, New York. “Chemung Group”; Senecan
Series. Conularia sp.
1 52 Anonymous. NYSM location 390, west side of Cayuga Lake inlet, New York. Ithaca Shale; Senecan Series. Conularia congregata.
153 Anonymous. NYSM location 392, 1.5 km southeast of Ithaca, New York. Ithaca Group; Senecan Series. Conularia congregata.
1 54 J. W. Hall and G. B. Simpson, 1 870. NYSM location 425, Ithaca, New York. Ithaca Shale, lower part of formation; Senecan Series. Conularia congregata.
155 Anonymous. NYSM location 514, Catskill Turnpike, 3-4.5 km east of Stamford, Delaware County, New York. Ithaca Group; Senecan Series. Conularia congregata.
156 J. W. Hall and C. Van Deloo, 1866. “Mr, Cornell’s Quarry,” 1.5 km northeast of Ithaca; also from Cemetary quarry and Cascadilla Creek, Ithaca, New York. Ithaca Shale; Senecan Series. Conularia congregata.
157 D. D. Luther, 1900. NYSM locality 2439, West Hill, near Naples, New York. “Naples Group”; Senecan Series. Conularia congregata.
1 58 Anonymous. South Hill, Ithaca, New York. Ithaca Shale; Senecan Series. Conularia congregata.
1 59 P. Zell, 1 982-1 983. Collins Hill Road quarry, near Sherburne, New York, Sherburne 7.5' Quadrangle. Hamilton Group; formation unknown; probably Senecan Series. Conularia congregata.
160 L. E. Babcock, E. L. Yochelson, and W. T. Kirchgasser, 1982. Big Sister Creek, Angola, Erie County, New York. Float in Angola Shale; Chautauquan Series. Con- ularia cf. C. congregata.
161 E. B. Hall. E. B. Hall locality I, Wellsville, New York. Wellsville Formation; Chau- tauquan Series. Paraconularia wellsvillia.
162 E. B. Hall. E. B. Hall locality XVIII, Almond, New York. Wellsville Formation; Chautauquan Series. Paraconularia wellsvillia.
Nova Scotia
1 63 Anonymous. Cape Breton, Nova Scotia. Lower Windsor Group; Osagian-Chesterian
Series. Paraconularia planicostata.
1 64 W. Dawson. Irish Cove, Cape Breton, Nova Scotia. “Lower Carboniferous” lime- stone, probably Windsor Group; Osagian-Chesterian Series. Paraconularia plani- costata.
165 Anonymous. Windsor, Nova Scotia. Lower Windsor Group; Osagian-Chesterian Series. Paraconularia planicostata.
166 D. G. Kelley, 1954. GSC location 24841, about 100 m east of comer of Route 5 and Buckwheat Road, Nyonza, Cape Breton Island, Nova Scotia. Lower Windsor Group; Osagian-Chesterian Series. Paraconularia planicostata.
167 D. G. Kelley, 1954. GSC location 24844, limestone at bridge on Lewis Mountain Road, 0.8 km from Route 19, Cape Breton Island, Nova Scotia. Lower? part of Windsor Group; Osagian-Chesterian Series. Paraconularia planicostata.
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Annals of Carnegie Museum
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168 Anonymous. Nova Scotia. Lower Windsor Group; Osagian-Chesterian Series. Par- aconularia planicostata.
169 Anonymous. Cape Breton, Nova Scotia. Windsor Group; Osagian-Chesterian Se- ries. Paraconularia planicostata.
170 M. J. Copeland, 1962. Shore of Bros d’Or Lake, Irish Cove, Cape Breton, Nova Scotia. Windsor Group; Osagian-Chesterian Series. Paraconularia planicostata.
171 Anonymous. Brookfield, Colchester County, Nova Scotia. Lower Windsor Group; Osagian-Chesterian Series. Paraconularia planicostata.
172 Anonymous. Harts County, Nova Scotia. Basal Windsor Group; Osagian-Cheste- rian Series. Paraconularia planicostata.
173 Anonymous. Maxner Point, Nova Scotia. Probably lower part of Windsor For- mation; Osagian-Chesterian Series. Paraconularia planicostata.
Ohio
1 74 Anonymous. Delaware, Ohio. “Comiferous Group,” probably Delaware Limestone; Ulsterian Series. Conularia elegantula.
175 G. Meszaros, pre-1982. Rathbone, Ohio. Columbus Limestone; Ulsterian Series. Conularia elegantula.
176 Hyatt Brothers, Dublin, Franklin County, Ohio. Columbus or Delaware Limestone; Ulsterian Series. Conularia elegantula.
177 Anonymous. Quarry 4 km southeast of Sylvania, Ohio. Silica Shale; Erian Series.
Conularia sp.
178 G, Meszaros, pre- 1 982. Leroy, Ohio. Chagrin Shale; Chautauquan Series, Conularia multicostata, Paraconularia chagrinensis.
179 E. Roeser, 1978. Float along Mill Creek, at and near Camp Koinonia, Lake and Ashtabula counties, Ohio, north and south of Ross Road bridge. Chagrin Shale; Chautauquan Series. Paraconularia chagrinensis.
180 D. Strock, 1985? Along Mill Creek, from Hidden Valley Park to the church camp. Lake County, Ohio. Chagrin Shale; Chautauquan Series. Paraconularia chagrinen- sis.
181 C. Talerico, 1984; L. E. Babcock et al., 1984. Float along Mill Creek, between Ross Road bridge and small dam upstream of Ross Road, Ashtabula County, New York. Chagrin Shale; Chautauquan Series. Paraconularia chagrinensis.
182 T. Stanley, 1984. Mill Creek, within 165 m downstream (north) of Ross Road bridge. Lake and Ashtabula counties, Ohio. Chagrin Shale; Chautauquan Series. Paraconularia chagrinensis.
183 M. E. Williams, 1981; S. McKenzie, pre-1982. Float along Mill Creek, Ashtabula County, Ohio. Chagrin Shale; Chautauquan Series. Paraconularia chagrinensis.
184 M. E, Williams, 1981; J. Hannibal etal., 1985. Stebbins Gulch, Holden Arboretum, Geauga County, Ohio, Contact between the Chagrin Shale and the Cleveland Shale Member of the Ohio Formation; Chautauquan Series. Paraconularia chagrinensis.
185 A. J. Weiss, 1984, Landfill on north side of Ohio Route 82, approximately 1.2 km west of 1-77 interchange, Broadview Heights, Ohio. Cuyahoga Formation, Meadville Shale Member; Kinderhookian Series. Paraconularia byblis, P. subulata.
186 J. Hall? Alexander, Licking County, Ohio. “Berea Shale” (=Sunbury Shale Sub- member, Orangeville Member, Cuyahoga Formation); Kinderhookian Series, Par- aconularia subulata.
187 Herrick? Alexander, Licking County, Ohio. “Berea Shale” (=Sunbury Submember of the Orangeville Member of the Cuyahoga Formation?); Kinderhookian Series. Paraconularia subulata.
188 L. E. Babcock, 1984. Sunbury Shale Submember of the Orangeville Member of the Cuyahoga Formation; 1-3 cm thick silty zone with abundant pyrite at Sunbury-
1986 Babcock and Feldmann—P/I^ coiV[/iv4i?L4 and Reticulaconularia
471
Berea Sandstone contact; Kinderhookian Series. Quarry Rock picnic area, north of Chagrin River, South Chagrin Reservation, east of Solon Road, Bentleyville, Cuy- ahoga County, Ohio. Paraconularia subulata.
189 M. Ciccarone, 1984. Sunbury Shale Submember of the Orangeville Member of the Cuyahoga Formation; Kinderhookian Series. Quarry Rock picnic area, north of Chagrin River, South Chagrin Reservation, east of Solon Road, Bentleyville, Cuy- ahoga County, Ohio. Paraconularia subulata.
190 L. E. Babcock, 1984. Sunbury Shale Submember of the Orangeville Member of the Cuyahoga Formation; approximately 1 m above the top of the Berea Sandstone; Kinderhookian Series. Quarry Rock picnic area, north of Chagrin River, South Chagrin Reservation, east of Solon Road, Bentleyville, Cuyahoga County, Ohio. Paraconularia subulata.
191 G. Meszaros, pre-1982. Weymouth, Ohio. Meadville Shale Member of Cuyahoga Formation; Kinderhookian Series. Paraconularia subulata.
192 J. Burke, W. J. Hlavin et al., 1967. North Branch of Rocky River near bridge at junction of Bagdad and Hood Roads, Bagdad, Ohio. Meadville Shale Member of Cuyahoga Formation; Kinderhookian Series. Paraconularia subulata.
193 Anonymous. Probably northeast Ohio. Probably Meadville Member of the Cuy- ahoga Formation; Kinderhookian Series? Conularia multicostata.
1 94 Anonymous. Voorhes Cemetary outcrop, west of Lodi, Ohio. Meadville Shale Mem- ber of Cuyahoga Formation; Kinderhookian Series. Paraconularia subulata.
195 R. W. Scott. Lodi, Medina County, Ohio. Cuyahoga Formation, probably Meadville Member; Kinderhookian Series. Paraconularia byblis.
196 G. Meszaros, pre-1982. Lodi, Ohio. Meadville Shale Member of the Cuyahoga Formation; Kinderhookian Series. Conularia multicostata, Paraconularia subulata.
197 R. Fisher. Creeks in and near Lodi, Medina County, Ohio. Meadville Member of the Cuyahoga Formation; Kinderhookian Series. Conularia multicostata, Paracon- ularia subulata.
198 R. Segedi et al., 1974. Streambed olf Pawnee Road, about 200 m south of U.S. Route 224, Lodi, Medina County, Ohio. Meadville Shale Member of Cuyahoga Formation; Kinderhookian Series. Conularia multicostata, Paraconularia subulata.
199 R. Segedi. About 50 m east of bridge on Pawnee Road, just south of U.S. Route 224, Lodi, Medina County, Ohio. Float and in situ specimens from the Meadville Member of the Cuyahoga Formation; Kinderhookian Series. Conularia multicos- tata, Paraconularia subulata.
200 L. E. Babcock, 1985. West fork of East Branch of Black River, south of Route 224, near intersection with Pawnee Road, Homer Township, about 3 km west of center of Lodi, Medina County, Ohio, Lodi 7.5' Quadrangle. Meadville Shale Member of Cuyahoga Formation; collected in situ in lowermost bed of siderite concretions upstream of Pawnee Road bridge; Kinderhookian Series. Conularia multicostata, Paraconularia subulata.
20 1 A. H. Worthen. Richfield, Ohio. “Kinderhook Formation” (=Cuyahoga Formation, probably Meadville Member); Kinderhookian Series. Paraconularia subulata.
202 Anonymous. NYSM location 110, Richfield, Summit County, Ohio. “Waverly Group” (probably Meadville Member of the Cuyahoga Formation); Kinderhookian Series. Paraconularia subulata.
203 Anonymous. Richfield, Ohio. Waverly Group, probably Meadville Member of the Cuyahoga Formation; Kinderhookian Series. Paraconularia subulata.
204 L. E. Babcock, 1984. Float in Meadville Member of the Cuyahoga Formation; Kinderhookian Series. Tributary to the Cuyahoga River, Furnace Run Metro Park, west off Route 2 1 , about 1.5 km south of Summit County-Cuyahoga County bound- ary, Summit County, Ohio. Paraconularia cf P. byblis.
205 Anonymous. Richfield, Ohio. “Waverly Group” (=Meadville Member of the Cuy-
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Annals of Carnegie Museum
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ahoga Formation); Kinderhookian or Osagean Series. Conularia multicostata, Para- conularia subulata.
206 A. Winchell. Cuyahoga River gorge, Cuyahoga Falls, Summit County, Ohio. “Near top of the Waverly Group, water limestone below conglomerate” (=Cuyahoga For- mation, possibly Meadville Member); Kinderhookian-Osagean Series. Pamconu- laria subulata.
207 J. Weiss, 1961. Gravel in Akron, Ohio area? Possibly Cuyahoga Formation; Kin- derhookian or Osagian Series. Paraconularia subulata.
208 Anonymous. “Rocky River bed,” probably Cuyahoga Formation; Kinderhookian or Osagean Series. Medina, Ohio. Paraconularia subulata.
209 Anonymous. Bagdad, Ohio. Probably Cuyahoga Formation; Kinderhookian or Osa- gean Series. Conularia multicostata.
210 C. L. Herrick. Near Lyon Falls, Richland County, Ohio. 18-24 m above “Con- glomerate n”: Osagean Series. Paraconularia byblis.
211 W. P. Cooper, 1890; Anonymous. Portsmouth, Ohio. Waverly Group, probably the Cuyahoga Formation; Osagean Series. Paraconularia missouriensis.
2 1 2 Anonymous. Wooster, Wayne County, Ohio. “Near the top of the Waverly Group” (=Wooster Member of the Cuyahoga Formation); Osagean Series. Paraconularia subulata.
213 Anonymous. Wooster, Wayne County, Ohio. Near top of Waverly Group, possibly the Wooster Member of Cuyahoga Formation; Osagean Series. Paraconularia su- bulata.
214 G. Meszaros, pre- 1 982. Wooster, Ohio. Cuyahoga Formation, Wooster Shale Mem- ber; Osagean Series. Conularia multicostata, Paraconularia subulata.
215 H. E. Wilson. 7.2 km south of Loudonville, Ashland County, Ohio. Probably Woos- ter Member, Cuyahoga Formation; Osagean Series. Paraconularia subulata.
216 F. Plutte, 1 964. About 4.5 km south of Loudonville, Ashland County, Ohio. Wooster Member of the Cuyahoga Formation; Osagean Series. Paraconularia subulata.
217 L. E. Babcock, 1984. East facing borrow pit on west side of Route 3, 0.9 km north of junction with Route 97, just south of Loudonville, Ashland County, Ohio, Greer 7.5' Quadrangle. Wooster Member of the Cuyahoga Formation; Osagean Series. Conularia multicostata, Paraconularia subulata.
218 J. Hall? Water works in Newark, Ohio. Waverly Group, “base of Division IIP’; probably Cuyahoga Formation; probably Osagean Series. Conularia multicostata.
219 G. Meszaros, pre- 1982. Rushville, Ohio. Allensville Member of Logan Formation; Osagian Series. Paraconularia byblis.
220 Bowsher, Savage, and Allen, 1952, Approximately 300 m of elevation below Old Maid’s Kitchen, in gully about 110 m west of Ohio Edison Dam on north side of Cuyahoga Gorge, Akron, Summit County, Ohio. Float in Meadville Shale Member of the Cuyahoga Formation; Kinderhookian Series. Paraconularia subulata.
22 1 Anonymous. Ohio. Possibly Cuyahoga Formation; Mississippian, possibly Osagean. Conularia multicostata, Paraconularia subulata.
Ill Anonymous. Ohio. “Lower Waverly Group,” Cuyahoga Formation; Osagean Series. Paraconularia subulata.
223 Stout and Girty, 1 897. Dixon’s Mill, on the Little Scioto River, about 5 km northeast of Sciotoville, Scioto County, Ohio. “Waverly Group,” probably Wooster Member of Cuyahoga Formation; Osagian Series. Conularia multicostata, Paraconularia byblis, P. missouriensis.
224 Carman, Stout, and Carney. Sciotoville, Ohio. Upper part of Cuyahoga Formation, 16.5-23 m below base of the Logan Formation; Osagian Series. Paraconularia byblisl, P. missouriensis.
225 F. B. Meek; Anonymous. Sciotoville, Ohio. “Waverly Group,” probably Cuyahoga Formation; Osagean Series. Conularia multicostata.
226 E. B. Andrews, 1869. Sciotoville, Ohio. Uppermost Cuyahoga Formation, Black
1986 Babcock and Feldmann— /’A?L4C6>NCZL4i?/^ and Reticulaconularia
473
Hand Member or lowermost Logan Formation, Byer Member; Osagean Series. Paraconularia missouriensis.
227 G. Meszaros, pre-1982. Sciotoville, Ohio. Portsmouth Shale; Osagean Series. Con- ularia muliicostata, Paraconularia missouriensis.
228 Cooper. James Hall’s location 385, Moot’s Run, Licking County, Ohio. Cuyahoga Formation, probably Wooster Member; Osagean Series. Conularia multicostata.
Oklahoma
229 G. A. Cooper et al., 1952. NE'A Sec. 7 T22N R20E, 5.3 km south of Adair, Mayes County, Oklahoma. Fayetteville Formation; Chesterian Series. Paraconularia okla-
homaensis.
Ontario
230 C. S. 9.6 km west of Cayuga, Ontario. Upper part of Oriskany Sandstone; Ulsterian Series. Conularia undulata.
Pennsylvania
23 1 F. M. Swartz. Near Curtin, Pennsylvania. Shriver Chert; Ulsterian Series. Conularia ulsterensis.
232 F. M. Swartz. Intersection of Delaware and New York State Railroad, 0.8 km west of mill of Mimsruk Paper Company Experimental Mills, near Curtin, Pennsylvania. Shriver Chert; 1.05 m below Oriskany Shale; Ulsterian Series. Conularia cf C.
desiderata.
233 F. M. Swartz, 1937. Float on roadcut on road leading through gap east of War- fordsburg, Pennsylvania. Shriver Formation?; 0.3”1 .7 m above conglomeratic sand- stone at middle of Shriver-like beds; Ulsterian Series. Conularia ulsterensis.
234 F. M. Swartz, 1937. Roadcut along road leading through gap east of Waifordsburg, Pennsylvania. 0.3- 1.5 m above conglomeratic sandstone at middle of Shriver-like beds; Ulsterian Series. Conularia ulsterensis.
235 F. M. Swartz. Road leading north from Schellsburg, Bedford County, Pennsylvania. Onondaga Shale; Erian Series. Conularia cf C desiderata.
236 S. Albright, 1 98 1 ; B. White, 1 983. Large roadcut on north side of Johnny Bee Road, about 0.2 km north of intersection with road to Dingmans Falls, Delaware Town- ship, Pike County, Pennsylvania. Mahantango Formation, approximately Center- field biostrome level; Erian Series. Conularia desiderata, C. ulsterensis, C. undulata.
237 D. Parris, 1982. Large roadcut on north side of Johnny Bee Road, about 0.2 km north of intersection with road to Dingmans Falls, Delaware Township, Pike Coun- ty, Pennsylvania. Manhantango Formation; Erian Series. Conularia undulata.
238 L. Klensch and J. Valenti, 1981. Roadcut across from Bushkill Country Store, 2 km from U.S. Route 209, Lehman Township, Pike County, Pennsylvania. Mahan- tango Formation; Erian Series. Conularia undulata.
239 L. Decina, 1983. Roadcut on north side of Pennsylvania Route 895, approximately 0.8 km west of Auburn, Schuylkill County, Pennsylvania. Mahantango Formation; Erian Series. Conularia desiderata.
240 Anonymous. Huntingdon, Huntingdon County, Pennsylvania. Frame Shale Mem- ber of the Mahantango Formation; Erian Series. No conulariids collected; Hy- olithes sp.
Quebec
24 1 R. B., 1 862; Anonymous. Grande Greve, Gaspe, Quebec. Grand Greve Limestone; Ulsterian Series. Conularia cf C desiderata, C. cf C. undulata.
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Annals of Carnegie Museum
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242 Anonymous. High Falls, Dartmouth River, Gaspe Peninsula. Grande Greve Lime- stone; Ulsterian Series. Reticulaconularia penouili.
243 Anonymous. Little Gaspe, Quebec. Grande Greve Limestone; Ulsterian Series. Conularia cf. C. desiderata.
244 Anonymous. Perce Rock, Gaspe, Quebec. Grande Greve Limestone; Ulsterian Se- ries. Conularia tuzoi.
245. Cape Barre, Quebec. “Cape Barre beds” (=Cape Bon Ami Formation); Ulsterian Series. Conularia cf. C. desiderata.
246 Anonymous. Gaspe Peninsula, Quebec. Float block of limestone, probably Grande Greve Limestone; Ulsterian Series. Reticulaconularia penouili.
247 J. W. Beede? Magdelen Islands, Quebec. Formation and series unknown; Missis- sippian Subsystem. Paraconularia sorrocula.
248 J. W. Beede? Cape le Tron, Grindstone Island, Magdelen Islands, Quebec. For- mation and series unknown; Mississippian Subsystem. Paraconularia planicostata.
South Dakota
249 L. E. Babcock, 1984. “Slagpile section,” overlooking bridge of Route 14A over Whitewood Creek, SW ‘A Sec. 13, T5N R3E, Deadwood, Lawrence County, South Dakota, Deadwood 7.5' Quadrangle. Englewood Formation, dolostone about 20 cm above top of shale-dolostone transitional zone. Paraconularia sp.
Tennessee
250 J. M. Safford. Hickman County, Tennessee. “Waverly Group”; Kinderhookian or Osagean Series. Paraconularia byblisl
251 A. S. Horowitz, 1966, etc. Roadcuts on both sides of Interstate Highway 40, 8.3 km west of junction of Interstate Highway 40 and U.S. Highway 70 at Monterey, Putnam County, Tennessee, Monterey 7.5' Quadrangle. Top of Pennington For- mation; “Kinkaid level,” zone of Pterocrinus tridecibrachiatus Gutschick, just below a quartz pebble conglomerate of the Pennsylvanian System; Chesterian Series. Par- aconularia chesterensis.
Utah
252 C. D. Walcott? Divide Bet, American Fork and Snake Creek, Wasatch Mountains, Utah, Kinderhookian or Osagean Series. Paraconularia chesterensis?
Wisconsin
253 E. E. Teller. Milwaukee, Wisconsin. Probably Milwaukee Formation; Erian Series. Conularia milwaukeensis.
254 E. E. Teller? Estabrook Park, Milwaukee, Wisconsin. Lindwurm Member of the Milwaukee Formation; Erian Series. Conularia milwaukeensis.
255 E. E. Teller. Milwaukee Cement Quarry, Berthelet, Wisconsin. Milwaukee For- mation; Erian Series. Conularia milwaukeenensis.
Appendix B — Measurements
The following values are measurements of selected type and other conulariid specimens considered representative of each taxon treated herein and in Part A. Species are listed alphabetically according to species as they are recognized herein. Measurements are listed in columns across each page. In cases where replicate measurements have been taken at varying distances from the hypothetical apex, they are listed vertically under the appro- priate columns. Terms are defined in the “Morphology” section. Abbreviations: L, length.
1986 Babcock and Feldmann --Paeaconularia and Reticulaconularia
475
cm; HL, hypothetical length, cm; AAMj, major apical angle, degrees; AAMn, minor apical angle, degrees; N, nodes/mm; RLAMj, ratio of right superior : left superior : abut- ting rods per ten on major face; RLAMn, ratio of right superior : left superior : abutting rods per ten on minor face; D, distance from hypothetical apex at apicad limit of mea- surement, applicable to values for R and RA; R, rods/cm; RA, rod angle, degrees; h, holotype; plh, plastoholotype; 1, lectotype; n, neotype; p, paratype; pi, paralectotype. Values given in parentheses are approximate; values followed by question marks are accurate, but it is not certain that they have been properly classified as pertaining to either the major or minor face.
Conularia congregata Hall
|
SPECIMEN NO. |
L |
HL |
AAMj AAMn N |
RLAMj |
RLAMn |
D |
R |
RA |
||
|
NYSM 3483 1 |
6.1 |
9.4 |
17 |
12 |
6 |
0:0:10 |
0:0:10 |
5.5 |
18 |
10 |
|
8.0 |
16 |
13 |
||||||||
|
NYSM 3483 pi |
6.7 |
10.7 |
17 |
14 |
7 |
0:0:10 |
0:0:10 |
5.5 |
17 |
10 |
|
8.5 |
21 |
9 |
||||||||
|
NYSM 3483 pi |
6.5 |
10.1 |
16 |
14 |
6 |
0:0:10 |
0:0:10 |
5.0 |
17 |
10 |
|
8.5 |
20 |
12 |
||||||||
|
NYSM 3483 pi |
6.3 |
9.9 |
17 |
13 |
6 |
0:0:10 |
0:0:10 |
8.0 |
16 |
10 |
|
Conularia delphiensis (Maroney and Orr) |
||||||||||
|
SPECIMEN NO. |
L |
HL |
AAMj AAMn N |
RLAMj |
RLAMn |
D |
R |
RA |
||
|
lUPC 14470-1 h |
2.0 |
(4.8) |
— |
— |
6 |
1:1:8 |
1:1:8 |
6.0 |
36 |
10 |
|
lUPC 14470-2 p |
2.1 |
(5.1) |
~ |
— |
1:0:9 |
0:1:9 |
4.5 |
36 |
11 |
|
|
lUPC 14470-4 p |
2.7 |
(3.4) |
15? |
- |
- |
0:0:10 |
0:0:10 |
1.6 |
42 |
|
|
lUPC 14470-6 p |
- |
- |
- |
7 |
- |
- |
“ |
- |
||
|
Conularia desiderata Hall |
||||||||||
|
SPECIMEN NO. |
L |
HL |
AAMj AAMn N |
RLAMj |
RLAMn |
D |
R |
RA |
||
|
AMNH 2697 h |
1.6 |
2.3 |
23 |
20? |
— |
0:10:0 |
0:10:0 |
3.5 |
27 |
15 |
|
NYSM 3487 |
5.8 |
6.4 |
27 |
22 |
3 |
1:8:2 |
1:9:0 |
5.0 |
17 |
12 |
|
7.0 |
14 |
7 |
||||||||
|
NYSM 3485 |
6.2 |
8.2 |
16 |
15 |
3 |
2:7:1 |
3:6:1 |
3.5 |
24 |
9 |
|
6.5 |
20 |
10 |
||||||||
|
USNM 395827 |
0.9 |
0.9 |
22 |
21 |
0:0:10 |
0:0:10 |
0.6 |
41 |
16 |
|
|
Conularia elegantula Meek |
||||||||||
|
SPECIMEN NO. |
L |
HL |
AAMj |
AAMn N |
RLAMj |
RLAMn |
D |
R |
RA |
|
|
AMNH CU 282G h |
3.9 |
6.7 |
20 |
17? |
6 |
0:4:6 |
— |
1.0 |
39 |
8 |
|
4.0 |
37 |
13 |
||||||||
|
CMNH 4584 |
2.7 |
3.2 |
(30?) |
— |
6 |
0:6:4? |
— |
3.5 |
32 |
3 |
|
CMNH 4648 |
3.1 |
(4.5) |
- |
5 |
- |
- |
(4.0) (24) |
- |
||
|
Conularia milwaukeensis Cleland |
||||||||||
|
SPECIMEN NO. |
L |
HL |
AAMj AAMn N |
RLAMj |
RLAMn |
D |
R |
RA |
||
|
USNM 85988 h |
3.1 |
5.0 |
15 |
14 |
6 |
0:0:10 |
0:0:10 |
6.0 |
18 |
5 |
|
MPM 20252 |
4.9 |
6.0 |
12 |
11 |
6 |
0:0:10 |
0:0:10 |
4.0 |
21 |
16 |
|
MPM 22974 |
4.5 |
4.6 |
15 |
13 |
5 |
0:0:10 |
0:0:10 |
4.0 |
24 |
18 |
|
6.0 |
18 |
8 |
||||||||
|
8.0 |
20 |
9 |
|
476 |
Annals of Carnegie Museum |
VOL. 55 |
||||||||
|
Conularia multicostata Meek and Worthen |
||||||||||
|
SPECIMEN NO. |
L |
HL |
AAMj AAMn N |
RLAMj |
RLAMn |
D |
R |
RA |
||
|
USNM 50157 plh |
2.8 |
6.7 |
20 |
18 |
— |
0:0:10 |
0:0:10 |
4.5 |
32 |
16 |
|
UK 6089 |
6.3 |
8.5 |
— |
11? |
3 |
— |
0:0:10? |
9.5 |
52 |
12 |
|
AMNH 6713 |
6.5 |
9.3 |
23 |
22 |
2 |
0:0:10 |
0:0:10 |
5.0 |
40 |
17 |
|
8.0 |
44 |
9 |
||||||||
|
CM 34533 |
8.2 |
11.2 |
16 |
15 |
2 |
0:0:10 |
0:0:10 |
4.0 |
25 |
— |
|
5.5 |
29 |
12 |
||||||||
|
Conularia pyramidalis Hall |
||||||||||
|
SPECIMEN NO. |
L |
HL |
AAMj AAMn N |
RLAMj |
RLAMn |
D |
R |
RA |
||
|
AMNH 33017 1 |
2.3 |
3.3 |
18 |
17 |
3 |
0:0:10 |
0:0:10 |
3.0 |
14 |
11 |
|
5.5 |
11 |
16 |
||||||||
|
7.5 |
9 |
9 |
||||||||
|
NYSM 3490 |
11.8 |
19.2 |
23 |
17 |
1 |
0:0:10 |
0:0:10 |
8.0 |
7 |
22 |
|
12.0 |
10 |
9 |
||||||||
|
15.0 |
10 |
5 |
||||||||
|
Conularia subcarbonaria Meek |
and Worthen |
|||||||||
|
SPECIMEN NO. |
L |
HL |
AAMj AAMn N |
RLAMj |
RLAMn |
D |
R |
RA |
||
|
UIPC 10680 h |
8.6 |
11.9 |
21 |
19 |
4 |
0:0:10 |
0:0:10 |
8.0 |
18 |
4 |
|
FMNH UC 18494 |
24.5 |
33.5 |
10 |
9 |
4 |
0:0:10 |
0:0:10 |
18.0 |
27 |
5 |
|
25.0 |
31 |
6 |
||||||||
|
FMNH UC 6289 |
7.7 |
- |
- |
- |
3 |
0:0:10 |
0:0:10 |
- |
- |
- |
|
FMNH UC 6610 |
3.1 |
- |
- |
- |
2 |
- |
- |
- |
60 |
- |
|
Conularia tuzoi Clarke |
||||||||||
|
SPECIMEN NO. |
L |
HL |
AAMj AAMn N |
RLAMj |
RLAMn |
D |
R |
RA |
||
|
NYSM 9404 h |
6.7 |
11.3 |
(10) |
— |
— |
(0:0:10) |
— |
6.0 |
26 |
9 |
|
9.0 |
36 |
10 |
||||||||
|
Conularia ulsterensis Howell |
||||||||||
|
SPECIMEN NO. |
L |
HL |
AAMj AAMn N |
RLAMj |
RLAMn |
D |
R |
RA |
||
|
PU 42071 h |
1.8 |
2.5 |
18? |
— |
7 |
2:0:8? |
— |
4.5 |
45 |
13 |
|
CM 34528 |
1.9 |
3.2 |
17? |
— |
7 |
0:0:10? |
— |
2.2 |
60 |
12 |
|
2.8 |
84 |
11 |
||||||||
|
CM 34529 |
1.8 |
(2.8) |
(18) |
(15) |
6 |
0:0:10 |
0:0:10 |
(2.0) |
62 |
12 |
|
Conularia undulata Conrad |
||||||||||
|
SPECIMEN NO. |
L |
HL |
AAMj AAMn N |
RLAMj |
RLAMn |
D |
R |
RA |
||
|
AMNH 41093 1 |
10.0 |
15.5 |
13 |
10 |
6 |
1:0:9 |
1:0:9 |
6.0 |
21 |
18 |
|
10.0 |
20 |
11 |
||||||||
|
13.0 |
27 |
4 |
||||||||
|
AMNH 5440 |
8.3 |
— |
(8) |
4 |
— |
— |
- |
30 |
- |
|
|
NYSM 3482 |
10.2 |
11.8 |
18 |
15 |
5 |
0:0:10 |
0:0:10 |
4.5 |
17 |
9 |
|
9.5 |
24 |
10 |
||||||||
|
CM 34532 |
7.5 (18.0) |
21 |
14 |
5 |
0:0:10 |
0:0:10 |
18.0 |
32 |
9 |
|
|
Paraconularia alpenensis Babcock and Feldmann, new species |
||||||||||
|
SPECIMEN NO. |
L |
HL |
AAMj |
AAMn N |
RLAMj |
RLAMn |
D |
R |
RA |
|
|
GSC 85060 h |
1.5 |
3.6 |
21 |
16 |
_ |
2:2:6 |
4:2:4 |
3.0 |
14 |
9 |
|
4.0 |
14 |
10 |
1986 Babcock and Feldmann— and Reticulaconularia
All
Paraconularia alternistriata (Shimer)
|
SPECIMEN NO. |
L |
HL |
AAMj AAMn N |
RLAMj |
RLAMn |
D |
R |
RA |
||
|
GSC 5111 h |
0.8 |
1.9 |
11 |
10 |
0 |
10:0:0 |
3:7:0 |
1.0 |
28 |
11 |
|
Paraconularia blairi (Miller and Gurley) |
||||||||||
|
SPECIMEN NO. |
L |
HL |
AAMj AAMn N |
RLAMj |
RLAMn |
D |
R |
RA |
||
|
UCGM 3986 1 |
7.4 |
11.2 |
(22) |
20 |
3 |
6:4:0 |
3:5:2 |
5.5 |
11 |
8 |
|
8.5 |
7 |
9 |
||||||||
|
11.0 |
9 |
15 |
||||||||
|
UMC 4270 |
3.4 (11.5) |
23? |
— |
3 |
7:2:1? |
- |
(16.0) |
6 |
19 |
|
|
UCGM 3985 |
6.8 (16.0) |
16 |
2 |
- |
7:3:1 |
12.5 |
6 |
- |
||
|
UCGM 3984 pi |
9.2 |
15.5 |
— |
18 |
3 |
- |
5:4:1 |
8.0 |
8 |
15 |
|
11.0 |
6 |
14 |
||||||||
|
14.0 |
6 |
15 |
||||||||
|
AMNH 25056 |
13.0 |
19.0 |
11 |
10 |
2 |
8:1:1 |
2:6:2 |
7.0 |
13 |
9 |
|
12.5 |
8 |
12 |
||||||||
|
16.0 |
7 |
17 |
||||||||
|
Paraconularia byblis (White) |
||||||||||
|
SPECIMEN NO. |
L |
HL |
AAMj AAMn N |
RLAMj |
RLAMn |
D |
R |
RA |
||
|
UMMP2167 h |
6.7 |
10.0 |
22 |
(14) |
1 |
1:4:6 |
— |
6.5 |
15 |
12 |
|
CMNH 4691 |
2.8 |
3.8 |
26 |
— |
1 |
1:7:2 |
— |
2.5 |
28 |
13 |
|
USNM 409489 |
1.7 |
2.1 |
19 |
14 |
1 |
1:6:3 |
6:2:2 |
1.0 |
29 |
18 |
|
Paraconularia chagrinensis Babcock and Feldmann, |
new species |
|||||||||
|
SPECIMEN NO. |
L |
HL |
AAMj AAMn N |
RLAMj |
RLAMn |
D |
R |
RA |
||
|
CMNH 6633 h |
3.9 |
4.9 |
28 |
21 |
4 |
0:9:1 |
0:9:1 |
2.0 |
20 |
12 |
|
3.0 |
18 |
9 |
||||||||
|
4.2 |
16 |
10 |
||||||||
|
CMNH 1622 p |
3.4 |
8.3 |
-- |
20? |
— |
0:7:3 |
0:8:2 |
5.0 |
16 |
9 |
|
CMNH 1818 p |
2.0 |
- |
- |
— |
4 |
— |
- |
- |
- |
- |
|
CMNH 1674 p |
4.5 (10.0) |
- |
3 |
- |
- |
- |
- |
|||
|
Paraconularia chesterensis (Worthen) |
||||||||||
|
SPECIMEN NO. |
L |
HL |
AAMj AAMn N |
RLAMj |
RLAMn |
D |
R |
RA |
||
|
ISGS 2489 h |
11.7 |
17.2 |
14 |
10 |
4 |
4:5:1 |
4:2:4 |
7.5 |
11 |
10 |
|
10.0 |
10 |
9 |
||||||||
|
14.0 |
8 |
11 |
||||||||
|
USNM 50156 |
2.7 |
3.9 |
21 |
17 |
5 |
4:3:3 |
8:2:2 |
1.0 |
25 |
8 |
|
2.0 |
20 |
8 |
||||||||
|
lUPC 17413 |
9.7 |
16.6 |
(14) |
5 |
(4:5:1) |
— |
15.5 |
7 |
15 |
|
|
lUPC 11313 |
12.8 |
25.5 |
9 |
9 |
4 |
4:5:1 |
4:5:1 |
14.0 |
6 |
9 |
|
20.0 |
5 |
9 |
||||||||
|
23.0 |
6 |
13 |
||||||||
|
lUPC 6458 |
9.8 |
35.5 |
8 |
(7) |
4 |
5:2:3 |
2:6:3 |
26.0 |
5 |
9 |
|
30.0 |
4 |
11 |
||||||||
|
Paraconularia missouriensis (Swallow) |
||||||||||
|
SPECIMEN NO. |
L |
HL |
AAMj AAMn N |
RLAMj |
RLAMn |
D |
R |
RA |
||
|
FMNH UC 6639 plh |
11.0 |
14.6 |
19 |
_ |
— |
1:9:0 |
— |
6.5 |
6 |
9 |
|
9.5 |
4 |
6 |
478
Annals of Carnegie Museum
VOL. 55
|
FMNH UC 6628 |
11.2 |
15.5 |
15 |
11 2 |
1:9:0 |
1:8:1 |
6.0 |
6 |
6 |
|
13.0 |
5 |
10 |
|||||||
|
FMNH UC 6627 |
12.8 |
19.5 |
14 |
11 3 |
2:8:0 |
3:7:0 |
10.0 |
6 |
11 |
|
14.5 |
5 |
17 |
|||||||
|
19.0 |
5 |
11 |
|||||||
|
AMNH 28692 |
5.7 |
13.8 |
- |
10 - |
— |
9:1:0 |
10.0 |
6 |
8 |
|
UMMP 26740 |
3.6 |
6.4 |
21 |
15 2 |
3:5:2 |
4:4:2 |
4.0 |
10 |
9 |
|
Paraconularia oklahomaensis Babcock and Feldmann, new species |
|||||||||
|
SPECIMEN NO. |
L |
HE |
AAMj AAMn N |
RLAMj |
RLAMn |
D |
R |
RA |
|
|
USNM 409801 h |
4.4 |
5.4 |
19 |
17 0 |
4:1:5 |
2:4:4 |
2.0 |
24 |
12 |
|
4.0 |
12 |
13 |
|||||||
|
Paraconularia planicostata (Dawson) |
|||||||||
|
SPECIMEN NO. |
L |
HL |
AAMj AAMn N |
RLAMj |
RLAMn |
D |
R |
RA |
|
|
RM(MU) 2749 h |
4.5 |
7.3 |
1 1? |
9 0 |
6:3:1 |
5:4:1 |
3.5 |
19 |
12 |
|
6.0 |
13 |
15 |
|||||||
|
GSC 7715 |
3.3 |
4.9 |
18 |
14 0 |
5:4:2 |
5:3:2 |
3.0 |
14 |
15 |
|
GSC 24644 |
1.6 |
5.3 |
13 |
12 0 |
3:6:1 |
5:3:2 |
3.5 |
20 |
12 |
|
Paraconularia recurvatus Babcock and Eeldmann, |
new species |
||||||||
|
SPECIMEN NO. |
L |
HL |
AAMj AAMn N |
RLAMj |
RLAMn |
D |
R |
RA |
|
|
USNM 409806 h |
4.8 |
7.8 |
16 |
15 3 |
2:5:3 |
2:4:4 |
4.0 |
18 |
12 |
|
6.0 |
(26) |
8 |
|||||||
|
USNM 409807 p |
0.8 |
- |
- |
- 2 |
- |
- |
- |
28 |
- |
|
USNM 409808 p |
1.1 |
2.6 |
16 |
- 3 |
3:6:1 |
3:5:2 |
1.5 |
26 |
8 |
|
USNM 409809 p |
0.7 |
- |
- |
- 3 |
- |
- |
- |
24 |
- |
|
Paraconularia salinensis (Whiteaves) |
|||||||||
|
SPECIMEN NO. |
L |
HL |
AAMj AAMn N |
RLAMj |
RLAMn |
D |
R |
RA |
|
|
GSC 4292 h |
2.6 |
3.0 |
(24) |
21 4 |
7:2:1 |
5:3:2 |
1.0 |
24 |
13 |
|
1.5 |
23 |
8 |
|||||||
|
Paraconularia sorrocula (Beede) |
|||||||||
|
SPECIMEN NO. |
L |
HL |
AAMj AAMn N |
RLAMj |
RLAMn |
D |
R |
RA |
|
|
NYSM 9414 h |
2.3 |
2.9 |
23 |
- 6 |
1:7:2 |
— |
0.8 |
20 |
11 |
|
1.2 |
19 |
14 |
|||||||
|
Paraconularia subulata (Hall) |
|||||||||
|
SPECIMEN NO. |
L |
HL |
AAMj AAMn N |
RLAMj |
RLAMn |
D |
R |
RA |
|
|
AMNH 32404 1 |
0.6 |
0.8 |
21 |
18 4 |
5: 2:3 |
6:1:3 |
0.5 |
56 |
11 |
|
FMNH UC 6961 |
1.6 |
2.4 |
18 |
12 5 |
0:10:0 |
3:7:0 |
1.0 |
35 |
18 |
|
1.5 |
35 |
11 |
|||||||
|
UIPC 10866 |
6.6 |
7.8 |
__ |
12 4 |
7:1:2 |
5.0 |
8 |
22 |
|
|
UMMP 2178 |
0.9 |
(5.5) |
— |
= 4 |
— |
24 |
(19) |
||
|
UMMP 245 |
0.5 |
1.4 |
22 |
(20) 4 |
0:7:3 |
6:3:1 |
1.0 |
30 |
11 |
|
OC 8309 |
5.4 |
12.8 |
12 |
10 2 |
1:8:1 |
4:5:1 |
11.0 |
8 |
15 |
|
Paraconularia |
wellsvillia Babcock and Feldmann, |
new species |
|||||||
|
SPECIMEN NO. |
L |
HL |
AAMj AAMn N |
RLAMj |
RLAMn |
D |
R |
RA |
|
|
CM 35001 h |
5.7 |
12.5 |
(14) |
(13) 3 |
1:8:1 |
5:4:1 |
8.5 |
4 |
26 |
|
10.0 |
5 |
31 |
1986 Babcock and Feldmann— il4iL4COivc/i^i?/y4 and Reticulaconularia 479
|
CM 34502 p |
5.8 (12.5) |
(18)? |
3 |
3:6:1? |
- |
7.5 |
9 |
(24) |
||
|
12.0 |
7 |
(18) |
||||||||
|
Paraconularia yochelsoni Babcock and Feldmann, : |
new species |
|||||||||
|
SPECIMEN NO. |
L |
HL |
AAMj AAMn N |
RLAMj |
RLAMn |
D |
R |
RA |
||
|
UMMP 45499 h |
2.2 |
2.6 |
20 |
18 |
0 |
5:3:2 |
10:0:0 |
0.6 |
15 |
18 |
|
1.5 |
14 |
17 |
||||||||
|
UUMP 45500 p |
2.6 |
3.1 |
17 |
0 |
6:1:2 |
— |
0.6 |
18 |
15 |
|
|
1.5 |
13 |
15 |
||||||||
|
UMMP 65509 o |
1.3 |
1.7 |
15 |
- |
0 |
8:1:1 |
- |
0.6 |
18 |
20 |
|
Reticulaconularia penouili (Clarke) |
||||||||||
|
SPECIMEN NO. |
L |
HL |
AAMj AAMn N |
RLAMj |
RLAMn |
D |
R |
RA |
||
|
NYSM 9412 h |
4.6 |
6.7 |
59 |
(22) |
2 |
2:5:3 |
3:5:2 |
3.0 |
14 |
4 |
|
4.5 |
21 |
3 |
||||||||
|
6.0 |
14 |
0 |
||||||||
|
GSC 87242 |
5.5 |
6.4 |
— |
30? |
2 |
— |
5:4:1? |
1.3 |
15 |
7 |
|
3.3 |
12 |
8 |
||||||||
|
Reticulaconularia sussexensis |
(Herpers) |
|||||||||
|
SPECIMEN NO. |
L |
HL |
AAMj AAMn N |
RLAMj |
RLAMn |
D |
R |
RA |
||
|
NJSM 10749 h |
2.7 |
3.1 |
32 |
26 |
1 |
0:2:8 |
— |
2.5 |
20 |
11 |
|
3.5 |
36 |
8 |
||||||||
|
NJSM 10750 p |
1.5 |
1.8 |
(27) |
24 |
2 |
1:2:7 |
2:1:7 |
0.6 |
39 |
14 |
1.5 16 11
Back issues of many Annals of Carnegie Museum articles are available, and a few early complete volumes and parts are listed at half price. Orders and inquiries should be addressed to: Publications Secretary, Carnegie Museum, 4400 Forbes Avenue, Pittsburgh, Pa. 15213.
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0/ CARNEGIE MUSEUM
CARNEGIE MUSEUM OF NATURAL HISTORY 4400 FORBES AVENUE • PITTSBURGH, PENNSYLVANIA 15213 VOLUME 55 31 DECEMBER 1986 ARTICLE 17
FLORAL MORPHOLOGY AND VASCULAR ANATOMY OF AMIANTHIUM MUSCAETOXICUM (WALTER) A. GRAY (LILIACEAEWERATREAE) WITH NOTES ON DISTRIBUTION AND TAXONOMY
Frederick H. Utech Abstract
Presentation of pedicel to stigma vasculature of the monotypic Amianthium muscae- toxicum as a Veratrean example documents the perigyny, spiral insertion of floral parts and an apocarpous gynoecium which are encountered. Total floral vascularization is derived from three lower pedicel bundles. A spiral series of divisions and fusions in the upper pedicel produces three compound outer tepal and three inner tepal bundles. From the former, outer tepal medians, tepal laterals, stamen and dorsal bundles result, while from the latter, inner tepal medians, tepal laterals and stamen bundles result. Due to perigyny, their formation is above that at which the ventral and ovular supplies are established. The ventral supply is spirally derived from continuing bundles following the formation of the compound inner tepal bundles. Within each septal arm, a compound lateral is associated with a compound ventral and both undergo radial divisions. Opposing lateral and ventral products fuse as the perigynous condition ends and the three carpels are freed. Each carpel has two simple ventrals, two laterals and a dorsal in the upper perigynous zone and two fusion ventrals and a dorsal in the upper freed zone. Neither tepal glands nor nectaries occur in this species. Epidermal cells characterized by ho- mogenous tannins occur in the pedicel, receptacle wall, abaxial tepal surfaces, filaments and freed carpels. Raphides commonly occur in cells along the lower margins of the inner tepals.
Introduction
Amianthium muscaetoxicum (Walter) A. Gray is a monotypic species of eastern North America with centers of distribution in both the Ozark-
Submitted 15 February 1986.
481
482
Annals of Carnegie Museum
VOL. 55
Fig. 1. — Distribution of Amianthium muscaetoxicum in eastern United States based on published maps (Steyermark, 1963; Radford et al., 1968; Johnson, 1969; Smith, 1978; Wherry et al., 1979) and collections at Carnegie Museum of Natural History (open circles). Johnson (1969) surveyed 46 herbaria in his southeastern United States treatment of the Liliaceae. Sampled populations are indicated by circled stars.
ian and Appalachian regions (Fig. 1). It occurs in mesic to dry wooded slopes and coastal plain savannas and pinelands. Familial and subfa- milial names associated with A. muscaetoxicum have changed during the last century, but the tribal association with Melanthium, Schoen- caulon, Stenanthium, Veratrum and Zigadenus “sensu lato” (including Toxicoscordion, Anticela, Tracyanthus and Oceanoros) has remained. Bentham and Hooker (1883) included these six genera in the tribe
1986
Utech —Amianthium
483
Veratreae, as did Engler (1889), Krause (1930), Melchior (1964), Thome (1968), Takhtajan (1969), and Hutchinson (1934, 1959, 1973). Within this tribal grouping, two lines —the Veratrum-Melanthium and the Zig- adenus lines— are generally recognized (Anderson, 1940; Preece, 1956; Zimmerman, 1958; Kupchan et al., 1961; Ambrose, 1975, 1980; Ster- ling, 1982). Amianthium and Stenanthium are generally placed some- where between these two evolutionary lines. Engler (1 889) initiated the association of the Veratreae with the subfamily Melanthioideae. This subfamily has been twice segregated as a separate family, the Melan- thiaceae (Gates, 1918; Small, 1933; Dahlgren, 1980; Dahlgren and Clifford, 1982; Dahlgren and Rasmussen, 1983; Dahlgren et al., 1985) and the Colchicaceae (Baker, 1879).
Genera in the other tribes of the Englerian Melanthioideae share separate styles and septicidal capsules with the genera in the Veratreae, though the latter possesses unusual extrorse anthers with valvular de- hiscence that open into peltate discs (Krause, 1930; Zimmerman, 1958; Kupchan et al., 1961) and usually many bitegmic, basitropic and cam- pylotropous ovules per carpel (Sterling, 1982), except for Amianthium which has two, rarely four.
Amianthium of Asa Gray (1837) is a conserved generic name (Farr et al., 1979; Voss, 1983) and its monotypic species, A. muscaetoxicum (Walter) A. Gray (1837), has gone under various names since it was first described by Walter in 1788 as Melanthium muscaetoxicum. Later synonyms include: Melanthium laetum Solander in Aiton (1789), Me- lanthium myoctonum J. F. Gmelin (1796), Helonias erythrosperma Michaux (1803), Helonias laeta (Solander in Aiton) Kew-Gawler (Cur- tis, 1805), Amiantanthus muscaetoxicum (Walter) Kunth (1843), Zig- adenus muscaetoxicus (Walter) Regel (1883), and Chrosperma mus- caetoxicum (Walter) Kuntze (1891). Excluded species of Amianthium and their current status include: A. nuttallii A. Gray var. alpha (1837) {Zigadenus nuttallii A. Gray ex S. Watson), A. nuttallii A. Gray var. beta (1837) (Z. paniculatus (Nutt.) S. Watson), A. angustifolium A. Gray (1837) (Z. densus (Desr.) Femald), A. leimanthoides A. Gray (1837) (Z. leimanthoides A. Gray), and^. texanum (Bush) Gates (1918) (Z. leimanthoides A. Gray).
Considerable biological information is implied in this species bi- nomial. Amianthium is derived from the Greek amianthos for “un- spotted” and anthos for “flower,” an allusion to a major generic char- acter, that is the lack of glands or nectaries on the perianth. The specific name, muscaetoxicum, translates literally as “fly poison.” In 1883, a note in Gardeners’ Chronicle “stated that the root, when bmised and mixed with honey, acts as a poison to flies.” While the species is commonly known as “fly poison,” it has also been known as “crow poison” and “swagger grass” (Muenscher, 1939, 1960). Several poi-
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sonous alkaloids are associated with the leaves and underground bulbs of the species and are reported to cause death in cattle and sheep (Pammel, 191 1; Marsh etal., 1918; Muenscher, 1939, 1960; Kingsbury, 1964). Amianthine, a steroidal or modified steroidal alkamine alkaloid with a C27H41O2N formula, has been described from the roots and leaves of A. muscaetoxicum as well as jervine and two unidentified ester alkaloids (Neuss, 1953). The latter alkaloids were shown to be responsible for the high toxicity of the species (Neuss, 1953). Although numerous and highly poisonous alkaloids commonly occur throughout the tribe Veratreae, amianthine has only been reported in Amianthium (Kupchan et al., 1961; Willaman and Schubert, 1961; Hegnauer, 1 963).
Two meiotic chromosome counts of « = 16 have been reported for A. muscaetoxicum: Ambrose (1975) from Bear Creek, Pennsylvania and Preece (1956) from Big Laurel Gap, Yancey County, North Car- olina. Excluding various polyploid multiples, Melanthium, Veratrum and Zigadenus (^Zygadenus; Preece, 1956) all share n = 16 (Fedorov, 1969; Moore, 1973; Goldblatt, 1981). It is quite apparent that x ^ 8 is the basic number unifying the Veratreae.
In focusing on both the floral morphology and vascular anatomy of A. muscaetoxicum a comparative model of a Veratrean gynoecium will be established in this paper. Similar treatments for the other tribal members of the Veratreae, such as Melanthium, Stenanthium, Vera- trum and Zigadenus, are in preparation and will continue the studies of Buxbaum (1925, 1927), Anderson (1940), El-Hamidi (1952), Am- brose (1975, 1980) and Sterling (1982) on the vascularization of the Veratrean carpel.
Materials and Methods
Flowering and fruiting inflorescences of Amianthium muscaetoxicum were collected from two different populations— Pennsylvania: Clinton County, Mt. Tableland, ca. 4.5 mi E of Sinnemahoning, 20 July 1979, Utech 79-241 (CM), and North Carolina: Macon County, ca. 3.5 mi N of Highlands, near Whiteside Mt., Nantahala National Forest, 28 July 1982, Utech and Ohara 82-270 (CM). The collected materials were fixed in acetic- ethanol (1:3) for 10 h with subsequent storage in 70% ethanol. Standardized paraffin sectioning (14-16 microns) and staining (saffarin-methylene blue) techniques (Johansen, 1940; Sass, 1958) were used on samples 00 flowers and 10 young fruits of varying ages) from both populations. As an additional check on these serial sections, whole flowers and fruits were cleared and stained in a NaOH-1% fuchsin mixture (Fuchs, 1963).
Composite photomicrographs (Figs. 3-5, 7-8, 10) present the vascular floral anatomy and morphology of A. muscaetoxicum, whereas Figs. 6 and 9 are summary line diagrams for the species. No teleological implications are intended in the descriptive ascent and departure of the various floral bundles which are letter-coded for ease in comparison. This coding parallels that used in our previous liliaceous studies (Utech, 1978^-1978^, 1979a, \919b, 1982, 1 984; Utech and Kawano, 1975, 1976, 1980, 1981).
Observations
Amianthium muscaetoxicum is a glabrous, subscapose, perennial herb from a thick bulb (Fig. 2). At anthesis, the linear basal leaves are
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Fig. 2.— -Flowering individuals of A. muscaetoxicum in Macon County, North Carolina (Utech and Ohara 82-270, CM) (scale indicated).
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Fig. 3. -Cross-sections from the pedicel and lower receptalar areas of A. muscaetoxicum showing the spiral opening of the locules, the degree of perigyny and distribution of epidermal tannin cells. A. Mid-pedicel section showing the formation of three compound outer tepal (OT) bundles. B. Upper pedicel section showing formation of three compound inner tepal (IT) bundles and departure of compound OT bundles. Due to spiral bundle formation, a fusion bundle is opposite the upper OT bundle and a gap opposite the lower left OT bundle (arrows). C. Transition between upper pedicel and lower receptacle areas showing he opening of two locules, the departure of three IT bundles and the remaining central bundles which form the ventral supply. D. Lower receptacle area with perigyny
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shorter than the stem. The few cauline leaves are progressively reduced and bract-like. The bracteate raceme is at first ovoid to conic, but at maturity it becomes cyclindric, measuring 4-14 cm long by 2-4 cm wide. Initially the flowers are white, but following anthesis the persis- tent tepals turn yellowish green or greenish purple and present a most striking color pattern.
The flowering pedicels are normally 7-17 mm long and ascending. In fruit, the pedicels are decidedly horizontal and much elongated. Internally, the fruiting and flowering pedicels have the same bundle number, though the vasculature in fruit is surrounded by a scleren- chymatous sheath. Lower flowering pedicel cross-sections are broadly triangular with three large, centrally arranged bundles. These three bundles establish the complete floral vasculature and are located on the radii which run from the middle of the “flat side” to the section’s center. These three radii are designated the outer tepal (OT) radii. The three radii from the “comers” to the center are designated the inner tepal (IT) radii. Mid-pedicel cross-sections are characterized by broad fluting and ridges (Figs. 3 and 9) which continue through the perigynous zone.
The three lower pedicel bundles along OT radii undergo tri-parted, radial divisions with three resulting bundles from each division. These divisions occur at slightly different levels, that is they are not co-planar, but rather in a spiral pattern. Within each division a gap is created by the outward departure of a central bundle product. Three such central bundles, designated compound outer tepal (OT) bundles (dorsal-com- pound bundle; Sterling, 1982) depart along OT radii and remain free of other vasculature. Eventually they establish the dorsals (D), the outer tepal medians (OTM), the outer tepal laterals (OTL), and the outer stamen (OS) bundles (Figs. 6, 9A-D).
The two resulting lateral bundles which are opposite a gap following the tri-parted divisions fuse with similar adjacent laterals along the IT radii. These fusion bundles are formed in a spiral pattern and undergo a tri-parted division similar to that observed at a lower level among the three original bundles. The central bundles of this second set of divisions depart along the IT radii and establish the compound inner tepal (IT) bundles (“zwischenbundel”; Sterling, 1982).
evident and three open locules with formation of the central ventral supply. E. Mid- receptacle area showing the central hole and the ventral supply. F. Upper receptacle area showing the spiral septal arm formation within the perigynous zone, inter-locular con- nections and placental supply. Dorsal bundles are not established at this level (scale indicated).
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Fig, 4.— Cross-sections from the lower receptacle to upper perigynous areas of A. mus- caetoxicum. A. Lower receptacle section showing the departure of compound OT and IT bundles, opening of two locules and formation of three bundles opposite IT bundles which establish the ventral supply. B, Section above A showing three open locules and the central hole. While three IT bundles depart, fusion and division occur among the remaining septal arm bundles. C. Section above B showing three freed septal arms between the inter-connected locules. Each septal arm has two laterals (L) and a compound ventral (V) from which placental bundles (P) arise. D. Section above C showing ovule supply via placental (P) bundles and the paired laterals and ventrals within the septal arms. E. Section above D showing the division of the compound ventral bundle (V) and the two laterals (L) within a septal arm. Papilloid nurse cells lining the inner septal arm
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From the second set of tri-parted divisions, the two remaining lateral bundles fuse laterally with adjacent laterals, close the OT radii gaps formed when the compound OT bundles departed and eventually es- tablish the total ventral supply (Fig. 6). These three fusion bundles undergo simple radial divisions in a spiral pattern. The two resulting products of this radial division fuse laterally with a similar adjacent lateral and form three fusion bundles along IT radii which close the gaps formed during the departure of the compound inner tepal (IT) bundles. Subsequent divisions among the compound OT and IT bun- dles will be discussed later. A given pedicel cross-section will usually show one bundle departing, another being formed and a gap in an area where a third will be formed.
The gynoecium and its associated vasculature can best be described in two parts: that in the lower perigynous zone and that in the upper freed zone. Ventral supply formation, locule opening and ovule pla- centation all occur within the lower perigynous zone (Figs. 3C“F; 4, 6). The dorsal bundles, on the other hand, which are derived from compound outer tepal (OT) bundles are established in the upper limits of the peripheral perigynous zone.
The ridged outline observed in the lower pedicel occurs up through the upper perigynous region where the tepals and stamens are freed (Figs. 3, 4F). Following the formation and departure of both the com- pound OT and IT bundles, a triangular vascular zone (“stele”) remains in the central area (Figs. 3C-D, 4A-B). The three large bundles along IT radii establish the triangle’s comers. These three complex fusion bundles which closed the gaps formed by the departure of compound IT bundles (Fig. 6) generally have two phloem caps or poles. Once these three comer bundles are established, the three locules which are perpendicular to OT radii open spirally.
With locular opening, each of the three compound bundles along IT radii undergoes a simple tangential division (Figs. 4B-C, 5A, C). Both resulting bundles are along the IT radii. The outermost division product has normally arranged xylem (adaxial) and phloem (abaxial), while the inner product has reversed conducting elements. Phloem strands fre- quently are observed to lag and may rarely and irregularly anastomose with other bundles (Fig. 5A-B).
margins are weakly appressed. F. Upper periygnous section showing the spiral arrange- ment of tepals, stamens and partially, freed carpels. Dorsals (D) are established at this level. Paired lateral (L) and ventral (V) products within septal arms are indicated as are outer tepal medians (otm), inner tepal medians (itm), outer stamen (os) and inner stamen (is) bundles. Epitepaly between an inner tepal and inner stamen is shown in the upper right (scale indicated).
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Fig. 5. -Cross-section of the middle to upper gynoecium of A. muscaetoxicum. A. Central zone showing three appressed septal arm tips and radial arm separation with paired lateral (L) and ventral (V) bundles. B. Section, same level as A, showing an outer stamen (OS) bundle and dorsal (D). An inner carpellary wall indentation, not a notch, is associated with each dorsal. C. Section above A showing the separated gynoecium and perianth. Within a septal arm, a given ventral and opposing lateral fuse to form a new, compound central in the same position as the lower, simple ventral. D. Section above C showing two compound ventrals (V) and dorsal (D) per carpel. The inner septal arm tips are further divided (arrow) compared with A. E. Section above D showing three free carpels each with a dorsal (D) and two ventrals (V). F. Stylar zone with the dorsal (D) and two ventrals (V) still present in each carpel (scale indicated).
The appearance at this level of a central opening or ‘"hole” along the central floral axis (Figs. 3E, 4B) indicates inter-connection of the three locules will follow. Openings develop from this hole along the OT radii to each of the three locules (Fig. 3F). As openings inter-connect the
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three locules, three septal arms or wings are established along the IT radii. Within these septal arms further divisions occur among the paired ventral supply bundles. The outermost member of each septal bundle pair, that is the one with normally arranged conducting elements, di- vides radially and establishes two lateral bundles (L) (Figs. 4C-D, 5A, 6). (These bundles, here designated as laterals (L), could also be called septal axials.) The innermost member of each septal bundle pair, that is the one with reversed conducting elements, also undergoes a radial division which establish two placental (P) or ovule supplying bundles. The parental bundle which established the two placental (P) bundles also undergoes a radial division and establishes two ventral (V) bundles. Each septal arm at this level has a pair of laterals (L) with normally arranged conducting elements and a pair of ventrals (V) with reversed elements (Figs. 4C-F, 6).
Each locule has two, rarely four, bitegmic, basitropic, campylotro- pous ovules. The placental (P) bundles supplying these ovules depart quite horizontally. Usually one, rarely two, ellipsoidal, lustrous, dark reddish brown to black seeds are found in each carpel. The wingless seeds measure 1.5--2.0 mm wide by 4. 5-6.0 mm long. Dehiscence begins in the upper stylar area and continues along the zones where the septal arms or margins meet (Fig. 5D-F).
The three dorsals (D) are the last carpellary vascular elements to be established and this occurs in the upper perigynous zone. The dorsals are associated with a zone of parenchyma cells which protrude as a locular indentation, not a notch (Figs. 4F, 5B, D-E, 7B). As the septal arms are subdivided along IT radii (Figs. 4E-F, 5 A), the outer carpellary wall is freed from the perigynous zone. Septal arm subdivision follows the changing distribution of epidemal tannin cells which line the outer carpellary wall margins (Fig. 5A, C, D). Within each subdivided septal arm, two sets of vascular fusions occur. The two laterals (L) fuse with two opposite ventral bundles (V) which are along the same radii. The two resulting fusion bundles, here designated as ventrals (V), are in the same location as the two lower, simple ventrals (V) (Figs. 5A, C-D, 6). It should be noted that while the ventrals prior to fusion and after fusion are designated in the same way, there is a difference. The later ventral is a terminal fusion product. With the formation of these fusion ventrals (V), the three carpels are freed from one another (Fig. 5D-F) in what can best be described as an apocarpous condition. Each freed carpel has a dorsal (D) and two ventrals (V) which continue into the stylar zone (Fig. 5F). There is no terminal carpellary fusion between the ventrals or between the ventrals and the dorsal.
In most liliaceous species with a superior ovary, both the tepal and stamen vasculatures are well established before the locules open. This is not the case in A. muscaetoxicum (Figs. 3C-F, 9) and other members
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of the Veratreae due to the basal perigynous condition. The six freed perianth parts, that is, the three outer and three inner tepals which are spirally inserted (Fig. 4F), are separate to their respective bases and spread widely. The inner tepals which measure 5-7 mm are slightly longer than the outer tepals. There are no glands or claws associated with these perianth parts as is frequently reported in Melanthium and Veratrum. The abaxial tepal surfaces of all six tepals are characterized by a single epidermal layer containing homogenous tannins (Fig. 8D), This epidermal tannin layer also occurs in the filaments and freed gynoecium. In addition to these tannin cells, there are randomly scat- tered tannin cells throughout the floral tissue. The adaxial tepal sur- faces, on the other hand, lack these tannin cells and instead have papilloid cells (Fig. 8D). These papilloid cells occur from the basal regions of the freed tepals to the apices.
There are other differences between the inner and outer tepals (Figs. 7, 8). The inner tepals are differentiated from the outer in having a short vertical zone of enlarged cells along the basal tepal margins (Figs. 4F, 8B, C). These enlarged cells frequently contain long raphides. Ep- itepaly between the inner stamens and tepals is the rule (Figs. 4F, 8A, B). Epitepaly between the outer stamens and tepals occurs, but it is not as pronounced as the former.
Vascularization of the six tepals and six stamens occurs above the level at which the ovules are supplied (Figs. 3C-F, 4F, 7-9). This is due to the typical perigynous condition within the Veratreae. Six com- pound bundles, derived via fusion and established in the pedicel and lower receptacle levels, are ultimately responsible for the complete vascularization of the tepals and stamens (Figs. 6, 9). These compound bundles have been designated as the compound outer tepal (OT) and compound inner tepal (IT) bundles, respectively, for they are located along those respective radii. Insertion and departure of tepals and stamens follow a spiral pattern (Figs. 4F, 6, 9). A vascular description for one compound OT and IT bundle will illustrate the patterns for both sets since they are free from each other.
In the upper perigynous zone, a compound OT bundle appears tri- angular in cross-section (Figs. 7A, 9). Several complex subdivisions occur within the compound OT bundles which results in the formation
Fig. 6. — Roll-out longitudinal summary diagram for the floral vasculature of A. mus- caetoxicum. A spiral pattern is indicated, in part, by the shifted levels at which the compound OT and IT bundles depart. Various text discussed bundles have been given the following code: L = lateral, V = ventral (simple and compound) and P = placental. Those bundles derived from the compound OT and IT bundles are not shown, see Figs. 7-9.
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of an outer tepal median (OTM), two outer tepal laterals (OTL), an outer stamen bundle (OS) and a dorsal (D). The OTM is established first and has normally arranged xylem (adaxial) and phloem (abaxial) (Figs. 7A, 9B). The remaining product, after the departure of the OTM, undergoes a rapid and complex subdivision in which two OTLs, an OS and a D are established (Figs. 7, 9). The OTM, OS and D bundles all lie along the same radius. It should be noted that the outer tepal laterals are derived from the remaining product bundles, not the OTM. Basally each outer tepal receives three bundles, an OTM and two OTLs. The laterals undergo further radial divisions to establish additional laterals (Fig. 7C). In freed outer tepals, a seven bundled condition is typical, that is three OTLs + OTM + three OTLs.
The origin of the inner tepal median (ITM), the two inner tepal laterals (ITL), and an inner stamen bundle (IS) from a compound inner tepal bundle (IT) is similar to the outer series (Figs. 8, 9). The vascu- larization of the inner tepals and stamens is in a spiral pattern. The ITM, two ITLs and IS bundles have normally arranged xylem (adaxial) and phloem (abaxial). Basally each inner tepal is supplied with three bundles, an ITM and two ITLs. The laterals undergo further radial divisions, as in the outer series, to establish additional laterals (Fig. 8C). In freed inner tepals, a five bundled condition, rarely seven, is encountered, that is two ITLs + ITM + two ITLs. There is no fusion between the laterals or between laterals and medians in either tepal set. Tepal bundles follow a parallel course and end along tepal margins.
The six equal stamens are nearly as long as the inner tepals. The filament’s epidermal layer contains tannins (Figs. 7D, 8C, lOA), while the anther walls do not. The basifixed, extrorse anthers have a valvular (lateral) dehiscence between the confluent thecae which open into a peltate disc (Fig. 10). The endothecium has wall thickenings or bands of the girdle type (Dahlgren and Clifford, 1982). This type of anther and mode of dehiscence is characteristic of the Veratreae.
The difference between outer and inner stamen vascularization which arose from compound OT and IT bundles, respectively, is that a dorsal
Fig. 7.— Vascularization of the outer tepals and stamens in A. muscaetoxicum. A. Mid- perigynous zone showing the departure of an outer tepal median (OTM) from a compound OT complex. B. Section above A showing the further division of the compound OT bundle into two outer tepal laterals (OTL) and an outer stamen (OS) bundle. The locular indentation associated with the dorsal (D) is evident. C. Upper perigynous section above B showing further division among the outer tepal laterals (OTL). D. Section above C showing the freed outer tepal and outer stamen (scale indicated).
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is associated with division of the OT parental bundle and there is no counterpart with the IT compound bundle (Figs, 7B-D, SA-C, 9),
Discussion and Conclusions
In describing the vascular floral anatomy and carpel morphology within selected members of the liliaceous tribe Veratreae, Anderson (1940), El-Hamidi (1952), Ambrose (1975), Sterling (1982), and Utech in this report have noted characteristic similarities and differences be- tween the flowers and fruits of Amianthium muscaetoxicum and the other members of the Veratreae. A common pattern of vascularization appears to be consistently observed within the tribe, while major dif- ferences which are frequently used to differentiate genera are reported. These anatomical and morphological characters include the presence or absence of sutural openings and a central carpellary hole at the lowermost level of ovular insertion, the degree of carpellary separation below the locular apex (an apocarpous tendency), the number of car- pellary lateral or septal axial bundles and a hypogynous or perigynous versus epigynous condition at the lowermost level of ovular insertion. The floral vascular anatomy and carpel morphology of A. muscaetox- icum as reported here will serve as a case study of the continuous vascularization of a Veratrean gynoecium for further comparative work within the tribe.
The pedicel to stigma vasculature of A. muscaetoxicum is most un- usual in that perigyny, spiral insertion of floral parts and apocarpous gynoecia are encountered. Through a complex series of divisions and fusions the complete floral vasculature is derived from three lower pedicel bundles. In the middle to upper pedicel a series of successive divisions and fusions in a spiral sequence produces three compound outer tepal (OT) bundles and three compound inner tepal (IT) bundles. From each compound OT bundle (dorsal-composite bundle; Sterling (1982)), an outer tepal median (OTM), several outer tepal laterals (OTL),
Fig. 8.— Vascularization of the inner tepals and stamens in A. muscaetoxicum with a tepal surface comparison. A. Epitepaly between an inner tepal and stamen is shown as an inner tepal median (ITM) is derived from a compound IT bundle. Subsequent division of the IT bundle establishes two inner tepal laterals (ITL) and inner stamen (IS) bundle. An epidermal tannin layer surrounds both the tepal and stamen. B. Section above A showing epitepaly as well as specialized cells which frequently contain raphides along the inner tepal margin (arrow). These cells are lacking in the outer tepals. C. Section above B showing a freed inner stamen and tepal with specialized cells (arrow). D. Section above C showing the difference in adaxial (tannins) and adaxial (pa = papilloid) tepal surfaces. These papilloid cells occur throughout both adaxial surfaces and extend to the tepafs tips (scale indicated).
so a
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an outer stamen (OS) and a dorsal (D) result. Similarly, from each compound IT bundle (‘'zwischenbundel”; Sterling (1982)), an inner tepal median (ITM), and an inner stamen (IS) bundle result. Due to the perigynous condition, the formation of these tepal, stamen and dorsal bundles is at a level above that at which the ventral and ovular supplies are established.
The spiral origin of the ventral supply is via successive divisions and fusions of continuing lateral branches following the formation of the compound IT bundles. Within each of the three undivided septal arms, that is, the perigynous zone where the three locules are not inter- connected, a lateral bundle (L) (compound septal axial bundle; Sterling (1982)) is associated with a ventral (V) (compound placental bundle; Sterling (1982)). The lateral bundle has normally arranged xylem and phloem while the ventral bundle has reversed conducting elements. There is a radial subdivision within each septal arm of both the ventral and lateral bundles. Each ventral bundle fuses with an opposite lateral bundle as the perigynous condition ends and the three carpels are freed.
It is noteworthy that at a lower level each carpel has five bundles, that is, a dorsal (D), two laterals (L) and two ventrals (V) while at a higher and freed carpellary level there is only a dorsal (D) and two fusion ventrals (V). Terminally, there is no fusion within a carpel be- tween the dorsal and ventrals or between the ventrals. While the ter- minology varies, the above observations are similar to those of Sterling (1982). Anderson (1940) noted that the carpels in Amianthium, Me- lanthium, Veratrum, and Zigadenus were supplied by a dorsal, two laterals and two ventrals. The cross-section drawings of Amianthium by Ambrose (1975; fig. 32G-K) were described as having one dorsal and four ventrals. Simply counting carpellary bundles from selected cross-sections is not adequate for detailed comparative purposes, rather the continuity of the complete floral pattern including origins, fusions and divisions must be followed.
The combination of spirally inserted floral parts as well as their spirally derived vasculature coupled to both a perigynous and apocar- pous gynoecium is most unusual among the “supposedly primitive lilies.” Furthermore, there is a central carpellary hole associated with the gynoecial base. This central hole is internally continuous with the
Fig. 9. “Line drawing showing cross-sections from the upper perigynous zone and the resulting vascularization from both a compound OT bundle (A-D) and a compound IT bundle (E-H). The outer tepal median (OTM), outer tepal laterals (OTL), outer stamen (OS) and dorsal (D) bundles are all derived from the compound OT bundle, while the inner tepal median (ITM), inner tepal laterals (ITL), and inner stamen (IS) bundles are derived from the compound IT bundle.
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three locules and the open stylar canal. Such a central hole according to Sterling (1982) could be taken to represent a partially closed suture or a remnant of an open portion of the stylar canal. The three carpels are essentially free above the perigynous zone. The inner septal wing tips that form the stylar canal are weakly differentiated into papilloid nurse cells. In this inner zone, dehiscence occurs exposing the single, rarely two, wingless seeds of each carpel. Such inner septal margin separation is a variation of the typical septicidal capsule dehiscence where splitting occurs between two adjacent carpels along a common septal radius.
Neither tepal glands nor nectaries occur in Amianthium though they are commonly encountered throughout the Veratreae. The epidermal layers of both abaxial tepal surfaces as well as that of the pedicel, the complete perigynous zone, the filaments and the freed carpels are char- acterized by cells with homogenous tannins. However, the adaxial sur- faces of both the inner and outer tepals have a generalized epidermis of small papilloid cells. Furthermore, as a possible defensive adapta- tion, raphides are observed in cells along the lower, outer margins of the inner tepals. Though there has been limited differentiation between the adaxial and abaxial tepal surfaces, they are not specialized as nec- taries or glands. Travis (1984) observed that foraging beetles are the chief pollinators for A. muscaetoxicum and experimentally demon- strated that this species is nearly self-incompatible and the fecundity and fruit set levels are at least partly pollinator limited. Furthermore, only a small percentage of seeds from self-pollinated plants are viable.
Acknowledgements
The author would like to recognize and thank the M. Graham Netting Research Fund and the O’Neil Botany Field Fund of the Carnegie Museum of Natural Flistory for supporting the field work and the lab related preparation of materials and photographs in the new Biosystematics Laboratory. Mr. William W. Brown and Ms. Nancy J. Perkins deserve special thanks for their artistic aid in figure production.
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