3C ISSN 0097-4463 “ANNALS. of CARNEGIE MUSEUM THE CARNEGIE MUSEUM OF NATURAL HISTORY 4400 FORBES AVENUE • PITTSBURGH, PENNSYLVANIA 15213 VOLUME 67 12 FEBRUARY 1998 NUMBER 1 CONTENTS ARTICLES Significance of nearshore trace-fossil assemblages of the Cambro-Ordovi- cian Deadwood Formation and Aladdin Sandstone, South Dakota .... Thomas M. Stanley and Rodney M. Feldmann 1 Diadectes (Diadectomorpha: Diadectidae) from the early Permian of central Germany, with description of a new species David S Berman, Stuart S. Sumida, and Thomas Martens 53 FEB 2 3 1* C/bRARlti Editors, ANNALS, BULLETIN and SPECIAL PUBLICATIONS: J. L. Carter, Ph.D B. C. Livezey, Ph.D D. R. Watters, Ph.D M. A. Schmidt, ELS, Assistant Editor Manuscripts, subscriptions, orders for individual numbers, and changes of address should be sent to: Office of Scientific Publications Carnegie Museum of Natural History 4400 Forbes Avenue Pittsburgh, PA 15213-4080 Phone (412) 622-3287 Fax (412) 622-8837 E-mail: scipubs@clpgh.org ANNALS OF CARNEGIE MUSEUM is published quarterly by The Carnegie Museum of Natural History, 4400 Forbes Avenue, Pittsburgh, Pennsylvania 15213-4080, by the authority of the Board of Trustees of Carnegie Institute. © 1998 Carnegie Institute. @ This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper). ANNALS OF CARNEGIE MUSEUM VoL. 67, Number 1, Pp. 1-51 12 February 1998 SIGNIFICANCE OF NEARSHORE TRACE^FOSSIL ASSEMBLAGES OF THE CAMBRO-ORDOVICIAN DEADWOOD FORMATION AND ALADDIN SANDSTONE, SOUTH DAKOTA Thomas M. Stanley’ Rodney M. Feldmann^ Research Associate, Section of Invertebrate Paleontology Abstract The Cambro-Ordovician Deadwood Form.ation and Aladdin Sandstone represent intertidal and sub- tidal, nearshore deposystems that contain few well-preserved body fossils, but contain abundant trace fossils. The present study uses the much neglected trace-fossil fauna to describe the diverse paleoen- viroemeets represented in the Deadwood-Aladdin deposystems, and to better understand the environ- mental conditions that controlled benthic life in the Early Paleozoic. The Deadwood-Aladdin ichnotaxa can be separated into three distinct assemblages based on the changing sedimentologic and hydrodynamic conditions that existed across the Cambro-Ordovician shelf. Trace-fossil assemblages and corresponding lithofacies characteristics indicate that the Dead- wood-Aladdin deposystems formed within an intertidal-flat and subtidal-shelf environment. Based on the distribution and numbers of preserved ichnotaxa, the intertidal flat can be subdivided further into an ecologically stressful inner sand-flat environment, and a more normal marine outer sand-flat environment, both of which belong to a mixed, SkoUthos-Cruziana softground ichnofacies. The inner sand flat is characterized by low diversity, low numbers, and a general lack of complexly constructed ichnotaxa. Trace fossils common to both assemblages tend to be smaller in the inner flat compared to the outer sand flat. Taphonomic effects, such as substrate type and sediment heterogeneity, also aid in differentiating between the inner and outer sand-flat assemblages. The subtidal shelf environment is categorized in the Cruziana ichnofacies. Ichnological evidence of periodic tempestite deposition and hardground development within this subtidal regime is mani- fested by high diversity and low abundance of ichnogenera. Key Words: trace fossils, ichnofossils, Cambrian, Ordovician, South Dakota, ichnofacies Introduction and Historical Perspective The Deadwood Formation and Aladdin Sandstone comprise all the Upper Cam- brian and most of the lowermost Ordovician rocks in western North Dakota and South Dakota, and eastern Montana and Wyoming. Surface exposures are limited to the Black Hills region of western South Dakota and eastern Wyoming, where they form an elliptical outcrop pattern within the interior of the Black Hills uplift (Fig. 1). The Deadwood Formation was formally named by Darton (1909) for exposures at Deadwood, South Dakota. The stratotype included the Deadwood Formation, Aladdin Sandstone, and basal lithotypes of what is now known as the lowermost Ordovician Winnipeg Formation (Fig. 1, 3). Because of the paucity of body fos- sils, Darton (1909) could only ascribe a Cambrian age to the type section. With continued paleontologic study of enclosed trilobites, the “Deadwood Formation” ' Kansas Geological Survey and the University of Kansas, 1930 Constant Avenue, Campus West, Lawrence, Kansas 66047. 2 Department of Geology, Kent State University, Kent, Ohio 44242. Submitted 17 January 1996. 1 Bridal Veil Falls 2 Annals of Carnegie Museum VOL. 67 Z UJ c ” 5 c m lo ® H Sco P OJ _ d o m o IS Q Z UJ ? CO I - "D 5 <0 O ^ c o I- ? ^ ^ lAJ lO ® DC ® . TO O O -J CO " > CD CO 1998 Stanley and Feldmann — Trace-fossil Assemblages 3 was given a Late Cambrian age by Darton and Paige (1925). Later, Furnish et al. (1936) separated the overlying Winnipeg Formation from the Deadwood For- mation based on the presence of Early Ordovician conodonts. Subsequently, Loch- man and Duncan (1950) found Early Ordovician trilobites in the upper intraclastic limestones in several “Deadwood” sections. Because no obvious unconformity exists between Cambrian and Ordovician strata, the Deadwood Eormation was reassigned a Late Cambrian-Early Ordovician age. The current stratigraphic pic- ture was completed by McCoy (1952) who designated the upper 4.4 m of the Deadwood Eormation as the Aladdin Sandstone based on the presence of numer- ous SkoUthos burrows. Since its first description, investigators of the Deadwood Eormation have com- mented on the abundant trace fossils that occur within the unit (Darton, 1909; Darton and Paige, 1925). Although trace fossils represent the dominant faunal evidence within the Deadwood Formation, paleontological and paleoecological studies have been limited to the meager occurrences of trilobites and inarticulate brachiopods (Lochman-Balk, 1964, 1970, 1971). Only one taxonomic study has dealt directly with the Deadwood-Aladdin trace fossils, and that was the desig- nation of a new ichnogenus, Ixalichnus, by Callison (1970). Consequently, this present work addresses not only the systematic ichnology, but also the paleoen- vironmental and paleoecological aspects of the Deadwood-Aladdin ichnofauna. The institutional abbreviation used in text is KSU, Kent State University, Kent, Ohio. Regional Paleogeography Throughout most of the Late Cambrian and Early Ordovician, two major facies belts developed across the cratonic shelf (Palmer, 1960; Fig. 2). In the region of the Black Hills, the Deadwood Formation and Aladdin Sandstone were deposited within the inner detrital belt, representing marginal-marine deposition. The trans- continental arch was the dominant tectonic element during much of the Paleozoic, and greatly influenced the type and distribution of the Deadwood-Aladdin de- posystems (Lochman-Balk and Wilson, 1967). As with many deposits of this time, initial submergence of the Cambrian shelf in the Black Hills area occurred on Precambrian basement composed principally of schists and metaquartzites. To- pographic relief on the basement surface consisted of metaquartzite monadnocks averaging no more than 30 m high. During the initial marine transgression into the Black Hills region these monadnocks became sources of metaquartzite boul- ders deposited as the basal conglomerate of the Deadwood Formation (Lochman- Balk and Wilson, 1967). Continued encroachment of the Cambro-Ordovician seas eastward was largely impeded by the transcontinental arch, which also supplied most of the terrigenous sediment of the inner detrital belt. The Black Hills area also was undergoing continual epeirogenic uplift throughout the Paleozoic, and may have represented a distal arm of the transcontinental arch (Cries, 1975). Fig. 1. — Generalized geologic map of the northern Black Hills in western South Dakota and eastern Wyoming. Outcrop pattern of the Deadwood Formation and Aladdin Sandstone shown as black band bounded by Precambrian basement and Lower Paleozoic carbonates. Location of measured sections are indicated by the white circles, and township and range coordinates for sections are given in lower left of this figure. 4 Annals of Carnegie Museum VOL. 67 Outer detrital ■ Middle carbonate belt □ Inner detrital belt Transcontinental arch MILES 0 300 j— — J 0 300 KM A A ' WESTERN MONTANA BLACK HILLS PERITIDAL CARBONATE RESTRICTED COASTAL SAND AND ALGAL BANKS SHELF FLATS Fig. 2. — Paleogeographic map of the Cambro-Ordovician craton showing relative positions of the shelf edge and transcontinental arch, and lateral distribution of major facies belts. Cross-section A- A' shows basic depositional environments across the cratonic shelf, superimposed with Irwin’s (1965) model of epeiric sedimentation, and distribution of energy zones in nearshore cratonic settings. The high energy zone in Irwin’s (1965) model corresponds to lithofacies 2 and 4 of the Deadwood For- mation and Aladdin Sandstone. Small circles on cross section represent zone of effective wave base. Diagram modified from Palmer (1960) and Sepkoski (1982). 1998 Stanley and Feldmann— Trace-fossil Assemblages 5 Continual epeirogeeic upwarping is suggested by a general thinning of Paleozoic strata as they onlap the Black Hills region (Gries, 1975; Lisenbee, 1975). Farther to the west, rocks characteristic of nearshore coastal deposystems of the Dead- wood Formation grade into restricted subtidal-shelf deposystems of the middle carbonate belt, represented by Dead wood time-equivalent units of the Du Noir and Emerson formations of Wyoming and Montana (Miller, 1936; Sepkoski, 1982; Fig. 2). Both formations are composed of thick shale and intraclastic limestone sequences, lithologically similar to the middle parts of the Deadwood Formation (Fig. 3), and indicate that shelf-like conditions extended into the Black Hills during times of maximum transgression. However, open-ocean circulation across the shelf was restricted by algal buildups along the shelf margin (Sepkoski, 1982; Fig. 2). West of the algal banks, restricted shelf sedimentation of the middle carbonate belt abruptly grades into outer-shelf and shelf-slope sedimentation of the outer detrital belt (Palmer, 1960). Deadwood and Aladdin Lithofacies In the northern Black Hills (Fig. 1), five lithofacies were deposited throughout Cambro-Ordovician time in this region. These are: 1) irregularly bedded hematitic sandstones, 2) interbedded sandstone and siltstones, 3) interbedded shale and in- traclastic limestones, 4) crossbedded sandstones, and 5) hematitic sandy lime- stones (Fig. 3, 4; Table 1). Irregularly Bedded Hematitic Sandstone Lithofacies (Lithofacies ij.- — Litho- facies 1 is characterized by dark reddish brown, medium-grained sandstone, with minor reddish green shale partings at the base. Bedding thickness is highly vari- able and irregular, ranging from thin-bedded (3-“5 cm) to thick-bedded (25-66 cm) intervals. Bed thickness increases toward the top of the lithofacies. Primary sedimentary structures are rare within thick-bedded strata, but consist of multi- directional ripple marks and small-scale crossbed sets in the thin-bedded intervals. Glauconite is a minor constituent of the sandstone although hematite is rather abundant, giving the facies its characteristic red color. Dolomite and calcite al- ternate as the main cements, with calcite being predominant in the basal parts of the lithofacies. Interbedded Sandstone and Siltstone Lithofacies (Lithofacies 2j.— Lithofacies 2 is composed predominantly of light-colored, interbedded fine-grained sandstone and siltstone beds that are 0.5~-2.0 cm thick. Current laminations, wavy crossbed- ding showing multiple current directions, parting lineations (Fig. 7D), and current- rippled bedding surfaces are common. Mudcracked surfaces also occur on some of the siltstone beds. Subordinate lithologies include shale as thin partings and clay drapes between sandstone and siltstone beds, which produced a flaser bedding texture, and fine-crystalline argillaceous limestone intervals normally found in the basal and upper parts of the lithofacies. Distinct grains or lenticular patches of glauconite are common within the sandstones and siltstones. Calcite is the dom- inant cement throughout. This facies represents the best exposed and most easily recognizable lithotype of the Deadwood Formation, and totally incorporates the lower 2.4 m of the overlying Aladdin Sandstone (Fig. 3). Interbedded Shale and Intraclastic Limestone Lithofacies (Lithofacies 3).- — Lithofacies 3 is composed of thin-bedded (3-10 cm), lenticular intraclastic lime- stone interbedded with thick (1-7 m) intervals of fissile shale, and thin-bedded (1-30 cm), finely crystalline to micritic limestone. Intraclasts are normally oblate 130 120 110 100- 90- 80 7 0- 60- 50- 4 0- 30 20 1 0- Annals of Carnegie Museum VOL. 67 (K) Archaeonassa 0-0 Arthraria ^ 1 Arthrophycus ^ Crossopodia Cruziana Didymaulichnus (2 GyroUthes Palaeophycus Planolites HI Psammichnites ^ Rusophycus ^ IScoUcia I Skolithos Taenidium 7Teichichnus O Thalassinoides Trichophycus ■77 Uchirites — f- Branched Burrow H* Ribbed Trail O Gently Curved Trace LITHOLOGY tto>| Conglomerate EI3 Sandstone ES Calcareous Sandstone [5^ Dolomitic Sandstone Limestone Dolomitic Limestone Argillaceous Limestone Sandy Limestone ^§lntraclastic Limestone Siltstone Shale Glauconite TEXTUP-E/BEDDINQ Massive/Thick Bedded Thin/Laminated Bedding Wavy Bedded Irregular Bedded Ke>I Lenticular Bedded Cross-Bedding [77?] Coarse/Medium-Grained Fine-Grained [V® Skolithos Burrows 1998 Stanley and Feldmann — Trace-fossil Assemblages 7 to tabular in shape, and set within a fine-grained crystalline or micritic limestone matrix. No current orientation of the intraclasts was evident. Upper surfaces of the intraclastic limestones commonly were cracked, pitted, and accompanied by protrusion of intraclasts above the bedding surface, suggesting possible hard- ground or firmgroued development from early subaqueous cementation (Leeder, 1982:291), or from exposure of partially lithified sediments from periodic storm erosion (Bromley, 1990:19). Lower contacts of intraclastic limestones were curved or appeared erosive into the underlying shale lithologies. Shale intervals are composed of gray, black, green, and sometimes purple calcareous fissile (1- 4 mm thick) shale, along with subordinate amounts of lenticular-bedded calcar- eous siltstone. Crossbedded Sandstone Lithofacies (Lithofacies Lithofacies 4 is composed of a basal trough crossbedded sandstone and conglomerate facies deposited in a high-energy, nearshore beach or barrier island system, that encompasses the basal 10 m of the Deadwood Formation, and supports a meager and poorly preserved trace-fossil assemblage consisting of Skolithos and scant PlanoUtes{l) burrows. This lithofacies also characterizes the upper 1.5-2 m of the Aladdin Sandstone, which exhibits the typical Skolithos ichnofabric of lower Paleozoic orthoquartzites (Droser, 1991; Droser and Bottjer, 1993). Because of the density of the Skolithos burrows, this upper part of the Aladdin has an average ichnofabric index of 4 based on Droser and Bottjer (1993). Both the sedimentological and ichnological nature of lithofacies 4 support a high-energy interpretation for this lithofacies (Droser and Bottjer, 1989; Bockelie, 1991; Droser, 1991). Hematitic Sandy Limestone Lithofacies (Lithofacies 5j.— Lithofacies 5 is com- posed of a wavy bedded, very argillaceous to sandy, hematitic, crystalline lime- stone that contains no trace fossils. Laterally, this lithofacies is discontinuous, but is always associated with lithofacies 1 or 2, and is interpreted to have formed on an intertidal carbonate flat in areas undergoing relatively low siliciclastic influx (Fig. 4). Because of the importance of their enclosed trace fossils, only lithofacies 1, 2, and 3 will be discussed in detail in the following sections. All three of these main lithofacies types fall within the typical Seiiacherian, marine softground ichnofacies (Bromley, 1990:215; Bromley and Asgaard, 1991). Systematic Ichnology Of the 18 formally designated ichnogenera and 27 ichnospecies collected from the Deadwood and Aladdin formations, ten ichnogenera are diagnostic for pa- leoenvironmental interpretations due to their restricted occurrences (Fig. 3, 16; Table 1). Ichnotaxa important in paleoenvironmental interpretations include Ar- chaeonassa, Skolithos, IScoUcia, GyroUthes, and Arthraria in differentiating outer sand-flat environments from inner sand flats and the subtidal shelf. The presence of certain ichnospecies, such as Planolites beverleyensis and Palaeophycus tub- ularis, can also aid in differentiating the inner sand flats from the outer sand flats. Fig. 3. — Composite weathered profile of the Deadwood Formation and Aladdin Sandstone showing lithofacies and stratigraphic distribution of the collected trace fossils. Numbers to left of section correspond to lithofacies types described in text. BRIDAL VEIL FALLS DEADWOOD LITTLE ELK CREEK 8 Annals of Carnegie Museum VOL. 67 1998 Stanley and Feldmann— Trace-fossil Assemblages 9 Ichnogenera such as Cruziana, Rusophycus, Crossopodia, and DidymauUchnus are more indicative of the sobtidal shelf environments. Ichnogenos Archaeonassa Fenton and Fenton, 1937 Archaeonassa fossulata Fenton and Fenton, 1937 (Fig. 5A-C) Material Examined. — Archaeonassa fossulata is one of the most common trac- es collected, being found at all three measured sections. Only two are illustrated showing the range in both morphology and preservation. A large slab (KSU 4556) with over ten individuals was collected at the Deadwood type locality. KSU 4597 was collected at the Little Elk Creek locality. Description. — Straight to gently curving, trace preserved in concave epirelief; width 2-5 mm, max- imum length 110 mm; trace consists of a smooth to transversely striated groove, bounded by variably defined lateral ridges; groove typically three-fourths total trace width, semicircular in cross section; bounding ridges smooth, ranging from sharp-crested to broadly-rounded semicircles in cross section; ridges are as high as groove is deep; trace may tangently intersect shallow, circular depression aver- aging 15 mm in diameter and no more than 5 mm deep. Discussion.— A much needed taxonomic review of Archaeonassa has been done by Buckman (1994). The Deadwood forms conform to two of Buckman's preservational variates, the smooth and standard type, of A. fossulata (Buckman, 1994:188, text-fig. 5 A, B2), with the smooth variate being the more common of the two forms. The ichnogenus has been attributed to the locomotor activities of gastropods, trilobites, and even irregular echieoids (Fenton and Fenton, 1931, 1937; Buckman, 1994). Concerning the Deadwood material, echinoids may be excluded as potential trace makers, because they did not evolve until the Late Ordovician. Although trilobites cannot be entirely excluded as possible excavators of the Deadwood Archaeonassa, they seem unlikely given that A. fossulata is a true epirelief trace in construction (Buckman, 1994). Trilobite repichnion, such as Cruziana, are rarely preserved as epireiiefs (Seilacher, 1970; Goldrieg, 1985). Gastropods seem to be the best candidates as trace makers, based on observations of trails left by the Recent snails Littorina and Ilyanassa on intertidal and subtidal flats in Washington and California (Fenton and Fenton, 1931, 1937). Important to this interpretation is that the Recent snails would occasionally excavate a small circular depression, terminating the epichnial groove. The Deadwood specimens of Archaeonassa imply a similar behavior as some of the trails terminate at shal- low burrows (Fig. 5B), and continue on the reverse side of the bed (Fig. 5C). Similar epichnial grooves associated with circular depressions have been de- scribed as ‘'annelid trails” by Hall (1852:pL 14, fig. 3). These forms have sub- sequently been assigned to A. fossulata by Buckman (1994). If HalFs specimens from the Silurian of New York are truly conicheogeeeric with the Deadwood forms of Archaeonassa, then both represent the only recorded specimens exhib- iting this particular behavior in the ichnogenus. All previously recorded occur- rences of Archaeonassa were collected from intertidal deposits (Buckman, 1992, Fig. 4. — Cross section from the Little Elk Creek section in the east to the Bridal Veil Falls section in the west, illustrating vertical and lateral distribution of Deadwood Formation and Aladdin Sandstone paleoeevironments, along with corresponding Seilacherian, marine soft-ground ichnofacies. Horizontal line (“T”) represents hypothetical time line used in Figure 16. Table I. — Summary of major lithofacies, associated trace fossils, and interpreted environments of deposition of the Deadwood Formation and Aladdin Sandstone. Under the column headed by “Trace Fossil Assemblages, “ D. T. = Dominant Trace, and S. T. = Subordinate Trace. Ethological interpretations of trace fossils from Frey and Pemberton (1984, 1985). 10 Annals of Carnegie Museum VOL. 67 s a • i: (u > T3 c W gj s s "C O a. > X .. 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'I o o g -S PQ °t i.s ^ ^ c 0 "S 3 1 ^ s ffl •O A Cj 5 £ ^ 00 s 0^ Q C •S b n) 3 5 .2 C S ^ "g ° ■| .1 ^ .J • : ^ o • ^ u Q ^ I" a o ^ S., O' o s C 4- -2 o G s .2 -a Q S G ^.2 rs S ^ Q "'3 'S £ g| 2 'a o — c bfl •£ G O cd C flj •- 'G o- £ o ^ c 4-1 O O -.G i-l rS .2 'w c o £ a o > >:.G o w o ,s g 8 ts ~o flj -o ^ S 0) 'e 2 cd w -a •a "O D B. e ’fi wo § 2 S 2 t y c HH A O >. .3 ed o w O ^ 'o o 10 cd .. e . b -d . cc ^ c D D dj Q cd ?P> § G ^ .3 ^ 2 ..2 cfl O w e c£) 3 u .2 « 2 3 fi « ^ E ‘3 o S Q £ o -d .§ « — 2 ^ -P cd ■S 'd cd « w e o o A ^ cS "O o S fH Cd •S w Fodinichnion of Thalassino- ides, Planolites, Taenidium, and “Gently Curved Trace.” 1998 Stanley and Feldmann — Trace-fossil Assemblages 11 ^ C3 u E E i ■5 3 (U O w « g X) b «3 >.x bO 3 U C W) > u > v o o .£ 2) G o C« 3 0) £ tu O o -o e w C/1 cd e 3 £ o 2 cA 2 O 0) o 2 o o C/3 <3J « d o £ M 3 o o > X .£ 2 (U 2 o X H 2 W) c u S c/3 3 X 2 £ 2 E O X "2 .s 5.— KSU 4671 was collected 29 m above the base of lithofacies 3. KSU 4542 was collected 1.5 m above the base of lithofacies 2. IRusophycus ichnospecies (Fig. 12C) Material Examined.— A single specimen (KSU 4669) was collected as float at the Little Elk Creek locality. 30 Annals of Carnegie Museum VOL. 67 Fig. 12. — A, B. IScolicia ichnospecies. A. KSU 4590, concave epireiief, lithofacies 2, Little Elk Creek section. B. Note arcuate ridges leading to distinct hypichnial ridge; KSU 4681, convex hyporelief, lithofacies 2, Deadwood type section. C. IRusophycus ichnospecies, KSU 4669, convex hyporelief, lithofacies 3, Little Elk Creek section. D. SkoUthos ichnospecies, typical form found in upper 2 m of the Aladdin Sandstone; KSU 4677, Deadwood type section. Bar scales represent 1 cm. 1998 Stanley and Feldmann-— Trace-fossil Assemblages 31 Description.-— Biloh&d trace preserved in convex hyporelief; width 17 mm, tapering to 15 mm at posteriori?) end; incomplete length. Lobes parallel, approximately one-half total trace width; separated by shallow medial groove that varies between 2-3 mm wide; groove distinct posterioriy(?), indistinct anteriorlyC?). No surface striae evident. In transverse section trace extends 5 mm above bedding; lobes quadrate in cross section, flat crested, and form a steep angle to bedding surface. Discussion,— this specimen is incomplete and poorly preserved, it is uncertain whether it represents a true Rusophycus or a broken Cruziana. Although collected as float, the slab on which this specimen is pre- served is similar to the thin, lenticular siltstone beds described from lithofacies 3. Ichnogenus Scolicia De Quatrefages, 1849 IScolicia icheospecies (Fig. 12A, B) Material Examined.— Two specimens on a single slab (KSU 4590) and another specimen (KSU 4681) were collected from the Little Elk Creek locality. Speci- mens KSU 4557 and 4572 were collected from the Deadwood type locality. Description.—SmglG, large, deep furrow that may become weakly bilobed along its length; pre- served in concave epirelief or convex hyporelief; width varies from 10-17 mm, averages 14 mm, remaining constant along trace length within individuals; length variable, maximum measured at 175 mm; furrow floor smooth (KSU 4590; Fig. 12A), composed of a finer-grained material than host matrix that may represent a lining. KSU 4681 (Fig. 12B) represents a single hypichnial ridge that becomes weakly bilobed at terminations; surface of furrow ornamented with fine, arcuate to straight transverse ridges that do not extend to trace margins; ridges grouped five to seven per centimeter. In transverse section epichnial furrow and hypichnial ridge has a quadrate cross section; depth or height of main structure ranges from 5-7 mm. Discussion.— Th&^& specimens were originally assigned to PalaeobulUa Got- zinger and Becker based on similar morphology and mode of preservation with Pennsylvanian forms from Tennessee (Miller and Knox, 1985), and because of similarities with modern gastropod epichnia (Knox and Miller, 1985). Uchman (1995) has subsequently synonymized PalaeobulUa, Taphrhelminthopsis, and Subphyllochordia into Scolicia. Most of these other forms, particularly Palaeo- bulUa and Taphrhelminthopsis, represent preservational variates of washed-out Scolicia. TwO' of the Deadwood specimens do have the appearance of postcon- straction erosion similar to that illustrated by Uchman (1995:text-fig. 2A, B; Fig. 12A). Uchman (1995) also suggests that similar washed-out traces (Scolicia stroz- zii) are possible preservational variates of Curvolithus or Cruziana. Neither Cur- voUthus nor Cruziana appear to be good candidates for the Deadwood specimens in that they represent a single hypichnial ridge or epichnial groove that is some- times preceded by arcuate transverse ribs representing backfilling of the burrow (top half of Fig. 12B). The specimen in Figure 12D superficially resembles Bea- conites illustrated in Hantzschel (T975:pL 28, fig. 1). Beaconites, however, is characterized by a distinct wall enclosing the backfill menisci (D’ Alessandro and Bioinlcv. 1987; Keighley and Pickerill, 1994). The Deadwood specimens do not exhibit a wall structure. All specimens from the Deadwood resemble in gross morphology epichnial or hypichnial expressions of washed-out Scolicia with poor- ly developed sediment strings along the bottom of the burrow. The lack of bilobate or trilobate morphology of the Deadwood traces is the reason only tentative as- signment to Scolicia is made. KSU 4557 and 4572 were collected 3.5 m above the base of litho- facies 2. KSU 4590 and 4681 were collected 4.5 m above the base of lithofacies 2. 32 Annals of Carnegie Museum VOL. 67 Ichnogenus Skolithos Haldeman, 1840 Skolithos ichnospecies (Fig. 7D, 8E, 12D) Material Examined. were collected from all localities and from most lithofacies except for lithofacies 3. Abundant Skolithos burrows are the most distinct feature of the upper 2 m of the Aladdin Sandstone. Description. — Specimens on a large block (KSU 4677; Fig. 12D) are narrow, uebranched, vertical to slightly inclined shafts preserved in full relief; shafts vary between 1-2 mm in diameter, remaining constant in individuals; normally straight, but may curve slightly at distal portions of tube; length variable, maximum observed at 45 mm; walls unornamented. Specimens on slabs KSU 4586 and 4676 (Fig. 7D, 8E) are narrow vertical shafts, having circular to oval cross sections; shafts preserved in convex or concave hyporeiief, concave epirelief, or full relief; shaft diameters range from 3-8 mm, average 5 mm, length incomplete; tube usually filled with sediment creating raised pimples in hy- poreiief and shallow depression in epirelief. Discussion.— Som& of the collected specimens of Skolithos from the Aladdin Sandstone (KSU 4677; Fig. 12D) appear conichnospecific to Skolithos verticalis (Alpert, 1974). The other specimens of Skolithos (KSU 4586 and 4676; Fig. 7D, 8E) are incomplete, but judging by their cross-sectional dimensions, may belong to S. linearis (Alpert, 1974). Skolithos is common throughout the outer sand-flat lithofacies (lithofacies 2) and ofUshore parts of the inner sand-flat lithofacies (lithofacies 1). It is the most diagnostic feature of the upper 2 m of the Aladdin Formation, making the Aladdin one of the most easily recognizable and mappable formations in the Black Hills. Facies. — Specimens collected and described from the Aladdin Sandstone (KSU 4677) occur within lithofacies 4 and represent a high-energy beach or barrier island complex. Other collected specimens usually occur in groups of ten or more individuals on a single slab and include KSU 4586 and 4676, collected 5 and 6.5 m, respectively, above the base of lithofacies 2 at the type section. At the Little Elk Creek section: KSU 4679 and 4681 were collected 3.3 and 4.5 m, respectively, above the base of lithofacies 2; KSU 4682 was collected 5.3 m above the base of lithofacies 1 , Ichnogenus Taenidium Heer, 1877 Taenidium serpentinum Heer, 1877 (Fig. 13 A) Material Examined.— Two specimens occur on a single slab (KSU 4544) col- lected from the Bridal Veil Falls locality. Description. — Narrow, unlined, linear trace preserved in convex hyporeiief; width 2 mm, maximum length 126 mm; trace margins slightly amiulated; trace interior with numerous, regularly spaced, fine, transverse arcuate menisci that represent backfilling of burrow; menisci average 1.6 mm thick in longitudinal direction; grouped 16 menisci per centimeter. In transverse section burrow has semicir- cular to quadrate cross section; height no more than 2 mm above bedding. Discussion.— The presence of evenly spaced, gently arcuate menisci that are about as thick as the burrow is wide, and lack of a wall support assignment of these specimens to Taenidium serpentinum (D’ Alessandro and Bromley, 1987; Keighley and Pickerill, 1994). Facies.— The specimens were collected 4.8 m above the base of lithofacies 3. 1998 Stanley and Feldmann — Trace-fossil Assemblages 33 Fig. 13. — A. Taenidium serpentinum Heer, KSU 4544, convex hyporelief, lithofacies 3, Bridal Veil Falls section. B, C. ITeichichnus ichnospecies. B. Full-relief partial burrow intersecting bedding on the left of figure; KSU 4574, lithofacies 2, Deadwood type section. C. Same specimen in transverse profile, note poorly preserved spreite. D. Thalassinoides ichnospecies, main burrow labeled as “M,” secondary branches labeled with an “S”; KSU 4589, full relief, lithofacies 3, Little Elk Creek section. Bar scales represent 1 cm. 34 Annals of Carnegie Museum VOL. 67 Ichnogenus Teichichnus Seilacher, 1955 ITeichichnus ichnospecies (Fig. 13B, C) Material Examined. — Specimen KSU 4574 was collected at the Deadwood type section, KSU 4682 from the Little Elk Creek locality. Description. — Cylindrical burrow oriented parallel to bedding, except at end where it gradually deflects into bedding, preserved in full relief; width 15 mm, remaining constant along burrow length; incomplete length of 55 mm; retrusive spreite arranged as stacked gutters, composed of 0.5-1 mm- thick packages of fine-grained sand and silt, oriented concave-up in cross section and intersect burrow floor at 10-15° in profile; burrow walls ornamented with fine, longitudinally directed striae, formed as exterior expression of the spreite. Discussion. — There is some possibility of misidentification of incomplete bur- rows of Teichichnus. It has been noted that Teichichnus may be transitional to Rhizocorallium (Chisholm, 1970), Ophiomorpha (Hester and Pryor, 1972; Frey et al., 1978), and Thalassinoides (Frey and Seilacher, 1980). None of these other ichnogenera were collected from the Deadwood Formation, except for one pos- sible specimen of IThalassinoides from the Little Elk Creek section. Because of the incomplete nature of the collected specimens only a tentative assignment is given to the Deadwood forms. Facies. — KSU 4574 was collected 3.5 m above the base of lithofacies 2. KSU 4682 was collected 8.7 m above the base of lithofacies 1. Ichnogenus Thalassinoides Ehrenberg, 1944 Thalassinoides ichnospecies (Fig. 13D) Material Examined. — A single specimen (KSU 4589) was collected from the Little Elk Creek section. Description. — Cylindrical-shaped burrow system preserved in full relief; burrows oriented horizon- tal to bedding; incomplete length of main burrow 30 mm; diameter of main burrow 15 mm except at burrow junctions, which swell to 19 mm; secondary burrows slightly smaller than main trunk, diameter 10 mm, oriented at 90° to main burrow; main trunk and secondary bifurcations show regularly spaced constrictions producing an annulated burrow margin; constrictions transverse, slightly arcuate, ranging between 2-5 mm thick in longitudinal direction, grouped two to three annuli per centimeter; internal composition of burrow system different than surrounding matrix. In transverse section burrow cross section oval, with long axis of ellipse parallel to bedding; faint, transverse laminations present, sug- gesting burrow collapse; bedding of host matrix near burrow system highly disturbed, with top of burrow showing gradational contact with host lithology. Discussion. — Assignment of this specimen to Thalassinoides ichnospecies is based on the branching character and slight swelling of the burrow at branch junctions. Burrow junctions of the Deadwood specimen, however, are at nearly 90°, which is atypical of Late Paleozoic and younger forms assigned to this ichno- genus (Myrow, 1995). However, similar T-shaped, rather than Y-shaped, bifur- cations have been reported by Howard and Frey (1984), Bromley (1990:159), Maples and Suttner (1990), Myrow (1995), and Uchman (1995). Until recently, Thalassinoides had not been recorded from units older than the Ordovician (Shee- han and Schiefelbein, 1984; Bromley, 1990:159; Myrow, 1995). Myrow (1995), however, has identified a new ichnospecies, T. horizontalis, from the Upper Cam- brian Peerless Formation of Colorado. There are also possible occurrences of Thalassinoides in rocks as old as the Lower Cambrian (Droser and Bottjer, 1988). Similarities between the Deadwood Thalassinoides and T. horizontalis consist of 1998 Stanley and Feldmann — Trace-fossil Assemblages 35 a burrow system oriented parallel to bedding with no vertical shafts, burrow junc- tions are mostly T-shaped and show little to no swelling, lack of definite scratches along burrow walls, and infilling material that is arranged as spreite that is com- positionally different from the host matrix. The Deadwood specimen, however, is larger and exhibits slight constrictions along the margins of interjunction seg- ments. Facies. — This specimen was collected 3 m above the base of lithofacies 3. Ichnogenus Trichophycus Miller and Dyer, 1878 Trichophycus pedum (Seilacher, 1955) (Fig. 14A) Material Examined. — ^Two specimens (KSU 4554, 4578) were collected from the Deadwood type section. A third specimen (KSU 4543) was collected at Bridal Veil Falls. Description. — Trace preserved in convex hyporelief; consisting of a linear main trunk, ranging from 3-5 mm wide; incomplete length of 65 mm; stout secondary branches arise from midline of main trunk, loop around, and extend upward into host bed; exposed parts of secondary branches not more than 4 mm wide and 6 mm long. Lateral margins of main trunk with faint undulations; margins of secondary branches straight. Discussion. — Geyer and Uchman (1995:185-187, fig. 5. 3-5. 5, 6, 7. 1-7.9) have recently reassigned this form to Trichophycus Miller and Dyer, based on differing behavioral patterns between this ichnospecies and other forms of Phycodes. Tri- chophycus pedum consists of a straight to gently winding main burrow oriented horizontal to bedding, with a variable number of secondary lateral probes ex- tending obliquely off the main burrow into the host bed. Geyer and Uchman (1995) note that T. pedum exhibits variable morphology that ranges from straight to curved main tunnels, coupled with different densities of the secondary probes. The Deadwood forms show a linear tunnel with sparse secondary probes, which closely resemble T. pedum collected from the Lower Cambrian Candana Quartzite of Spain (Crimes et al., 1977:fig. 7a, b), and a few forms collected from sandstone facies of the Lower Cambrian Gross Aub Formation (Geyer and Uchman, 1995: fig. 7.7). The prime similarity is the straight character of the main tunnel in both the Deadwood and Spanish material compared to the more common, gently curv- ing to palmate morphology represented by material from the Lower Cambrian of Pakistan (Seilacher, 1955), other forms from the Nama Group (Geyer and Uch- man, 1995), ?Cambro-Ordovician Kellys Island and Little Bell Island formations (Fillion and Pickerill, 1990), Middle Cambrian Oville Formation of northern Spain (Legg, 1985), Lower Cambrian Breivik Formation of Norway (Banks, 1970), and the Lower Cambrian Random Formation of Newfoundland (Crimes and Ander- son, 1985). The straight morphology may be a function of the coarse-grained texture of the hosting sediment, as well as the environment of deposition under which these variates were constructed. The Candana Quartzite and the hosting lithofacies of the Gross Aub and Deadwood formations are medium- to coarse- grained sandstones in association with desiccation cracks that formed under very shallow, intertidal conditions. Facies. — Specimen KSU 4554 was collected 2.5 m above the base, and KSU 4578 was collected 5.9 m above the base of lithofacies 1. KSU 4543 was collected at the base of lithofacies 2. 36 Annals of Carnegie Museum VOL. 67 Fig. 14. — A. Trichophycus pedum (Seilacher), KSU 4554, convex hyporelief, lithofacies 1, Deadwood type section. B, C. Uchirites ichnospecies. B. Full-relief burrow shown from above, note sharp ridge at apex of burrow; KSU 4573, lithofacies 2, Deadwood type section. C. Same specimen seen in cross section, note blunt lower apex of burrow. D. “Branched Burrow,” KSU 4562, convex epirelief, lith- ofacies 2, Deadwood type section. Bar scales represent 1 cm. Ichnogenus Uchirites Macsotay, 1967 Uchirites ichnospecies (Fig. 14B, C) Material Examined. — A single specimen (KSU 4573) collected at the Dead- wood type locality. Description. — Straight tube preserved in full relief; width 10 mm; incomplete length of 15 mm; surface of tube ornamented with faint grooves running parallel to long dimension of trace; trace cross section almond-shaped, with sharp upper apex and blunt lower apex; trace interior hollow, consisting 1998 Stanley and Feldmann — Trace-fossil Assemblages 37 of a circular tube encased in a sheath-like envelope; envelope composed of a coarser-grained material than host lithology. Discussion. — A diagnostic character of Uchirites is its almond-shaped cross section, consisting of a sharp upper apex and blunt lower apex. The burrow typ- ically consists of an inner structure with a smooth external surface surrounded by an outer structure having the appearance of a sheath (Chamberlain, 1971:231). In well-preserved specimens this outer sheath typically is ornamented by fine striae obliquely oriented to the midline. Although the Dead wood specimen lacks a stri- ated surface, it shows sufficient morphological characters to be classified under Uchirites. Facies. — This specimen was collected 7 m above the base of lithofacies 2. “Branched Burrow” (Fig. 14D) Material Examined.— KSU 4562 was collected from the Deadwood type section. Description. — Narrow, unornamented, branched-burrow system preserved in convex epirelief; main burrow 3.5 mm in diameter; second-order branches 2 mm in diameter, typically at 90° to main burrow; third-order burrows typically 1-1.5 mm in diameter; making a 20-25° angle with second-order bur- rows. Discussion. — This trace vaguely resembles Chondrites Sternberg and Hartsel- lea Rindsberg. Differences between it and Chondrites are the higher branch angles and the decrease in burrow diameter in the second- and third-order burrows seen in the Deadwood specimen. Hartsellea also represents a branching-burrow system like Chondrites, but individual burrows of Hartsellea have a distinctly lined wall and internal menisci (Rindsberg, 1994:47-50, pi. 9B, lOA, B, llA-D, 12C, D). The Deadwood specimen shows no lining or menisci fill. Facies. — This specimen was collected at the top of lithofacies 2. “Gently Curved Trace” (Fig. 15 A) Material Examined. — A single, incomplete specimen (KSU 4533) collected at Bridal Veil Falls. Description. — Smooth, narrow ridge preserved in convex epirelief. Ridge consistently 1.2 mm wide; circumference is 80 mm, incomplete; ridge gently curves to resemble a half circle. Discussion. — Because this specimen is incomplete no ichnogeneric designation is given, although it does resemble a partial Circulichnis montanus Vialov, 1971 (Hantzschel, 1975 :W52, pi. 31, fig. 4; Pickerill and Keppie, 1981:fig. 3). The trace does not appear to have been actively backfilled in that the filling material lacks distinct spreite and menisci, and is of the same composition as the host matrix. This excludes the form as a possible Planolites burrow based on taxo- nomic criteria (Pemberton and Frey, 1982; Keighley and Pickerill, 1995). The trace is also unlined, which would exclude it as a possible Palaeophycus (Pem- berton and Frey, 1982; Keighley and Pickerill, 1995). Facies.— Specimen was collected 1.8 m above the base of lithofacies 3. 38 Annals of Carnegie Museum VOL. 67 Fig. 15. — A. “Gently Curved Trace” shown by arrow, KSU 4533, lithofacies 3, Bridal Veil Falls section. B. “Ribbed Trail,” KSU 4573, concave epirelief, lithofacies 2, Deadwood type section. Bar scales represent 1 cm. “Ribbed Trail” (Fig. 15B) Material Examined. — A single specimen (KSU 4573) collected at the Dead- wood type section. Description. — Slightly curving furrow preserved in concave epirelief; furrow width approximately 20 mm; incomplete length 50 mm; furrow with regularly spaced transverse ridges; ridges consistently 2 mm thick in longitudinal direction, spaced 3 mm apart, and do not extend to lateral margins of furrow. Discussion. — The poor preservation of the specimen does not support a formal ichnogeneric designation. However, the trace looks very similar to the epichnial groove part of Plagiogmus Roedel (Glaessner, 1969:385, fig. 8). Facies. — Specimen was collected 7 m above the base of lithofacies 2. Results Aspects of Cambro— Ordovician Sedimentation Sedimentary structures consisting of bidirectional crossbedding, clay drapes, flaser, wavy, and current-rippled bedding, and the occurrence of mudcracks in 1998 Stanley and Feldmann — Trace-fossil Assemblages 39 siltstone and sandstone beds indicate that lithofacies 1 and 2 were deposited under intertidal conditions (Ginsburg, 1975:92; Klein, 1985). The fine-grained terrige- nous elastics intermixed with tempestite-derived intraclastic limestones in litho- facies 3 indicate that this unit was deposited further offshore on the Cambro- Ordovician shelf. The chaotic nature of the intraclasts suggests little postdeposi- tional reworking by tidal currents or waves, implying deposition below normal wave base, but not below storm wave base. The lenticular nature and curved lower contacts of the intraclastic beds suggest scouring of the substratum during tempestite deposition. A similar interpretation was made for Deadwood time- equivalent units in Montana (Emerson Formation) and Wyoming (Du Noir For- mation; Sepkoski, 1982), which also consist of interbedded shale and intraclastic limestone. Other published examples of Cambro-Ordovician siliciclastic tidal deposits in- clude the Eureka Quartzite of California and Nevada (Klein, 1975), the Monkman Quartzite of western Canada (Jansa, 1975), the Hickory Sandstone of central Texas (Cornish, 1986), the Theresa-March formations of New York and Ontario (Bjerstedt and Erickson, 1989), the upper Bell Island and Wabana groups of New- foundland (Fillion and Pickerill, 1990), and the Santa Rosita Formation of Ar- gentina (Mangano et al., in press). Except for the Bell Island and Wabana groups, all of these formations conform lithologically and texturally to the Deadwood- Aladdin tidal deposits. Similarity among these regionally separated deposits is illustrated by the lack of terrigenous mud in the upper intertidal to supratidal regimes. In contrast. Late Paleozoic (Carboniferous) to Recent tidal environments contain a predominance of mud in upper tidal facies (Ginsburg, 1975; Weimer et al., 1982; Klein, 1985); because tidal-flat facies models have been based on these Recent mud-dominated tidal sequences, recognition of Lower Paleozoic tidal de- posits have been difficult (Dott and Byers, 1980; Klein, 1980; Moiola, 1980). For example, units previously interpreted as simple blanket sands that formed under wave-dominated coastal systems such as the Middle Cambrian Flathead Sandstone of Wyoming and Montana, the Lower Ordovician St. Peter Sandstone of the mid- continent, and the Mt. Simon and Eau Claire formations of Wisconsin exhibit a greater complexity and intergradation of tidal and nonmarine fluvial sedimentation than previously reported (Dott and Byers, 1980; Moiola, 1980; Driese et al., 1981; Bjerstedt and Erickson, 1989). Lack of a dominant mud facies in these Cambro-Ordovician tidal orthoquartzite suites may be due to the extreme macrotidal range of the Cambrian epeiric seas, coupled with an absence of vascular plants. Given the low-lying topography of the Precambrian basement during the initial Cambrian transgressions onto the craton, frictional drag would have minimized wave processes while augmenting tidal ranges and currents in nearshore coastal settings (Irwin, 1965; Schopf, 1980; Hallam, 1981; see Fig. 2). Assuming horizontal time lines in Figure 4, a minimal breadth of the Black Hills Cambro-Ordovician shelf intertidal zone may have approximated 25-35 km, which far exceeds the 7-km average of modem macro- tidal coasts (Klein, 1980; Weimer et al, 1982). The closest modem analog that can be compared to the Deadwood-Aladdin intertidal zone is the Yellow Sea macrotidal flat of southwestern South Korea, with a tidal-flat width ranging from 8-25 km (Klein, 1980; Alexander et al., 1991). Current energy produced by tidal forces was probably fairly strong, especially across shelf areas within effective wave base. Lithofacies 2 is interpreted as forming in a moderate- to high-energy tidal regime due to the occurrence of parting lineations on some siltstone and 40 Annals of Carnegie Museum VOL. 67 sandstone bedding surfaces, and probably falls within the high-energy region of Irwin’s model for epicontinental seas (Irwin, 1965; Hallam, 1981; see Fig. 2). Current energy probably slackened over the inner parts of the tidal flats, exem- plified by lack of well-developed bedforms and tidal sedimentary structures in lithofacies 1, compared to lithofacies 2. Lack of fine-grained terrigenous elastics in lithofacies 1 suggests that tidal energy was strong enough to wash all but a coarser-grained sand fraction basinward. Transportation of muds basinward was also permitted by the lack of baffling by vascular plants in the upper intertidal and supratidal zones during the Early Paleozoic. Tidal deposits in Texas and New York also exhibit a coarsening of grain size in the upper peritidal to supratidal parts of the sand flats (Cornish, 1986; Bjerstedt and Erickson, 1989). Except for thin shale partings and clay drapes in lithofacies 2, the mud fraction by-passed the intertidal zones, and was deposited further offshore in lithofacies 3 (Fig. 16). Besides their trapping effects of fine-grained siliciclastics in upper tidal areas, the importance of vascular plants today can be inferred by changes in the type of physical and chemical weathering in terrestrial environments throughout geologic time. Without terrestrial plants, eolian processes probably dominated land areas in the Early Paleozoic, and without the stabilization effects of plants, would have deposited a greater amount of coarse-grained sand in nearshore marine environ- ments than observed today (Dott and Byers, 1980). At least for terrigenous tidal deposits, tidal sedimentation has changed since the Cambrian, and may be directly or indirectly related to the exploitation of the inner tidal zone by vascular plants, and strong tidal currents prevalent along ancient mesotidal coasts. The nature and trophic structure of tidal-flat benthic communities also must have changed with the evolution of vascular plants because of the indirect effect on substrate type within Cambro-Ordovician upper intertidal regimes. Modem macrotidal flats such as the Yellow Sea coast of South Korea and Gulf of Cali- fornia are characterized by large numbers of epifaunal, herbivorous grazers and deposit feeders in the inner tidal environments (Thompson, 1968, 1975; Frey et al., 1987; Fiirsich and Flessa, 1987). Herbivorous gastropods and bivalves dom- inate these modern peritidal and supratidal environments, feeding on algae and plant detritus deposited from suspension from waning tidal currents (Fiirsich and Flessa, 1987). Consequently, there is a preponderance of epifaunal repichnia- and infaunal vertical domichnia-type traces in Recent upper tidal regimes (Frey et al., 1987). Traces common to the upper tidal zones of the Deadwood Formation differ considerably from the modern counterparts (Table 1). Horizontal infaunal burrows are the most common traces found. These assemblages are dominated by Plan- olites, with minor occurrences of Palaeophycus, Trichophycus, and ITeichichnus. Although vertical domichnia burrows are present in the upper tidal flats, they are subordinate to the horizontal burrows. Also contrary to Recent tidal assemblages, repichnial surface trails are noticeably sparse in the upper sand flats of the Dead- wood (lithofacies 1). However, repichnion do increase in diversity and density in the outer sand-flat deposits (lithofacies 2). A similar situation occurs in the Cam- bro— Ordovician Marsh-Theresa tidal deposits as the epifaunal—infaunal ratio in- creases from peritidal to lower intertidal deposits (Bjerstedt and Erickson, 1989) and in the Ordovician Santa Rosita Formation of Argentina, which is characterized by a monoichnospecific Planolites montanus trace-fossil assemblage in the most shoreward tidal facies (Mangano et al., in press). 1998 Stanley and Feldmann- — Trace=fossil Assemblages 41 Trace-fossil Assemblages When coupled with the Deadwood and Aladdin lithofacies, the trace-fossil as- semblages provide a clearer picture of Cambro-Ordovician nearshore paleoenvi- ronments. The Deadwood^Aladdie trace fossils can be segregated into distinct assemblages related to the hydrodynamic and sedimentological conditions that persisted during the Cambro-Ordovician of the Black Hills. These assemblages also recur within their prescribed lithofacies during repeated shifting of strand lines (Fig. 4, 16). Two basic trace-fossil assemblages dominate: 1) an intertidal sand-flat assemblage that occurs in lithofacies 1 and 2, and 2) a restricted subtidal- shelf assemblage corresponding to lithofacies 3. Trace fossils also reflect sedi- mentologic and taphonomic regimes within the intertidal environment, and can be segregated into an inner sand-fiat assemblage (traces enclosed in lithofacies 1) and an outer sand-flat assemblage (traces enclosed in lithofacies 2). Table 1 sum- marizes the trace fossil distribution with corresponding lithofacies types. Inner Sand-flat Assemblage.— assemblage is characterized by very low ichnofaunal density and diversity, and occurs in lithofacies 1 and shoreward parts of lithofacies 2 (Table 1; Fig. 16). Ichnotaxa are dominated by Planolites. Pa- laeophycus, SkoUthos, Trichophycus, and ITeichichnus also occur, but are rare and normally found in the offshore parts of the assemblage (Fig. 3, 16). The paucity of trace fossils observed in these inner sand flats contrasts with modem tidal flats, which normally show low diversity but high density of traces within the inner-tidal regimes (Weimer et ah, 1982; Ekdale et ah, 1984:177; Frey et ah, 1987). Environmental parameters such as salinity, temperature, current en- ergy, and subaerial exposure probably varied greatly within these inner-tidal re- gimes as they do in modem environments, and probably influenced the number and distribution of the trace fossils. Also important are the potential taphonomic effects inherent in very nearshore marine settings (Bromley, 1990) in that the ichnological distinction between the inner and outer sand-flat assemblages may be as dependent on taphonomy, particularly depth of burrowing, as it is on the varying environmental conditions across the intertidal zone. Although there are noted exceptions in the depth and degree of bioturbation for Lower Paleozoic rocks (Miller and Byers, 1984; Sheehan and Schiefelbein, 1984), burrowing depths of the Cambro-Ordovician marine benthos were shallow (Ausich and Bottjer, 1982, 1990, 1991; Thayer, 1983; Bottjer and Ausich, 1986) and confined to the uppermost mixed-layer tiers (Bromley, 1990:128; Bromley and Asgaard, 1991). Shallow trace construction is also common in modem inter- tidal environments (Frey et aL, 1987). Added to the preponderance of shallow traces, Frey et al. (1987) noted that few of these epibenthic traces in the shoreward parts of the Yellow Sea tidal flats would have little potential for being preserved. It stands to reason that most biogenic activity in the Deadwood inner sand-flat regime stood a fairly poor chance of being preserved as well. The prevalence of disrapted bedding associated with the Deadwood inner sand-flat trace-fossil as- semblages may indicate that intense bioturbation occurred without subsequent preservation of discrete, recognizable trace fossils. When comparing the type and distribution of ichnotaxa between the inner and outer sand-flat assemblages, ich- nofaunal differences are not as much a change in ichnotaxa, but an exclusion of inferred, shallowly constructed biogenic stmctures. Full-relief burrows of Plan- oUtes, Palaeophycus, and Skolithos are the most common traces in the inner sand- 1998 Stanley and Feldmann — Trace-fossil Assemblages 43 flat assemblage. It is only toward the more offshore parts of the assemblage that shallower trace fossils such as Archaeonassa are preserved (Table 1; Fig. 3, 16). Substrate type also would have been a factor in trace-fossil preservation, par- ticularly the grain size of the hosting medium. The fact that the lithofacies as- sociated with this assemblage is coarser grained and monotextured compared to the outer sand-flat lithofacies would have contributed to the subtle ichnological and taphonomic differentiation of the intertidal assemblages. The course texture of the sediment common to the inner sand flats may have led to the exclusion of other ichnospecies of Palaeophycus (besides P. tubularis) where taxonomic cri- teria are based on the delicate ornamentation of the burrow lining, which may not have been preserved in the coarser sediment. Environmental parameters like current energy, oxygenation, and salinity also played an important role in defining the character of the inner sand-flat assem- blage. What is apparent in Figures 3 and 16 is the presence of complex fodinichnia like ITeichichnus and Trichophycus, and epichnia like Archaeonassa in the more offshore, and assumed more normal, marine parts of the assemblage. Absence of these trace fossils in shoreward parts of the assemblage is probably due to an environmental gradient occurring perpendicular to depositional strike. Ecological stress acting across the inner-flat benthos is also implied by the average size of the preserved ichnotaxa compared to those same ichnotaxa in the outer flat. Modem outer- and inner-flat areas, such as in the Gulf of California, exhibit a shoreward decrease in size of organisms, particularly mollusks, inhab- iting both tidal regimes (Fiirsich and Flessa, 1987). Inner-flat gastropods and bi- valves were considerably smaller, even within the same species, than their outer- flat counterparts (Fiirsich and Flessa, 1987). The authors did not have a definitive explanation for this phenomenon, but stated it could be attributed to either the ecological factors inherent of the stressful inner-flat environments, or to waning current energy across this nearshore area that transported only the smaller species from the outer flat. Within the Deadwood Formation, Planolites occurred with high frequency in all lithofacies, represented by P. montanus and P. beverleyensis. Ichnospecific distinction of these two forms is based primarily on the size of the specimens. Planolites beverleyensis has trace diameters 5-8 mm larger than P. montanus (Pemberton and Frey, 1982). In all measured sections, P. montanus was the only Planolites ichnospecies collected from lithofacies 1, suggesting that larg- er forms of the Planolites-pxoducing animal were excluded from inner sand-flat environments. A similar situation occurs in the Lower Ordovician Santa Rosita Formation in Argentina, where high intertidal lithofacies consist of monoichnos- pecific trace-fossil assemblages of P. montanus (Mangano et al,, in press). Trace- fossil assemblages in the offshore parts of the intertidal zone of the Santa Rosita show a greater degree of trace-fossil complexity and diversity compared to the P. montanus assemblage. This contrast between inner and outer tidal-flat trace-fossil assemblages appears similar to the transition between lithofacies 1 and lithofacies 2 of the Deadwood Formation, and indicates similarities between many Cambro- Ordovician intertidal deposits. Size trends are also observed in burrows of Pa- Fig. 16. — Lateral distribution of collected ichnogenera and environments of deposition during time T of Figure 4. No vertical or lateral scale is implied. N.H.T. and N.L.T. represent normal high tide and normal low tide, respectively. E.W.B. represents effective wave base. 44 Annals of Carnegie Museum VOL. 67 laeophycus tubularis, which are 7 mm smaller in diameter in lithofacies 1 as compared to lithofacies 2 (see discussion under Palaeophycus tubularis in Sys- tematic Ichnology). Differences in the size of trace fossils also were noted by Hakes (1985) from Upper Pennsylvanian marginal-marine settings of Kansas. Hakes (1985:31, table 2) showed that many of the trace fossils from nearshore brackish-water facies were much smaller than those same trace fossils from nor- mal marine facies. This decrease in size was attributed to variable salinities en- countered in the marginal-marine trace-fossil assemblages, which probably inhib- ited the trace makers from attaining full adult size. It seems unlikely that selective destruction of larger burrows contributed to the size differences between the outer sand-flat and inner sand-flat trace fossils. The analogy can then be drawn between the Deadwood intertidal assemblages and those exhibited by bivalves and gastropods from the Gulf of California, suggesting that size variability in taxa across the intertidal zone is due to ecological factors. Outer Sand-flat Assemblage. — Ichnotaxa diagnostic of this assemblage includes Archaeonassa, Skolithos, Gyrolithes, IScolicia, Arthraria, and Planolites bever- leyensis (Table 1; Fig. 16). This assemblage is characterized by an increase in numbers and diversity of ichnogenera compared to the inner sand-flat assemblage. Endobenthic domichnia and fodinichnia of Palaeophycus, Skolithos, Planolites, and Trichophycus, and repichnion of Archaeonassa increase markedly. New ich- notaxa represented by domichnia {Arthraria) and fodinichnia {Psammichnites, Gy- rolithes, and IScolicia) are also encountered, and indicate more epibenthic and shallow endobenthic modes of life. Trilobite cubichnion and repichnion in the form of Rusophycus and Cruziana also occur, but are relegated to the more off- shore parts of the assemblage. Stronger current action and more normal-marine open circulation, implied by the presence of planar laminations and current-rippled bedding surfaces, accompanies the increased ichnogeneric diversity. The combi- nation of normal-marine circulation and tidal-dominated primary sedimentary structures suggest that this trace-fossil assemblage inhabited an offshore, moder- ate- to high-energy part of the tidal flat. Preservational acuity of the different ethological trace fossils increases in the outer sand-flat assemblage, and is directly related to increased sediment hetero- geneity found in lithofacies 2. Important to the preservation of surface and near- surface biogenic structures would have been clay drapes deposited from waning tidal currents. These clay drapes, represented as thin shale partings and flaser bedding, would have protected the epichnial and shallow endichnia from erosion at times of exposure and resubmergence of the assemblage during the transition from low to high tide. Conversely, lack of an appreciable mud or fine-grained sand component in the inner sand-flat environment probably contributed to the poor preservational quality of that assemblage. A strong ecological gradient still occurred across the outer sand flats as Ru- sophycus and Cruziana only occur in the offshore parts of the assemblage (Fig. 3, 16). Lack of these ichnogenera in the more landward parts of the outer sand flat suggests that environmental conditions were still too hostile for normal-marine trilobites. Other ichnogenera that overlap from the lower energy areas of the subtidal zone include Crossopodia and Didymaulichnus (Fig. 16), and imply that environmental conditions were more hospitable for their trace makers toward the offshore parts of the sand flats. Both the outer and inner sand-flat assemblages represent a mixed or overlapping Skolithos-Cruziana ichnofacies. Displacement or total absence of a particular ich- 1998 Stanley and Feldmann— Trace-fossil Assemblages 45 nofacies from the archetypical ichnofacies model has been well documented (Ek- dale, 1988; Bromley, 1990; Frey et ah, 1990). A notable example for a modem tidal flat is from the Yellow Sea, which falls entirely within the Cruziana ich- nofacies, with no Skolithos ichnofacies shoreward (Frey et ah, 1987). A mixed ichnofacies is commonly found in modem and ancient environments that exhibit fluctuating depositional and taphonomic conditions, such as intertidal and estua- rine environments (Ekdale et ah, 1984:172, 179; Ekdale, 1985; Bjerstedt, 1988). Within the outer sand-flat assemblage current energy was probably the most vari- able environmental parameter. Strong currents suggestive of bed-load transporta- tion are implied by the presence of parting lineations, cross-bedding, and current- rippled bedding surfaces, while the presence of shale partings and flaser bedding implies suspended-load deposition from waning tidal currents. The net result is a mixed trace-fossil assemblage and taphonomic signature characterizing both high and low current energy. Restricted Subtidal-shelf Assemblage. — This trace-fossil assemblage corre- sponds to lithofacies 3 (Fig. 3, 16). Cubichnia and repichnia (Rusophycus and Cruziana) dominate and are the most diagnostic ichnotaxa of this assemblage. Other ichnotaxa include repichnion of Archaeonassa and Didymaulichnus, and fodinichnia of Taenidium, lArthrophycus, Planolites, Psammichnites, Crossopo- dia, and Thalassinoides (Table 1; Fig. 16). The presence of fine-grained terrige- nous and carbonate sediments, well-developed, thin to fissile bedding, coupled with epibenthic cubichnia and repichnia suggests that this facies formed in a subtidal, normal-marine environment characteristic of the Cruziana ichnofacies. Siliciclastic deposition was sporadic, allowing deposition of carbonate sediment during times of low siliciclastic influx. Periodic storms eroded carbonate sediment from peritidal banks to the west (Fig. 2) and redeposited the flotsam as lenses of intraclastic limestones. Little or no wave reworking is evident in the intraclasts, indicating that this lithofacies formed below normal wave base but not below storm wave base. Previous paleoecological studies interpreted this lithofacies as forming on in- tertidal mud flats (Lochman-Balk, 1964, 1970, 1971; Lochman-Balk and Wilson, 1967). However, the taphonomy of the preserved trace fossils, characterized by finely preserved epibenthic and shallow endobenthic traces, coupled with the cha- otic nature of the intraclastic limestones, contradicts an intertidal interpretation of this lithofacies. The high diversity but low abundance of traces characteristic of this assemblage suggests periodic disruption of the normal-marine benthos. Low abundance of trace fossils is an ecological response of the benthic community to storm surges that inhibited the establishment of well-developed communities in the substrate. A similar association was noted by Pemberton and Frey (1984) and Vossler and Pemberton (1988) for mixed quiet-water and tempestite deposystems in the Upper Cretaceous of Alberta. One apparent irregularity of this assemblage is the low density of endobenthic fodinichnia compared to the outer sand-flat assemblage. The lack of strong bio- turbation is documented by the well-developed fissility of the shales and absence of burrow mottling in the micritic limestones. The absence of large numbers of infaunal traces may also have be a response to tempestite deposition, but it seems more likely due to hardground or firmground development, which tends to de- crease endobenthic activity and trace fossil diversity (Ekdale, 1988). Hardground to firmground development on intraclastic limestones and on some micritic lime- stone beds is suggested by the presence of expansion cracks on limestone bedding 46 Annals of Carnegie Museum VOL. 67 surfaces formed during early cementation, as well as by protrusion of intraclasts above bedding. The environmental interpretation of subtidal deposition coupled with early cementation in this lithofacies is also supported by Myrow (1995), who noted a strong correlation between the ichnogenera Thalassinoides and firmground to hardground development. Thalassinoides also appears to be restricted to inter- tidal and shallow subtidal environments of deposition. Although Thalassinoides is not common in this assemblage, its presence strengthens the interpretation that hardground development may have played a part in restricting bioturbation and trace-fossil density, Hardground development is not unique to this lithofacies, and has been documented from similar units in Montana and Wyoming (Brett et al., 1983). Conclusions Combining lithostratigraphic and ichnologic data is an extremely effective aid in deciphering Deadwood-Aladdin paleoenvironments. Sedimentological and hy- drodynamic responses of the enclosed ichnotaxa allowed for segregation of in- tertidal deposystems into two assemblage zones, an inner and outer tidal flat. Due to the extreme changes in environmental and taphonomic parameters, the trace- fossil assemblage enclosed in the inner-flat environment is characterized by low ichnofaunal diversity and abundance. Parameters such as salinity, water temper- ature, current energy, and sediment heterogeneity greatly influenced the character of this assemblage. Key ichnotaxa that differentiate the outer sand flat from the inner sand flat include: Archaeonassa, Skolithos, IScolicia, Gyrolithes, Arthraria, and Planolites beverleyensis. Taphonomic characteristics, as well as changes in the size of the preserved trace fossils, can also be used as guides in segregating the intertidal assemblages. The subtidal shelf assemblage is characterized by an increased number of shal- low repichnia and cubichnia compared to the intertidal assemblages. Main envi- ronmental controlling factors on this assemblage appear to be frequent tempestite deposition that may have temporarily disrupted the quiet- water benthos. Lack of deep burrowing ichnofauna within this assemblage may also have been due to the periodic storms that would have exposed firmground or hardground surfaces, and inhibit deep infaunal burrowing. Acknowledgments Dr. C. G. Maples read early drafts of this manuscript and made helpful suggestions, and two anonymous reviewers greatly improved the manuscript. Dr. M. Miller provided important German literature and Dr. L. Wiedman assisted in the translation. Field costs were partially defrayed with a grant from Sigma Xi. Contribution 362, Department of Geology, Kent State University, Kent, Ohio 44242. Literature Cited Alexander, C. R., C. A. Nittrouer, D. J. Demaster, Y. 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V »• ’'n<'#’>l4U< I %- ’.--f ■ >‘- '.' ■ -i^ i ^ -t ii .-•I ifT<^f - ..r( / ■: ■ u-: ■.■< '"'-V J;ipt.:., I . <^c-v ^ -t. -n^lwy^ ■ ' -- • , ti'i* f.i/Vf’' V JtJff ( i i '» ' i«,.< ' , K* ■ r>-t , f' wm^ ■ V-'' ^♦4y|»v.7'i.«-?r-Eip^.^ -T3 , y%?#* .,=-.*.-#*5 «l*,a ' ■■ - 'i* 'TWiHr-Wi.' 3 ♦-.A) * ^ ,irp . - - ’ e ^ ■ « •; - ••■' -t ' - j*. ... « F * . \ . '-a- ’ ■ . ■>«. :\ ^ ..y ■ W^t^ .;z.:'fi*iiv t '- f V d ' I »*-• If JJ. ' -^7'% # •v . *. >dSsS^ . . ‘%^*7 . ' ■ ■ ■ ; c. n ’ ■• \ v**Ji*i* • •■ "* '>ii' ■ ' ' ., ;■ '■ /- ' ^ , «»j.3 .'►.. Vi'^. <• ” '■“ ' i ^ ■ % ^ -f ■ *#-^5 1» * -^ r ** '■''(il3' SJ*4 - ^ ■,^ --T*, ,« . - - . 4 - ^ _tC^ "V „• j '-.,/ 5' J ^ • .« ■ r * . •^ 'i i . J ^ nr.:- v^ , r-*^'. ■ *. ' , . .. ,^- •, -4 , • /' .i. ■'- -..fi/ «?!•'''.> 1 •’ '• 4 ! ■>‘^i»^ * i ^’*. ■-* •> • * * " ***< • • • • ' '"* ■r- * f' 4 tt: . k-1 . " . f- I-. ^■7* \ [ *4*1^' ■* - - -4'* 34l*f f'.f’ff '’ P" 'f, f , * ; i. » *♦» h'“ ' > ' ■^J , ' tt ■ ."- .- . ■, . ^-‘' ".. ■ ,' ♦ f^' *4, ,%♦©;■ .' jF.t ». .„. ’. v-lf" ! ‘Je« 1 ANNALS OF CARNEGIE MUSEUM VoL. 67, Number 1, Pp. 53=93 12 February 1998 DIADECTES (DIADECTOMORPHA: DIADECTIDAE) FROM THE EARLY PERMIAN OF CENTRAL GERMANY, WITH DESCRIPTION OF A NEW SPECIES David S Berman Curator, Section of Vertebrate Paleontology Stuart S. Sumida' Research Associate, Section of Vertebrate Paleontology Thomas Martens^ Abstract A new species of the diadectomorph Diadectes, D. absitus, is described on the basis of a nearly complete, articulated skeleton that includes the skull, an isolated skull, and the greater portions of two articulated postcrania. All were collected from fluvial red=bed deposits immediately above the Tambach Sandstone, middle part of the Lower Permian Tambach Formation, lowermost formational unit of the Upper Rotliegend, of the Bromacker locality in the midregion of the Thuringian Forest near Gotha, central Germany. Diadectes absitus represents the first member of this genus to be described from outside of North America, A combination of autapomorphic and plesiomorphic characters provides a more substantial basis for recognizing D. absitus than is available for distinguishing between the North American species of Diadectes. Two possible conclusions are drawn from the relationship of D. absitus to the North American members of the genus: 1) its uniqueness reflects the wide geographic separation from the North American species, and 2) its greater primitiveness reinforces previous assessments of the biostratigraphic position and age of the Tambach Formation as earliest Permian Wolfcampian. Key Words: Diadectidae {Diadectes), Upper Rotliegend, skeleton, Bromacker locality, Germany, Pangaea Introduction Until very recently, the presence of diadectids in the Lower Permian of Europe has been limited to two very incomplete specimens described over a century ago from the Lower Rotliegend of Germany. A string of four presacrals and two sacrals from the Leukersdorf Formation of the Erzgebirge Basin near Zwickau was described by Meyer (1860) as Phanerosaurus naumanni, and Geinitz and Deichmueller (1882) described P. pugnax on the basis of disarticulated skull and postcranial elements from the Niederhaeslich in the Doehlen Basin near Dresden. In a lengthy restudy of the latter species, Stappenbeck (1905) reassigned it to a new genus as Stephanospondylus pugnax. Although the dentition and/or vertebrae of both forms clearly justifies a diadectid assignment, the lack of additional diS“ coveries has not permitted detailed comparisons with the better known diadectids, particularly Diadectes of North America, and both of the German taxa remain poorly defined (Romer, 1925). ' Department of Biology, California State University, 5500 University Parkway, San Bernardino, Cal- ifornia 92407. 2 Abteilung Palaontologie, Museum der Natur, PSF 217, D-99853 Gotha, Germany. Submitted 9 January 1997. 53 54 Annals of Carnegie Museum VOL. 67 During a period of nearly two decades, four specimens assignable to Diadectes as a new species have been recovered from a sandstone quarry known as the Bromacker locality that lies in the Lower Permian Tambach Formation, lowermost unit of the Upper Rotliegend, in the middle part of the Thuringian Forest near Gotha, central Germany (Berman and Martens, 1993; Sumida et aL, 1996). The first specimen discovered, consisting of a nearly entire, slightly disarticulated post- cranial skeleton, was collected by Martens and Schneider in 1979 during an ex- ploratory excavation at the Bromacker quarry, but was misidentified as possibly a limnoscelid diadectomorph (Martens, 1980, 1988, 1989; Martens et aL, 1981). In the summer of 1991, Martens discovered a nearly complete, isolated small skull of a juvenile specimen which he identified as a diadectid in a brief note (Martens, 1992). Two closely associated adult specimens were collected by the authors in the summer of 1993. One consists of a nearly complete, articulated skeleton with an excellently preserved skull (designated here as the holotype of a new species) and the other the greater portion of an articulated postcranium. Collectively, these specimens represent the first occurrence of Diadectes outside of the Lower Permian of North America. The Bromacker Diadectes is part of a diverse assemblage of terrestrial or sem- iterrestrial amphibians and reptiles represented by articulated skeletons from the Bromacker locality (Martens, 1980, 1988, 1989; Boy and Martens, 1991; Berman and Martens, 1993; Sumida et aL, 1994, 1996). To date, three Bromacker verte- brates have been described: 1) a single, nearly complete skeleton of a new genus of protorothyridid amniote, Thuringothyris mahlendorjfae Boy and Martens, 1991; 2) two specimens of the seymouriamorph amphibian Seymouria were ten- tatively referred to S. sanjuanensis Vaughn, 1966 (Berman and Martens, 1993); and 3) a skull and partial, articulated postcranium representing a new genus and species of the amphibian family Trematopidae (Sumida et aL, 1994). Seymouria and trematopids are known otherwise only from the Early Permian and Permo- Pennsylvanian, respectively, of North America. For over a century the Bromacker locality has been well known as a very important source of excellently preserved tetrapod trackways preserved in the Tambach Sandstone of the Tambach Formation (Pabst, 1896, 1908; Muller, 1954, 1969; Haubold 1971, 1973). Vertebrate skeletal remains, however, eluded detec- tion at Bromacker until 1974 (Martens, 1980). All of the vertebrate skeletal spec- imens so far collected from Bromacker have come from fluvial red-bed facies immediately above the sandstones yielding the trackways and are quite similar to those deposits which have yielded the vast majority of the Lower Permian and Permo-Pennsylvanian vertebrates of the United States (Martens, 1975, 1982, 1988, 1989; Martens et aL, 1981; Berman and Martens, 1993; Sumida et aL, 1996). With the exception of Bromacker, terrestrial or semiterrestrial vertebrates from Permian red-bed deposits of the Rotliegend or their equivalent in central and western Europe are quite rare and include most notably the primitive syn- apsids, Haptodus (Paton, 1974; Currie, 1979; Laurin, 1994), Ophiacodon (Paton, 1974), and Sphenacodon (Paton, 1974) from England and Casea (Sigogneau- Russell and Russell, 1974) from France. The apparent uniqueness of Bromacker among European localities has been explained as possibly an artifact of collecting (Martens, 1989; Berman and Mar- tens, 1993; Sumida et aL, 1996). The red-bed exposures of the Upper Rotliegend are very poor and have been commonly perceived as representing an inhospitable dry climate that was not conducive to preservation of skeletal remains. As a result. 1998 Berman et al.— Early Permian Diadectes from Germany 55 relatively little energy has been devoted to their exploration, and, as a conse- quence, they have yielded few vertebrates. Alternatively, intensive prospecting has focused on the highly fossiliferous lacustrine grey sediments and black shales of the Lower Rotliegend that reflect limnetic environments. These deposits, in which are found the well-known tetrapod localities as Niederhaeslich, Friedri- chroda, and various Saar-Nahe sites such as Lebach, have yielded a great number and variety of obligatory aquatic amphibians, but only very rarely terrestrial or semiterrestrial tetrapods (Milner and Panchen, 1973; Milner, 1993). Two important conclusions have been suggested (Berman and Martens 1993; Sumida et al., 1996; Berman et al., 1997) to explain the high degree of com- monality between the Bromacker tetrapod assemblage and those of the Lower Permian red-bed deposits throughout the United States: 1) similar environments, as represented by typical fluvial red-bed facies, are being sampled; and 2) an absence of major physical and biological barriers during the Early Permian al- lowed faunal interchange across northern Pangaea. The striking similarities between the Bromacker assemblage and those of North America have also brought into question the widely accepted assessments of the biostratigraphic position and age of the Tambach Formation, lowermost forma- tional unit of the Lower Permian Upper Rotliegend of the Tambach Basin in the midregion of the Thuringian Forest of central Germany. The Rotliegend, as well as its lower and upper subdivisions (in western Europe the lithostratigraphic terms Autunian and Saxonian are commonly used in place of Lower and Upper Rotlie- gend, respectively), is a traditional lithostratigraphic unit that refers to continental beds considered to be entirely or in great part Lower Permian and to overlie Upper Carboniferous Stephanian deposits in central Europe. In the region of the Thuringian Forest the Upper Permian marine Zechstein overlies in places the Rotliegend. Certain elements of the Bromacker assemblage, particularly Sey- mouria cf. S. sanjuanensis, Diadectes, and trematopid, strongly suggest an earliest Permian Wolfcampian age for the Tambach Formation. This, in turn, indicates that most or possibly all of the underlying Lower Rotliegend in the Thuringian Forest region should be reinterpreted as Upper Carboniferous (Berman and Mar- tens, 1993; Sumida et al., 1996), rather than the widely accepted Lower Permian. The following acronyms are used to refer to institutional repositories of spec- imens: CM, Carnegie Museum of Natural History, Pittsburgh, Pennsylvania; FMNH, Field Museum of Natural History, Chicago, Illinois; MCZ, Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts; MNG, Museum der Natur, Gotha, Germany; UCMP, Museum of Paleontology, University of California, Berkeley, California. Anatomical structures are identified by the following abbreviations: a, angular; af, anterior fenestra; ar, articular; ati, atlantal intercentrum; atn, atlantal neural arch; axe, axial centrum; axi + atp, axial intercentrum plus atlantal pleurocentrum; axn, axial neural arch; bo, basioccipital; c, coronoid; c3, centrum 3; d, dentary; ec, ectopterygoid; en, external naris; fp, footplate of stapes; i3, intercentrum 3; in, internal naris; j, jugal; 1, lacrimal; m, maxilla; mf, medial fenestra; n, nasal; na4, neural arch 4; oc, occipital condyle; osp, ossified plate of stapes; ot, otic trough; p, parietal; pal, palatine; pat, proatlas; pm, premaxilla; po, postorbital; pop, paroccipital process; pp, postparietal; pra, prearticular; pro, prootic; ps, para- sphenoid; pt, pterygoid; q, quadrate; qf, quadratojugal foramen; qj, quadratojugal; rl, atlantal rib; r2, axial rib; r3, r4, presacral ribs 3 and 4; sa, surangular; sf, stapedial foramen; sm, septomaxilla; sop, supraoccipital-opisthotic; sp, splenial; 56 Annals of Carnegie Museum VOL. 67 sq, squamosal; ss, stapedial shaft; st, supratemporal; t, tabular; v, vomer; I-V, carpal digits. Systematic Paleontology Order Diadectomorpha Family Diadectidae Genus Diadectes Cope 1878 Diadectes absitus, new species Holotype. — MNG 8853, greater part of an articulated skeleton with skull con- tained in two adjoining blocks and lacking the following: large portions of the tail; interclavicle; left shoulder girdle, forelimb, and manus; right radius, ulna, and manus; left hindlimb and pes; and most of the right pes. A disarticulated right clavicle preserved directly above the cervical vertebrae was removed. A string of four distal caudal vertebrae lying a short distance to the right of the midlength region of the presacral column almost certainly belongs to the holotype. Paratypes. — MNG 8747, greater part of skull with lower jaws; MNG 7721 and 8978, greater por- tions of articulated and partially articulated, respectively, postcranial skeletons. Horizon and Locality. — Uppermost level of 60 m-thick Tambach Sandstone of the Lower Permian Tambach Formation, Upper Rotliegend, in the Bromacker locality of the middle part of the Thuringian Forest near the village of Tambach- Dietharz and about 20 km south of the town of Gotha, central Germany. Diagnosis. — Can be distinguished from all other Diadectes species by the fob lowing autapomorphic features: 1) postfrontal chevron-shaped, with its posterior apex penetrating deeply into anterior margin of parietal; 2) postorbital has very narrow entrance into orbit; 3) abrupt, dorsalward, step-like retreat of anterior portion of jugal from the free, ventral margin of the skull; 4) lower jaw subcircular in cross-sectional shape at level of coronoid eminence with the height and width being nearly equal; and 5) dentary lacks a prominent labial parapet lateral to the cheek teeth and in its place is a wide, dorsally facing platform. The following plesiomorphic features distinguish Diadectes absitus from all other members of the genus: 1) dorsal process of premaxilla long and narrow, extending to a level beyond the posterior margin of the external naris; 2) prefrontal extends well be- yond the level of the anterior margin of the frontal; 3) postorbital forms for most of its posterior extent a tapering, triangular process that ends in a very narrow contact with the supratemporal; and 4) basicranial joint remains open and mobile in adults. Etymology. — Latin, absitus, meaning distant, apart, or remote, referring to its distant occurrence from all other known members of the genus that occur in North American. Description. — Skull Roof. The skull of Diadectes absitus conforms very closely to descriptions of North American members of this unique genus. For this reason the description that follows, including the postcranium, is mainly restricted to those features that are either at variance with previous accounts (Olson, 1947, 1950; Watson, 1954; Lewis and Vaughn, 1965; Berman et al., 1992) or are not obvious from the illustrations presented here or elsewhere of North American species. With this approach in mind, in all the figures presented here to illustrate the cranial anatomy of D. absitus the identification of structures has been limited mainly to those in which possible confusion may occur. In addition, cranial sutures have been drawn on the skulls with ink. The skulls of the holotype MNG 8853 and the paratype MNG 8747 (Fig. 1-8) are approximately the same length (13.0 cm), but in other measurements, particularly transverse dimensions, the holotype is 9-12% larger. The holotypic skull is complete and has undergone minor dorsoventral crushing of the right side and occiput and lateral crushing of the left side. The paratypic skull MNG 8747, on the 1998 Berman et al,- — Early Permian Diadectes from Germany 57 other hand, has been crushed moderately dorsoventrally, and considerable amounts of bone have been lost from the cheek regions. As in the North American species, the roofing bones of both skulls are thick and porous, with a high degree of relief, particularly on the table portion, where well-developed, irregular prominences are created by a network of prominent, smooth channels or grooves. Also as in the North American species, these features are more prominent in the smaller, less mature MNG 8747 than in the holotypic skull. The premaxillae are preserved in both skulls, but only in MNG 8853 (Fig. 3, 4) are the dorsal processes well preserved. They are narrow, tapering slightly to blunt points as they extend a consid- erable distance onto the dorsal surface of the skull to a level well beyond the posterior margin of the external nares. Each premaxilla possesses four procumbent, incisiform teeth which are subequal in size except for the fourth being considerably smaller. Although the premaxillary teeth of MNG 8853 are all present, only their labial surfaces are visible. In MNG 8747 only the bases of the teeth remain except in the right premaxilla, where the fully erupted third is missing the tip of the crown and only the crown of the partially erupted fourth is visible. The crowns of replacement teeth are also visible in the lingual pits of the second tooth of the right premaxilla and the first and third of the left. The teeth are distinctly procumbent and incisiform. The bases are oval in cross section with the long axis directed posteromedially, whereas the lingual surface of the distal portions are broadly concave, giving them a chisel-like appearance. The paired nasals and frontals are narrowly rectangular and nearly equal in their dimensions. The frontals, however, are slightly shorter and gradually widen slightly posteriorly. In both skulls the great width of the parietals is due to a pronounced, well-defined rectangular lateral lappet that replaces, or represents the incorporation of, the intertemporal bone (Berman et al., 1992). In MNG 8747 a narrow, spike-like projection of the posterior margin of the parietal partially separates the tabular and supra- temporal. In MNG 8853 the projection is much broader and subrectangular, but this may be due in part to crushing in the occipital region. The postparietal is a single, anteroposteriorly narrow, rectan- gular bone, which in MNG 8853 has a width approximately four times its midline length. Approxi- mately one-third of the postparietal surface area, extending along its anterior margin, is exposed on the skull table, whereas the remaining posterior portion of the bone is angled abruptly downward onto the occiput to overlap the dorsal margin of the supraoccipital. In MNG 8747 the posterior portion of the postparietal slopes ventrally from the skull table at about 45°, whereas in MNG 8853 the surface is vertical. Although the prefrontal extends noticeably beyond the level of the anterior margin of the frontal in both skulls, the extension is greater in MNG 8853. The chevron-shaped postfrontal forms the posterodorsal margin of the orbit, with the posterior apex penetrating deeply into the anterior margin of the parietal. Description of the cheek bones is based almost exclusively on MNG 8853. The postorbital has a narrow entrance into the posterior corner of the orbit. The greater portion of the postorbital forms a posteriorly narrowing, triangular process that extends along the lateral margin of the parietal lappet to within a short distance of the expanded otic, or temporal, notch. The postorbital ends in a narrow contact with the supratemporal, thus preventing contact between the lateral lappet of the parietal and squamosal. At its anterior contact with the premaxilla the maxilla has a short, narrow, dorsally directed, process-like extension that enters the external naris. Posteriorly the maxilla extends to about the level of the midlength of the orbit. Only the dentition of the right maxilla of the juvenile skull MNG 8747 is fully exposed and well preserved, and includes 1 1 teeth at various stages of replacement: teeth 2, 4, 6, 7, and 9 are fully erupted, ankylosed, and show wear on the lingual cusp and on some of the central cusps as well; teeth 1, 5, and 10 are fully erupted, ankylosed, and show no wear; and teeth 3, 8, and 11 are partially erupted and not ankylosed. The crowns of replacement teeth are visible in the lingual pits of the second, fourth, and ninth teeth. The maxillary dentition is essentially identical to that of other members of the genus. Yet, a description providing some details is warranted, because Diadectes exhibits marked developmental growth stages of the marginal dentition that differ from those in the very closely related. Late Pennsylvanian Desmatodon (Vaughn, 1969, 1972; Berman and Sumida, 1995). Although all the maxillary teeth in MNG 8747 are expanded transversely, they are positioned so that the labial edge of each tooth lies well anterior to the lingual edge. The first two teeth are slightly longer and more incisiform than the succeeding maxillary teeth, giving them a morphology intermediate between those of the premaxilla and the more posterior, molar-like cheek teeth. The labial cusp is essentially absent, the posteromedial surface of the well-developed central cusp is broadly concave, and the lingual cusp is weakly developed. The crown of the third tooth is only partially erupted, but also appears to be incisiform. The remaining teeth of the series increase in size to the sixth tooth and then decrease to the end of the series. The cheek teeth exhibit the typical molar-like structure seen in other species of Diadectes, although their degree of development or “molarization” clearly reflects a juvenile stage of growth (Berman and Sumida, 1995). The subconical central cusps of the cheek teeth are very pro- Annals of Carnegie Museum VOL. 67 Fig. 1. — Diadectes absitus, holotype (MNG 8853). Greater part of articulated skeleton exposed mainly in dorsal view as preserved in two blocks and showing the skull, vertebral column except for large 1998 Berman et al. — Early Permian Diadectes from Germany 59 nounced and represent possibly half the height of the crown, whereas the lingual and labial cusps are poorly developed and would be more accurately described as “shoulders.” The lingual cusps are much more sharply defined than the labial cusps, as the bases of the central cusps are much more expanded laterally than they are medially. The sixth and largest of the molariform cheek teeth measures 7.0 mm in transverse width, 3.7 mm in anteroposterior length, and 5.0 mm in height. The spacing between cheek teeth 4 through 7 is about 1.0 mm. Enough of the right maxillary dentition of MNG 8853 is exposed laterally to indicate the presence of at least ten teeth plus spaces for two more. The teeth appear to exhibit the same serial changes as in MNG 8747. The lacrimal and jugal are preserved only in MNG 8853. Their contribution to the suborbital bar on the right side of the skull is unusual, however, in that the suborbital process of the jugal appears to form the entire ventral border of the orbit. The typical condition for this structure is seen on the left side of the skull, where the suborbital process of the jugal is shorter, and the ventral orbital rim is completed anteriorly by a short posterior, suborbital process of the lacrimal. The jugal’s contribution to the ventral margin of the skull exhibits an abrupt, step-like dorsalward retreat just anterior to the quadratojugal. This feature is especially pronounced in the left jugal. Only the right septomaxilla of MNG 8853 is sufficiently well preserved and accessible for descrip- tion (Fig. 4), and in general conforms to that described in Early Permian synapsids (Romer and Price, 1940; Wible et al., 1990). It is subrectangular in outline and lies deep within the external naris. However, because its margins are extremely thin and a major crack through the skull extends along its ventral contact with the premaxilla, the exact outlines of this element are difficult to discern. The posterior margin appears to contact the lacrimal, the dorsal margin nearly reaches the nasal, but their proximity may have been exaggerated by dorsoventral crushing of the skull, and the anterior margin ends free at about midlength of the external naris. Basically the septomaxilla is deeply bowed medially, so that its dorsal and ventral margins curve outward to the facial surface of the skull. Most prominent is a narrow, pillar-like structure, described as the dorsal process in primitive synapsids, which extends posterodorsally, joining the lateral lips of the dorsal and ventral margins of the septomaxilla. The outer surface of the dorsal process may have had a facial exposure. The dorsal process is pierced by an anteroposterior canal, the septomaxillary canal, which in primitive synapsids is believed to have transmitted branches of the superior labial nerve and artery (Wible et al., 1990). Crushing of the skull appears to have resulted in a small amount of lateral overlap of the posterior margin of the septomaxilla by the lacrimal. This may have obscured the presence of a septomaxillary foramen which is normally present at the union of the two elements. MNG 8853 and 8747 exhibit features of the squamosal and quadratojugal (Fig. 5, 6, 7B, 9A) either overlooked or not preserved in North American specimens described by previous authors. Not men- tioned in previous descriptions is a broad flange of the squamosal that projects medially and slightly posteriorly from nearly the entire length of the posterior or otic margin of its facial exposure that can be referred to as the medial flange. In MNG 8747 most of the facial exposure of the right squamosal has been lost, revealing the medial flange. It has an extensive, overlapping suture with all but a narrow ventral margin of the anterior surface of the portion of the quadrate which forms the transversely wide, anterior wall of the otic notch. In Figure 9A, however, only the lateral edge of the medial flange of the squamosal is visible. As seen in the skull MNG 8853 (Fig. 5, 6, 7B), the posterior surface of the medial flange of the squamosal is not overlapped entirely by the quadrate, and a narrow strip along its lateral margin is exposed in occipital view. Further, the facial surface of the otic margin of the squamosal is extended posteriorly 2 or 3 mm into a knife-like edge whose inner surface curves abruptly medially to become the medial flange. The quadratojugals are represented by only those in the holo- typic skull MNG 8853, but only the right one is well preserved and accessible. It completes the sculptured, posteroventral rim of the otic notch, ending in a small, free, triangular, medially directed process. Below and just anterior to the sculptured medial process is a narrow, nonsculptured, sharply wedge-shaped medial extension of the quadratojugal that enters the anterior wall of the otic notch to form the medial margin of the quadratojugal foramen before ending just above the lateral condyle of the quadrate. In MNG 8747 the facial exposure of the right quadratojugal has been lost and only a broad medial flange like that of the squamosal remains. The medial flange overlaps not only the ventral margin of the anterior surface of the portion of the quadrate forming the anterior wall of the otic notch, but also the greater portion of the anterior surface of the medial flange of the squamosal. portions of caudal series, iliac blades, and right tibia, fibula, and proximal portion of pes. A series of four distal caudals presumed to belong to the holotype lies to the right of the midpresacral region of column. Scale = 10 cm. Annals of Carnegie Museum VOL. 67 Fig. 2. — Reverse side of two blocks seen in Figure 1 containing Diadectes absitus, holotype (MNG 8853) and showing the skull, anterior cervical vertebrae, right femur, and puboischiatic plate in ventral view, right cleithrum and scapulocoracoid in lateral view, and right humerus with distal expansion in ventral view. Scale = 10 cm. 1998 Berman et al, — Early Permian Diadectes from Germany 61 Fig. 3.— Diadectes absitus, holotype (MNG 8853). Skull in A, dorsal and B, lateral views. Scale = 4 cm. 62 Annals of Carnegie Museum VOL. 67 Fig. 4. — Diadectes absitus, holotype (MNG 8853). Right nasal region of skull in lateral view. Scale = 2 cm. Extending posteriorly from the end of the spike-like projection of the posterior margin of the parietal is a distinct suture that continues to the distal end of the sculptured, downturned, posterolateral, horn- like extension of the skull table. This suture marks the contact between the tabular and the supratem- poral. Whereas the tabular and supratemporal occupy subequal portions of the sculptured, horn-like extension of the skull table in MNG 8747, the supratemporal forms most of this structure in the larger MNG 8853. Both D. absitus skulls reveal that the ventral surface of the distal end of the horn-like extension of the supratemporal, that forms the posterodorsal margin of the otic notch, has a substantial contact with the dorsal edge of the distal end of the paroccipital process. From its sculptured contri- bution to the skull-table horn, the tabular extends medially and slightly anteriorly onto the occipital surface of the skull as a smooth plate of bone. This portion of the tabular contacts most of the dorsolateral margin of the supraoccipital-opisthotic complex of the braincase (Fig. 7B). From near the distal end of the paroccipital process, this contact arches slightly medially to reach the lateral margin of the occipital portion of the postparietal. Palate. The palate is possibly the most poorly known region of the skull of Diadectes. Olson (1947) presented the most recent and detailed reconstruction of the palate, based on several individuals, yet many features remain undescribed. The palate of D. absitus is best preserved and exposed on the right side of skull MNG 8747 (Fig. 10), although some breakage and loss of bone has slightly obscured the sutural contacts of the palatine and ectopterygoid. In MNG 8853 the right pterygoid, ectopterygoid, and posterior half of the palatine are exposed and well preserved, as well as part of the left pterygoid (Fig. 7A). Anteriorly the vomer forms a ventrally thickened, midline, ridge-like premaxillary process that ventrally overlaps the vomerine process of the premaxilla. The premaxillary processes are very narrowly separated and their distal ends are slightly forked. Although much of the posterior contact of the vomers with the pterygoids in MNG 8747 is poorly defined, it is still traceable. Posterior to the premaxillary process the vomer continues along the midline as a dorsoventrally thickened ridge to the level of the sixth maxillary tooth and supports a single row of eight teeth. This interpretation agrees with Olson’s (1947) reconstruction. From the dorsolateral margin of the premaxillary process a flat, wing-like, posterolateral extension of the vomer forms much of the anterior medial border of the long, narrow internal naris. The posterolateral wing of the vomer is not shown in Olson’s (1947) reconstruction. Anteriorly the palatal ramus of the pterygoid extends lateral to the adjacent, midline, tooth-bearing 1998 Berman et al. — Early Permian Diadectes from Germany 63 ridge of the vomer, contacting the posteromedial margin of the posterolateral wing of the vomer. Although most of the lateral margin of the pterygoid is hidden from view by the palatine, it is clearly highly unusual in contributing to the medial margin of the internal naris, forming as much as the posterior half of the medial border of the opening, and in preventing a vomer-palatine contact. The palatal rami of the pterygoids unite along the midline for much of their length, allowing for only a short, narrow interpterygoid vacuity posteriorly. The medial tooth-bearing ridge of the vomer is con- tinued posteriorly by the pterygoid, which contributes ten more well-developed teeth to the series. The double row of midpalatal teeth of the paired vomers and pterygoids have cylindrical bases and sharply pointed conical crowns. The teeth of both rows increase in size posteriorly from a vertical height of about 0.03 to 1.05 mm except for a marked decrease in the last three teeth. The midline vomer-pterygoid tooth rows and marginal dentitions lie at the same horizontal plane and below the slightly vaulted palatal surfaces. The transverse flange of the pterygoid is subrectangular in ventral outline, expanding somewhat distally along its free, posterior margin. In MNG 8747 the ventral surface of the process thickens so greatly distally, that in lateral view its distal end projects well below the plane of the palate as a broadly triangular, vertical shelf (Fig. 9A). The anterior margin of the flange is defined sharply, particularly in MNG 8853, by a shallow, dorsalward, step-like elevation to the ventral surface of the palate proper. The distal half of the anterior margin of the flange contacts the ectopterygoid. In both D. absitus skulls the transverse flange of the pterygoid lacks teeth and, whereas the ventral surface of the process in MNG 8853 is slightly textured, that in MNG 8747 is smoothly finished. A shagreen of minute denticles is visible along the medial margin of the right palatal ramus in MNG 8747 from the base of the transverse flange to the posterior end of the medial row of teeth. The interpterygoid vacuity is closed posteriorly on either side of the midline by a very broad, posteromedially directed, flange- like process of the pterygoid, referred to here as the basal process. All but a small, triangular, anter- omedial area of the ventral surface of the basal process is occupied by a slightly depressed articular facet that faces mainly ventrally and slightly posteriorly to receive the basipterygoid process of the braincase. The quadrate ramus of the pterygoid extends posteriorly to the level of the quadrate condyle. Beginning immediately behind the transverse flange the ventral edge of the ramus forms a narrow, shelf-like lateral flange that gradually tapers posteriorly to its termination at about three-fourths the length of the ramus. A short distance posterior to the basal articulation a small, narrow, convex flange projects nearly dorsally from near the ventral margin of the central portion of the internal surface of the quadrate ramus. This flange is similar to that described in Limnoscelis as the arcuate flange (Romer, 1946; Fracasso, 1983). The unique structure of the palatine in North American Diadectes (Olson, 1947; Berman and Sum- ida, 1995) is duplicated in the D. absitus (Fig. 7 A, 10). There is a pronounced, arcuate, secondary palatal shelf that extends medially from its contact with the inner margin of the maxillary alveolar shelf and ventral to the primary palatal shelf of the true palate. In palatal view the secondary shelf partially obscures most of the contact between the primary palatal shelf of the palatine and pterygoid. A ventrally thickened, rectangular process, referred to here as the posteromedial process, extends from the posterior end of the primary palatal shelf of the palatine. Proximally the posteromedial process of the palatine nearly closes the channel formed between the primary and secondary palatal shelves of the palatine. In MNG 8747 the posteromedial process of the palatine appears to reach nearly the transverse flange of the pterygoid, whereas in MNG 8853 it is continued to the transverse flange by a short, complementary process of the pterygoid. The secondary palatal shelf of the palatine is con- tinued a short distance posterolaterally by the ectopterygoid to the free, lateral margin of the palate. From here the shelf is continued by a short, narrow, posterior extension of the ectopterygoid that curves slightly ventrally to contact the distal end of the anterior margin of the transverse flange of the pterygoid. A small portion of the ectopterygoid posterior and medial to its contribution to the sec- ondary palate is depressed abruptly dorsally to form a shallow concavity that is bordered medially by the posteromedial process of the palatine and posteriorly by the transverse flange of the pterygoid. In Olson’s (1947:fig. 3) reconstruction of the palate of Diadectes he illustrated the depression at the junction of the ectopterygoid, pterygoid, and palatine as a circular pit, but noted that in some specimens this depression appears to penetrate the bone as a fenestra. Interestingly, he commented (p. 16) that the depression or pit lies in the position of the infraorbital fossa and may represent an incipient development of this opening. Palatoquadrate. Only the quadrate (Fig. 5-7, 9, 10) of this complex has been exposed and consists basically of three major components: the condyle and two vertical sheets or laminae that arise from the condyle. One lamina, which can be referred to simply as the otic or temporal lamina, extends for a considerable height above the condyle as the transversely wide, subrectangular plate that forms the anterior wall of the otic notch. Its summit contacts the ventral surface of the supratemporal and, as shown by Watson (1954), possesses a short process at its dorsomedial margin which ends in a gently 64 Annals of Carnegie Museum VOL, 67 convex oval face that inserts into a conforming facet on the lateral surface of the prootic. At the ventromedial corner of the posterior surface of the otic lamina, just above the medial lobe of the condyle, is a short, well-defined subcircular protuberance, the quadrate boss. Lateral to the boss the quadrate forms all but the lateral margin of the quadratojugal foramen. The second, vertical lamina of the quadrate, the dorsal process or lamina, extends anteriorly and slightly medially from the medial margin of the otic lamina, so that an approximate right angle is formed anterolaterally between them. Proximally the dorsal lamina may have reached the height of the otic lamina, but tapers somewhat anteriorly as the ventral margin of its medial surface overlaps the lateral surface of the quadrate ramus of the pterygoid. This contact is very extensive anteropos- teriorly, as the dorsal lamina of the quadrate extends to approximately the level of the posterior margin of the transverse flange of the pterygoid (Fig. 9A). The greater distal portion of the ventral edge of the dorsal lamina inserts in a very shallow groove on the dorsal surface of the narrow, lateral flange of the quadrate ramus of the pterygoid. Braincase. In MNG 8853 the only portion of the braincase visible in palatal view is the right side of the basiparasphenoid complex, whereas dorsoventral crushing has resulted in considerable tele- scoping and distortion of the braincase bones exposed on the occiput. On the other hand, these regions of the braincase in MNG 8747 are relatively well preserved and exposed (Fig. 9B, 10), and provide almost entirely the basis for the description that follows. The parasphenoidal rostrum is visible as a keel-like structure that extends a short distance across the interpterygoid vacuity. On the midline of its proximal end is a small, ventrally projecting protuberance. The well-developed basipterygoid pro- cesses are subrectangular in outline, constricting somewhat at their base. They project anterolaterally Fig. 5. — Diadectes absitus, holotype (MNG 8853). Right otic region in A, posterolateral and B, oc- cipital views. Scale = 2 cm. 1998 Berman et al. — Early Permian Diadectes from Germany 65 Fig. 6. — Outline sketches of views A and B of Figure 5 to indicate individual elements and other structures. Dotted line marks union of facial and occipital surfaces of squamosal. and slightly ventrally, with their articular facets facing dorsally and slightly anterolaterally. A mobile, basicranial joint was undoubtedly present in both MNG 8747 and 8853. Posterior to the basipterygoid processes the smooth, ventral surface of the basiparasphenoid continues a short distance before ex- panding on either side of the midline as thin, free-edged, ventrally arching sheets that extend poster- olaterally to form long, triangular projections, the cristae ventrolaterales. A very narrow space separates the posterior edges of the cristae from the overlying basiparasphenoid complex proper which continues a short distance posteriorly to a transverse, feathered contact with the basioccipital. In both D. absitus skulls the occipital condyle faces strongly posteroventrally, which may be due in small part to dor- soventral crushing. In posterior view the condyle is semicircular in outline, with rounded dorsolateral comers and a slight, middorsal concavity. In MNG 8853 the articular surface is essentially flat, whereas in MNG 8747 there is a deep, central notochordal pit. The prootics are exposed in palatal view of the skull MNG 8747 (Fig. 10). They arise anteriorly from near the base of the basipterygoid process, with which they are indistinguishably fused, and extend posterolaterally for a considerable distance immediately adjacent to the lateral margins of the cristae ventrolaterales as wing-like sheets. Distally their incompletely preserved medial margins be- come very thin and appear to have ended in an abutment contact with the lateral edges of the cristae ventrolaterales. The thickened lateral margins of the prootics curve dorsally, and, although their extent in this direction can be exposed only a short distance, it is assumed that they continued as the lateral walls of the braincase. Just posterior to the palatal exposure of the right prootic in MNG 8747 (Fig. 10) is a large, stout, crescent-shaped structure that projects ventrally from near the base of the par- 66 Annals of Carnegie Museum VOL. 67 Fig. 7. — Diadectes absitus, holotype (MNG 8853). Skull in A, ventral and B, occipital views. Scale = 4 cm. occipital process of the opisthotic. Its coarsely textured, concave surface faces posteromedially and ventrally. This structure, which has been termed the otic trough (Fracasso, 1983; Berman et al., 1992), has been identified in Limnoscelis and North American Diadectes, as well as in primitive synapsids, and described as being formed entirely by the opisthotic and projecting ventrolaterally from the pos- terior border of the fenestra ovalis (Fracasso, 1987; Berman and Sumida, 1990; Berman et al., 1992). Although a function has not been ascribed to the otic trough, the in-place exposure of the right stapes 1998 Berman et al. — Early Permian Diadectes from Germany 67 Fig. 8. — Diadectes absitus, paratype (MNG 8747). Skull in dorsal view. Scale = 2 cm. Annals of Carnegie Museum VOL. 67 Fig. 9. — Diadectes absitus, paratype (MNG 8747). Skull in A, lateral and B, occipital views. Scale — 2 cm. 1998 Berman et al. — Early Permian Diadectes from Germany 69 Fig. 10. — Diadectes absitus, paratype (MNG 8747). Skull in ventral view. Scale = 2 cm. 70 Annals of Carnegie Museum VOL. 67 in MNG 8747 indicates clearly that the stapedial footplate articulates with the posterolateral region of its outer, convex surface. In MNG 8747 the left exoccipital has been lost, revealing that it did not contribute to the articular surface of the condyle, but rather extended along its dorsal surface as a thin sheath flooring the foramen magnum. Immediately above its contact with the condyle the exoccipital is strongly waisted medially before continuing as a dorsomedially directed, wing-like process that borders the foramen magnum laterally. The exoccipital-opisthotic suture is quite distinct on the occiput, extending ventrolaterally onto the ventral margin of the braincase. Here it continues anteriorly, first intersecting the lateral margin of the large jugular foramen, presumably for cranial nerves IX-XI and the jugular vein, and then extends a short distance along the anteromedial basal margin of the otic trough before terminating. Although there is no visible supraoccipital-opisthotic suture, its approximate position, however, can be presumed. On the right lateral margin of the supraoccipital-opisthotic is a small opening bordered laterally by the tabular. This opening, visible on both sides in MNG 8853, is almost certainly the posttemporal fenestra, but is believed to be lost in very mature adults specimens (Berman et al., 1992). Typically in amniotes the supraoccipital-opisthotic suture extended dorsolaterally from the dorsolateral margin of the foramen magnum just above the contribution of the exoccipital to this opening and ended at or near the medial margin of the posttemporal fenestra. Immediately adjacent to and on either side of the foramen magnum in MNG 8747 is a shallow, 5 mm-long, horizontal groove occupying the presumed, approximate position of the proximal end of the supraoccipital-opisthotic contact. Stapes. In MNG 8853 both stapes are present, but only the right is fully exposed (Fig. 5, 6, 7B), whereas in MNG 8747 only the right is present and exposed (Fig. 9A, 10). In general the stapes is divisible into two parts: an expansive, flat, distal plate that occupies the otic (or temporal) notch and a short proximal shaft with an expanded footplate. Because it could not be determined whether the distal plate represents an ossified tympanic membrane or is completely or partially stapedial, it was termed simply the “ossified plate” by Olson (1966). In MNG 8853 the ossified plate occupies nearly the entire otic notch, with its lateral perimeter reaching to within 3 or 4 mm of and paralleling the otic notch. A short, stapedial shaft projects dorsomedially from the medial margin of the ossified plate at a level about one-fourth the distance down in its height. Below the shaft the dorsal two-thirds of the medial margin of the ossified plate thickens greatly and extends a short distance as it wraps around the medial margin of the quadrate. At the base of the shaft there is a small, hemispherical swelling of the ossified plate. The bluntly pointed ventral end of the ossified plate closely approaches the small, triangular, medial process of the quadratojugal at the posteroventral margin of the otic notch. In the juvenile MNG 8747 the precursor material of the ossified plate had apparently only partially ossified, and the plate occupies only the dorsal half of the otic notch. The dorsal, or posterodorsal, and ventral margins of the plate exhibit smoothly finished edges, whereas the anterior margin is incompletely preserved. The posteroventral corner of the ossified plate thickens as it continues into the medially directed stapedial shaft. The shaft is very short and has an anteroposteriorly elongated, suboval cross section. The footplate is subtriangular in ventral view, with the posteromedial corner being greatly elongated. The medially facing surface of the footplate is slightly concave, with an irregular, roughened surface that suggests it had a cartilage covering. It is reasonable to assume that this surface articulated with the posterolateral surface of the otic trough of the opisthotic, from which it is only very narrowly separated. A small stapedial foramen pierces the posteroventral surface of the shaft. Mandible. The greater part of the right lower jaw of MNG 8747 is preserved and has been separated from the skull (Fig. 11). It is missing large portions of the splenial, angular, and prearticular that border the medial fenestra ventrally and posteriorly, and almost the entire articular. The left jaw is very incomplete, but importantly includes the symphyseal portions of the dentary and splenial, as well as the first four teeth and the roots of the fifth and sixth. This portion has also been separated from the skull (Fig. HE), whereas a badly damaged midlength portion remains attached to the skull. Al- though the lower jaws of MNG 8853 are complete, their attachment to the skull allows only partial exposures of the lateral and medial surfaces. In MNG 8747 the cross-sectional shape of the jaw posterior to the symphyseal region is subcircular, with the mediolateral width being slightly greater than the dorsoventral depth at the level of the coronoid eminence. As a result, the adductor fossa faces almost directly dorsally, and the incompletely preserved medial fossa apparently faced primarily ven- trally. This arrangement nearly prevents the two openings from being visible in a single orientation of the jaw. On the other hand, in MNG 8853 jaw depth very slightly exceeds the width at the same level, and both fossae are clearly visible in medial view. These differences almost certainly reflect different ontogenetic stages. With the exception of proportions, the lower jaw of D. absitus specimens deviates in only a few minor ways from those of North American members of the genus. However, because the only detailed description of a lower jaw of Diadectes (Welles, 1941) was based on an imperfect specimen (UCMP 33903), it is necessary to comment on some aspects of its anatomy in D. absitus. In lateral view of Fig. 11. — Diadectes absitus, paratype (MNG 8747). A-D. Right jDwer jaw in lateral, medial, .dorsal, and ventrolateral views. E. Symphyseal region of left lower jaw in medial view. Scale = 2 cm. 72 Annals of Carnegie Museum VOL. 67 the jaw the surangular tapers anteriorly along its dorsal contact with the angular as a wedge-shaped process to a level about one-fourth the distance back in the length of the angular (Fig. IIA). An unusual feature clearly visible in the right lower jaw of MNG 8747 is the formation by the dentary of a wide, dorsally facing platform lateral to the cheek teeth (Fig. IIC). This structure is also visible in the right lower jaw of MNG 8853; however, it appears to be relatively narrower and possibly bordered by a low, rounded facial ridge of the dentary. The dentary dentition is exposed only in MNG 8747, and, as in the upper dentition, is typical of that in Diadectes. However, the reasons for giving the above brief account of the upper marginal dentition also apply here. The right dentary possesses 15 teeth, although only the bases of the first five, and the 13th and 15th remain; fortunately, the first four teeth of the symphyseal region of the left jaw are well preserved. The first three are distinctly procumbent and incisiform, decrease markedly in size posteriorly, are narrowly separated, and, as in the premaxillary teeth, are broadly concave on the distal portion of their lingual surface, giving them a chisel-like appearance in lateral view. The fourth tooth is smaller than the preceding tooth and intermediate in form between the anterior incisors and the succeeding molar-like teeth. The molar-like cheek teeth of the series increase in size posteriorly to the ninth or tenth, then decrease gradually to the 13th, with the last two decreasing markedly. Their degree of molar-like development clearly reflects a juvenile stage of growth (Berman and Sumida, 1995). The ninth tooth of the series measures 6.0 mm in transverse width, 3.0 mm in anteroposterior length, and 4.0 mm in estimated height (the crown is badly worn). Although transversely expanded, the labial sides of the teeth are positioned well in advance of the lingual sides. The lingual and labial cusps of the cheek teeth are poorly developed and perhaps would be more accurately described as “shoulders,” whereas the subconical central cusps are well developed and account for about half the height of the crown. The lingual cusps appear to be more pronounced than those of the labial side, as the bases of the central cusps are more expanded labially than they are lingually. The coronoid, which was absent in the Diadectes specimen described by Welles (1941), is well preserved and entirely exposed in the right jaw of MNG 8747 (Fig. IIA-C). The coronoid forms almost the entire coronoid eminence of the jaw, which attains a height slightly above the level of the cheek teeth. Although visible in both lateral and medial views of the jaw, the coronoid is more expansive medially and forms the anterolateral border of the adductor fossa. The coronoid eminence is much thicker transversely than the surangular portion of the adductor fossa rim. From the anterior corner of the adductor fossa the coronoid extends anteriorly as a mediolaterally flattened, narrow, rectangular plate that sheaths the medial surface of the alveolar shelf of the dentary while contacting the splenial and prearticular along its ventral margin. Anteriorly the coronoid extends to the level of the sixth maxillary tooth, although there is a small gap at about the level of the 11th tooth. In ventral view of the mandible of MNG 8853 (Fig. 7A) the splenials are exposed in part as narrow bands extending posteriorly from the symphysis along the ventromedial margin of the jaw. At about their midlength they taper posteriorly to a sharply pointed process that penetrates the anterior end of the angular. This results in the splenial being separated from the ventral border of the medial fenestra by a narrow, marginal splint of angular. Just posterior to the symphysis on the medial surface of the right jaw of MNG 8747 the dentary-splenial suture opens narrowly for a few millimeters. The margins of the opening are smoothly rounded, giving it the appearance of a fenestra. Welles (1941) identified and referred to this opening in Diadectes as the anterior fenestra and described it as communicating with the Meckelian canal or, as he preferred, the primordial canal. Axial Skeleton. The complete presacral column of MNG 8853 is visible for the most part in dorsal view (Fig. 1) and includes 21 vertebrae. The atlas-axis complex is complete, although the strong, lateral angulation of the skull to the left side of the vertebral column has resulted in the disarticulation between the atlantal intercentrum-neural arch and the fused atlantal pleurocentrum-axial intercentrum components (Fig. 12). In most of its key features the complex conforms closely with that in North American species (Sumida and Lombard, 1991). Both proatlases are exposed in dorsal view, but only the right one is well exposed and little distorted. Both contact the occiput adjacent to and at the midheight level of the foramen magnum, but it is uncertain that this represents their actual point of contact with the skull. They are disarticulated from and lie just lateral to their respective atlantal neural arch halves. The right proatlas, as seen in dorsal view, consists of two subequal parts: an expanded, subcircular anterior end from whose posterior margin projects a slightly tapering, spine-like process. The anterior expanded end is strongly inclined anterodorsally, so that its ventral surface, undoubtedly the site of an articulation facet, contacts the occiput of the skull. The spine-like posterior portion, the epipophysis, must have projected directly caudally. There is a ventral thickening at the base of the epipophysis, whose ventral surface presumably possesses a facet for articulation with the atlantal neural arch. The atlantal neural arch halves are preserved in very nearly their correct orientation and articulation with one another, the atlantal intercentrum, and the occipital condyle of the skull. The upper halves 1998 Berman et al.- — ^Early Permian Diadectes from Germany 73 axl + atp Fig. 12. — Diadectes absitus, holotype (MNG 8853). A. Posterior region of skull and first five cervical vertebrae in dorsal view. B, Same region as in A exposed on reverse side of block showing atlantal intercentrum and paired atlantal neural arches in posteromedial view, fused atlantal pleurocentrum and axial intercentrum in right ventrolateral view, central elements of cervical vertebrae 2-4 in ventral view, right atlantal rib in medial view, and right ribs 2-4 in lateral view. Scale = 3 cm. of the arches are exposed in dorsal view and appear as a larger version of the proatlas. The dorsal surface of its rounded, anterior end is occupied almost entirely by a flat, anterior zygapophyseal facet for the proatlas. Medial to the facet there is a small, rectangular extension by which the arch half contacted its mate on the midline, whereas laterally (visible only on the right arch half) there is a very small triangular protuberance. Posteriorly the arch halves consist of a long, spine-like process which tapers to a sharp point and, as in the proatlas, is identified as an epipophysis. Posteriorly the 74 Annals of Carnegie Museum VOL. 67 epipophyses diverge slightly from the midline. A ventral thickening at the base of the epipophysis buttresses the posterior zygapophysis. Below the level of the pre- and postzygapophyses the arch halves are visible only in posteromedial view (Fig. 12B). Each consists of a subrectangular plate with its long axis directed ventrolaterally and its internal surface facing strongly posteriorly and slightly ventromedially. The dorsomedial margins of the plates are slightly concave and diverge posterodorsally from the midsagittal plane, as they form the ventral half of the neural canal. The anterior margins clearly contacted the dorsolateral edges of the occipital condyle, whereas the ventral margins have a broad contact with the atlantal intercentrum. A short but vertically elongate diapophysis for the tu- berculum of the atlantal rib projects ventrolaterally and slightly posteriorly from very low on the posterior margin of the lateral surface. In posterior view (Fig. 12B) the atlantal intercentrum is in general crescent-shaped, with the apices of the dorsally directed lateral wings contacting the bases of the atlantal neural arches. The posterior face of the midcentral area is excavated into a broad, shallow concavity. The midventral length of the atlantal intercentrum is very short, perhaps only 2 mm. A short, vertically elongate parapophysis for the capitulum of the atlantal rib projects ventrolaterally from the lateral surface. The anterior rim of the atlantal intercentrum contacts the ventral half of the occipital condyle rim. As previously described (Sumida and Lombard, 1991; Sumida et al., 1992) in North American Diadectes, the atlantal pleurocentrum in MNG 8853 is fused to the dorsal surface of the axial inter- centrum. In lateral view (Fig. 12B) the pleurocentral portion has the appearance of a low, rectangular block. However, strong, laterally oblique crushing has resulted not only in some transverse narrowing of the element, but also in its right lateral surface facing ventrolaterally so as to be visible in ventral view of the complex. The flat dorsal surface of the centrum (Fig. 12A) is of unfinished bone except for a smooth, middorsal channel which ends just short of the anterior margin. The channel may represent the floor of the neural canal. These features suggest that the pleurocentrum was continued dorsally by cartilage that may have ossified later in ontogeny. In ventral view of the complex the intercentral portion has a subtriangular outline with the apex directed anteriorly and a very low, broadly rounded midline surface. The posterolateral corners are extended slightly and end in a truncated margin as the parapophyseal processes for the capitular head of the axial rib. The anteriorly directed apex is exaggerated slightly to form a bluntly rounded extension whose tip likely contacted the midventral concavity on the posterior surface of the atlantal intercentrum. Exposure of the axis in MNG 8853 is limited to the left side of the neural arch and the right side of the centrum, and the two elements appear to be fused (Fig. 12). There is no swelling or lateral expansion of the arch, as in the immediately succeeding vertebrae. The blade-like neural spine is long and low, not exceeding the height of the more posterior spines, and thickens considerably toward its distal and posterior margins. Close to the posterior margin on the lateral surface of the spine is a well- developed, sharp-edged ridge that extends ventrally from the crest to the anterior edge of the base of the postzygapophseal buttress. The zygapophyses are set close to the midline, although the posterior set is slightly wider apart. Whereas the anterior zygapophyses and their facets are oval, the posterior set are narrowly subrectangular. Both sets of facets slope slightly ventrolaterally. The partially exposed centrum exhibits a moderately developed midventral keel. In MNG 8853 the intercentrum of the third cervical (Fig. 12B) is about two-thirds the size of that of the axis and is weakly chevron-shaped with the apex directed anteriorly, as reported in Diadectes (Sumida and Lombard, 1991). Intercentrum 4 and those of succeeding vertebrae are much smaller and have a laterally expanded oval outline. The postaxial neural spines (Fig. 1) are low and roughly diamond-shaped in horizontal section, with the long axis oriented anteroposteriorly. They increase slightly in transverse width from the third to the seventh vertebra, becoming nearly quadrangular in horizontal section, then decrease in width gradually in the succeeding seven or eight vertebrae to a narrow diamond outline that is essentially maintained to the end of the presacral series. The summits of the spines are very coarsely textured, giving them an irregular margin. This is particularly true of those in the anterior half of the presacral series, where the summits are slightly expanded, giving the spines a slightly mushroom-like appear- ance. Additionally, in this region of the column there is also a bilateral constriction of the neural spine close to the posterior corner of its summit, so that in dorsal view this angle of the spine is converted into a process-like structure. A further variation in spine structure is seen in the vertebrae near the posterior end of the dorsal series in MNG 7721, where the lateral, angular margins of the spines are expanded laterally into short, irregular processes (Fig. 1, 13C). One noticeable exception to the general structure of the postaxial dorsal neural spines given above is that of the fourth vertebra in MNG 8853 (Fig. 1, 12A), which is greatly reduced in both its lateral and longitudinal dimensions to a nearly blade-like structure (a similar condition is seen in the amphibian Eryops; Moulton, 1974). Seemingly, longitudinal compression of this region of the column has brought the neural spines of these three vertebrae into contact. 1998 Berman et al, — -Early Permian Diadectes from Germany 75 Fig, 13.— Diadectes absitus, paratype (MNG 7721). A, Postaxial cervical and C, D, posterior dorsal vertebrae in lateral and ventral views (anterior to the left), respectively. Scale = 4 cm. 76 Annals of Carnegie Museum VOL. 67 The neural arches of all the presacrals of the postcervical region (Fig. 1) are swollen, with the zygapophyses extending laterally well beyond the lateral margins of the centra. In the first nine vertebrae there is a marked, successive increase in the lateral extent of the zygapophyses from very close to the midline in the atlas-axis complex to nearly equalling the lateral extent of the transverse processes. More posteriorly, the presacrals exhibit only a very gradual increase in this dimension. The low, squat structure of the presacrals is expressed particularly in the near-horizontal plane occupied by the dorsal crests of the posterior zygapophyseal buttresses. The zygapophyseal planes in the cervical region appear to slope slightly posteroventrally, whereas those of the rest of the presacrals are hori- zontal. Lateral and ventral views of the presacral vertebrae have been possible only in two short series in MNG 7721 (Fig. 13): one includes five postaxial cervicals lacking mainly the neural spines and the other includes six dorsals (only five are shown) from just anterior to the sacrum. Their differences are reflected mainly in the structure of the transverse processes. In the vertebrae from the cervical region the transverse processes are laterally projecting, anteroposteriorly thin, and dorsoventrally elongate ridges. In lateral view they extend diagonally anteroventrally from the posterior surface of the anterior zygapophysis to the upper portion of the anterior centrum rim. The narrow costal facets for the ribs face ventrolaterally and slightly posteriorly and show no noticeable expansions demarking capitular and tubercular areas. In end view of the vertebrae the processes shorten gradually in lateral extent as they extend anteroventrally, but still end with a rather long, nearly horizontal ventral margin. The transverse processes in the string of posterior dorsals are thicker, project directly laterally, and are restricted to the posterior surface of the anterior zygapophyseal buttress and do not extend onto the centrum. Their wider costal facets face mainly laterally and slightly posteroventrally and are slightly constricted into nearly equal capitular and tubercular regions, the former being a little wider. Disar- ticulation of the last or sixth vertebra of the string of far posterior dorsals reveals the absence of hypantrum and hyposphene accessory articulations. The last two vertebrae of the string are deeply amphicoelous, but do not appear to be notochordal. In some of the vertebrae there is an indication of a neurocentral suture which may suggest a less than fully mature condition. Except for the atlas-axis complex, the centra of the presacral vertebrae are spool-shaped, with horizontal diameters of the ends exceeding the lengths in the cervicals and farthest posterior presacrals in MNG 7721 by approximately 40 and 18%, respectively. The lateral surfaces of the centra are flared moderately outward to thickened, semicircular rim lips. Flaring of the ventral surface of the centrum is much less pronounced, and there are no midventral ridges. The midventral margin of the posterior rim of the centrum is bevelled to accommodate the small intercentrum. All that can be seen of the two sacral vertebrae in MNG 8853 (Fig. 1) are the poorly preserved neural arches. However, a large portion of the tail of MNG 8853 is preserved (Fig. 1,14) and includes a string of 18 vertebrae, although nine are represented only by intercentra or haemal arches. As preserved, the first five vertebrae of the series are represented by only the intercentra in dorsal view. They have a narrow, oval outline and are aligned as if retaining their original spacing and orientation. The caudal series is continued posteriorly by nine complete, laterally exposed vertebrae that diminish steadily in size posteriorly. Their neural arches are narrow, not swollen, with the zygapophyses po- sitioned close to the midline. The anteriormost neural spine of the series curves slightly posteriorly and is blade-like for most of its length, as it terminates distally in a cap-like lateral expansion that is narrowly oval in dorsal view. More posteriorly the spines gradually straighten to a vertical orientation, decrease dramatically in size, and quickly reduce and lose the cap-like expansion at their distal end. The oval zygapophyseal planes slope steeply ventromedially. The transverse processes are exposed only in dorsal view and are positioned at about midheight on the centra and adjacent to the anterior central rim. They exhibit a marked, successive decrease in size posteriorly. In the anteriormost pre- served neural arch, vertebra 6 of the series, the transverse process is rather broad and long, projects directly laterally, and narrows slightly distally along its posterior margin. By the posteriormost pre- served neural arch, vertebra 14 of the series, the process is reduced to a very small, ventrolaterally projecting, triangular nubbin. The centra of this series are spool-shaped with moderately concave lateral surfaces. The lips of the central rims are laterally flared, but that of the posterior rim is far more pronounced. The ventral margins of the anterior central rims are strongly bevelled to accommodate the haemal arches. The small, proximal, intercentral portions of the haemal arches are narrowly rect- angular in lateral view. As the long spines extend from the posteroventral corner of the intercentral portion they gradually double in anteroposterior width by their distal end. Directly following the nine complete, articulated caudals the remaining, preserved portion of the tail ends in a series of four haemal arches that also retain their approximate spatial relationships and alignment with the preceding arches. Their dorsal and lateral surfaces are exposed, as well as the proximal portion of their posterior surface. Additionally, the entire posterior surface of a disarticulated haemal arch is exposed in MNG 7721. In posterior view the haemal arch is Y-shaped, with the 1998 Berman et al, — Early Permian Diadectes from Germany 77 Fig. 14. — Incomplete tail of Diadectes absitus, holotype (MNG 8853; see also Fig. 1). Anteriormost and posteriormost vertebrae of series represented only by intercentra and haemal arches, respectively. Scale = 4 cm. 78 Annals of Carnegie Museum VOL. 67 proximal, crescentic intercentral crosspiece spanning the ends of the two arms of the Y to enclose a triangular space. The roughened, posterodorsal articular surface of the intercentral crosspiece contacted the bevelled ventral rim of the centrum. The spine ends in a laterally flattened, blade-like structure. An isolated string of four distal caudals with haemal arches is exposed in left lateral view a short distance to the right of the midlength region of the presacral column of MNG 8853 (Fig. 1). Their proximity to MNG 8853 and similarity to those of Diadectes leaves little doubt that they belong to this specimen. By way of comparison with Case’s (1911) description of Diadectes, their serial place- ment is probably near caudal 25. The neural spines are very small, dorsally directed, and laterally flattened with truncated tips. The zygapophyses are set very close together, and their oval facets are inclined very steeply ventromedially. Transverse processes are absent. The lateral surfaces of the centra are moderately concave, and the ventral margin of the posterior rim is bevelled to accommodate the haemal arch. The haemal arches are shorter versions of those more anterior in the caudal series. A complete set of presacral ribs is present and well preserved in MNG 8853 (Fig. 1), but a tightly overlapping arrangement permits adequate description of only those of the atlas and axis (Fig. 12B). The complete right atlantal rib is exposed in posterior or medial view and closely approximates its correct relationship with the atlantal neural arch and intercentrum. In this view the head and shaft, which are subequal in length, form a flat, planar surface. The broadly triangular rib head was probably dicephalous, as a weakly ossified region roughly defines a narrow gap between the well-developed capitulum and the much narrower tuberculum. The short shaft maintains a narrow width that curves slightly posteriorly to its transversely truncated distal end. The right axial rib is complete, exposed in anterior or lateral view, and nearly retains its proper association with the axis. It is essentially a larger version of the atlantal rib except for the gradual, distal expansion of the shaft to a width approximately 50% greater than its proximal end. There is a continued, progressive increase in the expansion of the rib shafts 3-7, with those of vertebrae 6 and 7 having a roughly banana-shaped outline. In the re- maining presacral ribs only the dorsal margins of the tubercula are visible, and the shafts maintain a nearly constant, narrow width throughout their length. The shafts exhibit a gradual increase in length to about the 1 4th rib, then decrease to a length nearly equalling that of the atlantal rib. Whereas the first four ribs are only slightly curved, the succeeding rib shafts exhibit a moderate posterior curvature that becomes pronounced in the last four or five presacral ribs. Shoulder Girdle. All the elements of the shoulder girdle are well represented except the interclav- icle. In the holotype the right scapulocoracoid and cleithrum are exposed below the cervical region of the skeleton (Fig. 2, 15), whereas the right clavicle was found isolated above the cervical region and was subsequently removed (Fig. 16). In the paratypes MNG 7721 and 8778 only remnants of the articulated interclavicle, clavicles, and left scapulocoracoid are preserved (Fig. 16B). The holotypic right cleithrum (Fig. 15) is complete, exposed in lateral view, and very nearly in its correct articulation along the anterior margin of the scapulocoracoid. The cleithrum of Diadectes absitus very closely duplicates that seen in North American specimens (Case, 1911; Romer, 1956). In lateral view it has a narrow, sickle-shaped appearance. The expanded, posterodorsally curving upper half of the element has separated slightly from the anterodorsal corner of the scapular blade, which it presumably overlapped laterally. The much narrower ventral stem of the cleithrum maintains a constant width as it extends ventrally, terminating at the level of the upper margin of the glenoid cavity. As shown by the posterior view of the complete right holotypic clavicle (Fig. 16A), the stem and ventral plate meet in a sharply defined internal angle of 120°. The ventral plate is bowed ventrally, with much of the posterior portion turned upward abruptly into a high, transverse, vertical wall that quickly diminishes in height as it joins the dorsal stem. In ventral view (Fig. 16B) the ventral plate is narrow and triangular, and a smooth, narrowly triangular area of the posteromedial corner is de- pressed dorsally. The plate ends medially in a feathered edge. The narrow dorsal stem gradually tapers distally to a blunt point. A moderate, rounded ridge extends the entire length of the medial edge of the posterior surface of the stem. Lateral to the ridge the posterior surface of the stem forms a shallow channel that probably received the ventral stem of the cleithrum. Although the right holotypic scapulocoracoid is complete, only the scapular blade and coracoid region anterior to the glenoid cavity are well exposed (Fig. 15). A suture cannot be found between the scapular blade and coracoid plate. The scapular blade is tall and narrow, and except for the weakly ossified, obliquely truncated anterodorsal corner, is subrectangular in outline. Most unusual, but typical of North American species (Romer, 1956), is the absence of a gradual, but pronounced posterior distal expansion of the blade. The full extent of the coracoid plate is clearly seen in MNG 7721 (Fig. 16B) by either remaining bone or lighter-colored, reduced areas of the matrix that indicate where bone had been present. The coracoid plate is strongly expanded anteroventrally into a smoothly curving, almost semicircular margin. The long, narrow glenoid cavity is essentially complete and extends directly to the posteriormost point on the margin of the scapulocoracoid. As is typical of early tetrapods, the glenoid is screw-shaped, facing posterolaterally and slightly ventrally at its anterior end and dorsally 1998 Berman et al. — Early Permian Diadectes from Germany 79 Fig. 15.- — Diadectes absitus, holotype (MNG 8853), Right cleithram and scapulocoracoid in lateral view (ventral margins toward top of page) and right humerus with distal expansion in ventral view (see also Fig. 2). Scale = 2 cm. 80 Annals of Carnegie Museum VOL. 67 1998 Berman et al. — Early Permian Diadectes from Germany 81 at its posterior end. A well-developed supraglenoid buttress supports the anterior end of the glenoid. Located beneath the anterior half of the glenoid is a partially preserved deep fossa. At the anterodorsal margin and deepest level of the fossa is a remnant of the coracoid foramen. Forelimb and Manus. The humerus is best exemplified by the right element of the holotype MNG 8853 (Fig. 15), where it is exposed with the distal expansion in ventral view. The humerus has undergone some dorsoventral crushing and erosion of the surface bone. Because the humerus conforms very closely to those described (Case, 1911; Romer, 1956; Sumida, 1997) in North American Dia- dectes, only a few minor comparisons are required. The humerus of MNG 8853 appears to differ from that of the North American species in having 1) a slightly greater development of the crest of the proximal head for the attachment of the pectoralis muscle, 2) a greater distal expansion of the supinator process so that the angle formed between it and the capitellum is greater, 3) a more strongly distally hooked ectepicondyle, 4) a more transversely expanded trochlea, and 5) a slightly greater truncation of the posterodistal comer of the entepicondyle. The only radius and ulna complete enough to warrant description are those articulated with the right manus in MNG 7721 (Fig. 17); an incomplete left epipodial pair is preserved in MNG 8978. Seen in dorsal (= anterior) view the radius has a relatively narrow, long shaft with moderately expanded ends. The proximal end is far more expanded than the distal end, and both expansions are slightly greater on the lateral margin of the bone. For most of its proximal length the cross-sectional outline of the radius is a mediolaterally expanded oval, with the medial margin flattened and the lateral margin drawn out into a narrowly pointed edge. This outline is continued to the concave proximal articular surface, whereas the poorly preserved distal end appears to have a subrectangular articular surface with the dorsoventral dimension being slightly greater. The ulna is essentially complete in MNG 7721 except for the loss of the olecranon process. The complete bone may have been as much as 25% longer than the radius. What remains of the sigmoid notch is well defined. The bone is dorsoventrally (anteroposteriorly) narrow and greatly expanded mediolaterally. Because the ends are much more strongly flared medially than laterally, the medial margin is deeply concave. The dorsal surface of the distal end is very slightly convex. A dorsoventrally narrow articular surface, occupying the entire end of the bone, is partially divided by a shallow excavation on the ventral surface that approximates the distal margin angulation. Both distal facets face slightly ventrally, with that for the intermedium facing slightly medially and that for the ulnare facing slightly laterally. The manus of MNG 7721 is nearly complete (Fig. 17), and the few incomplete elements, all phalanges, can be easily restored. The carpals are well articulated with no noticeable intervening spaces, suggesting a mature stage of development. The manus is exposed in dorsal view, but with the distal phalanges of digits 1-4 hyperflexed against the plantar surface. As far as known, this is the first complete carpus of Diadectes to be described, and it includes the expected elements, although there is no indication of a medial centrale or a pisiform ossification. The radiale is trapezoidal in outline and very thick, with a broad, flat proximal articular surface that appears to match that of the radius. The intermedium is subrectangular, but with a wedge-shaped medioproximal extension of the deep, oval ulnar articular margin. The oval ulnare is the dominant element of the carpus. Dorsal expansion of the proximal and distal margins produces a shallow, depressed central area. The lateral or proximal centrale is pentagonal in outline, with substantial contacts with the intermedium, ulnare, and distal carpals 2-4, and its proximomedial corner contacts narrowly the radiale. A series of five distal carpals is clearly defined. Distal carpal 1 appears as a small, equilateral triangle in outline and may have been displaced slightly laterally on its contact with the first metacarpal. The second distal carpal is a narrow, proximodistally elongated oval with the proximal end being wedge-shaped. Because distal carpal 2 contacts not only nearly the entire medial margin of the lateral centrale, but also the distolateral corner of the radiale, it is suspected that it may also include or has expanded to exclude the medial centrale. Distal carpal 3 is small and trapezoidal in outline. Distal carpal 4 is basically pentagonal in outline, with a proximal angulation that wedges partially between the medial centrale and the ulnare and a broad distal margin contact with metacarpal 4. The medial angulation of the fourth distal carpal contacts the lateral centrale and the third distal carpal, and its narrow, truncated lateral margin contacts the fifth distal carpal. The small, distal carpal 5 is a proximodistally short, lens-shaped element. An unidentified, subcircular element about 8 mm in maximum diameter and partially visible between Fig. 16. — Diadectes absitus. A, B. Right clavicle of holotype MNG 8853 in posterior and ventral views. C. Incomplete left scapulocoracoid of paratype MNG 7721 in lateral view (missing portions of bone indicated by reduced matrix). Scale = 2 cm. 82 Annals of Carnegie Museum VOL. 67 Fig. 17. — Diadectes absitus, paratype (MNG 7721). Articulated right radius, ulna, and manus in dorsal view. Distal phalanges of digits 1-4 are hyperflexed against plantar surface of manus and shown separately. Scale = 4 cm. metacarpals 2 and 3 is suggestive of a carpal. Of the two carpal elements not accounted for in MNG 7721, however, a pisiform seems the most likely identification, as the tightly articulated carpus could not have accommodated a medial centrale. Metacarpal 1 is very short and squat, whereas 2-5 are substantially larger and exhibit a gradual lengthening and narrowing serially. With two exceptions all five digits of MNG 7721 are well rep- resented. The two phalanges of the first digit were lost due to weathering and were restored using their enclosing matrix as a natural mold to cast them in epoxy. Secondly, incomplete preservation makes it impossible to determine whether the fifth digit consisted of two or three phalanges. The phalangeal formula is, therefore, 2-3-4-5-2 or 3, but the higher count is suspected to be correct, as it is the usual formula of late Paleozoic terrestrial vertebrates. As is typical in Diadectes, all of the phalanges are short and broad, with the terminal ones ending in a bluntly rounded margin. An unusual Diadectes feature that is also present in MNG 7721 is the extreme proximodistal compression of the penultimate phalanges of digits 2-4. Pelvis. The holotypic pelvis MNG 8853 is complete, although the ilia have suffered severe enough crushing to prevent detailed description (Fig. 1, 2, 18A). However, in MNG 8978 the disarticulated right ilium is complete and well exposed in lateral view (Fig. 18B). The anterior margin of the iliac blade is nearly vertical, with only a very slight anterodistal curvature. The posterior margin, however, is drawn out into a short, broadly triangular process that ends only a very short distance beyond the level of the expanded base of the ilium. There is no indication of an external iliac shelf, although such a structure is visible but poorly defined in the holotype. The long, quadrangular holotypic pu- boischiatic plate is well preserved and exposed in ventral view, with its lateral halves sloping ventro- medially slightly to their midventral union to form to a modest keel. The plate is strongly waisted by the semicircular ventral rims of the acetabula, which occupy 44% of the plate's length. The union of the pubis and ischium cannot be detected, but is assumed to have occupied its normal position at about the midlength level of the acetabulum. If this is correct, the ischium is about twice the length of the pubis. Whereas the paired pubes end anteriorly in a low, triangular margin, the slightly convex posterior margins of the paired ischia form a wide, V-shaped notch in the plate. The anteroposteriorly 1998 Berman et al. — Early Permian Diadectes from Germany 83 Fig. 18. — Diadectes absitus. A. Puboischiatic plate and femur of holotype MNG 8853 in ventral view (see also Fig. 2). B, C. Lateral view of right ilium and dorsal view of left femur of paratype MNG 8978. Scale = 4 cm. 84 Annals of Carnegie Museum VOL. 67 Fig. 19. — Diadectes absitus. A. Right tibia, fibula, and proximal portion of pes of holotype MNG 8853 in dorsal view (see also Fig. 1). B, C. Isolated left fibula of paratype MNG 7721 in dorsal and ventral views (distal end toward top of page). Scale = 4 cm. elongate obturator foramina, defined by poorly preserved margins, lie midway between the antero- ventral margin of the acetabulum and the midventral union of the pubes. Hindlimb and Pes. Femora are best preserved in the holotype MNG 8853 and MNG 8978, where the right of the former and the left of the latter are exposed in ventral and dorsal views (Fig. 18A, C), respectively. Both femora are complete, but the ends of MNG 8853 are not entirely exposed and those of MNG 8978 have been severely crushed dorsoventrally. The heads are widely expanded, subequal in length, and joined by a very short shaft in MNG 8853. In the smaller MNG 8978, however, there appears to be no intervening shaft. Dorsal view of the femur MNG 8978 shows the proximal and distal heads as angled posteriorly, so that the anterior margin of the bone is straight or slightly concave and the posterior margin is strongly waisted. The dorsal surface of the proximal head is broadly convex, and the narrow, terminal articular surface has a slightly crescentic outline, convex above, that thins more gradually posteriorly. As is customary, of the two distal condyles the posterior one is larger and extends a short distance farther distally. On the ventral surface of the femur MNG 8853 the depth of the intertrochanteric fossa is exaggerated along its anterior margin by a welF developed internal trochanter. The internal trochanter is continued to the distal corner of the fossa and onto to the shaft, where it becomes much more pronounced as the fourth trochanter. The best examples of the tibia and fibula are the right elements in the holotype MNG 8853 (Fig. 19A), exposed in dorsal (= anterior) view, and an isolated left fibula in MNG 7721 (Fig. 19B, C). Although other examples of these elements are present in MNG 7721 and 8978, they are too poorly preserved to be informative. Both elements in the holotype are strongly crushed dorsoventrally ( = anteroposteriorly), and most of the lateral half of the proximal head of the tibia is hidden by the overlying fibula. The fibula exceeds the tibia in length by 7%. Because the ends of the tibia are much more flared laterally than medially, the lateral margin of the bone is deeply concave, whereas the medial margin is only slightly concave. Despite the extreme crushing, the cnemial crest can be seen to end proximally in a prominent, knob-like rugosity. The heads of the fibula are much more greatly 1998 Berman et al. — Early Permian Diadectes from Germany 85 expanded medially than laterally, so that, whereas the lateral margin of the bone is only slightly concave, the medial margin is strongly concave with essentially no separation of the heads by a shaft. The proximal head terminates in a very slightly convex margin that is directed slightly posteromedially (= dorsomedially), whereas the moderately convex margin of the distal head is oriented slightly anteromedially (= ventromedially). As seen in MNG 7721, a very low ridge on both the dorsal and ventral surfaces of the distal head lies close to and parallel with the lateral margin. The ventral surfaces of both heads are broadly concave. The proximal articular surface is crescentic in outline, with the concave margin ventral, and narrows slightly from a thicker, bluntly rounded lateral margin. The distal articular surface is weakly sigmoidal, with a slight constriction dividing it into a thicker, ventrally convex medial portion for the astragulus and a thinner, dorsally convex lateral portion for the astragulus and calcaneum. Although elements of the pes are present in the holotype MNG 8853 (Fig. 1, 19A) and MNG 7721, they are so jumbled and poorly preserved that they are of little descriptive value. The dorsal ( = anterior) surface of the astragulus is, however, clearly visible in MNG 7721 (not figured) and exhibits the standard L-shaped outline. The dominant, subrectangular horizontal limb occupies about 80% of the proximodistal length of the astragulus and exceeds by 2.4 times the mediolateral width of the much smaller neck. The rectangular outline of the neck is altered by a broadly rounded proximolateral corner for contact with the fibula. At the distal end of the lateral margin opposing the calcaneum is a small, semicircular notch of the perforating foramen. A channel-like depression extends proximo- medially from the foramen to the posteromedial angle of the astragulus. Just distal to the foramen the distolateral comer of the astragalus is beveled for contact with the fourth distal tarsal. Comparisons and Discussion European Diadectids Of the vertebrates described from the Lower Permian of central Europe, only two genera can be assigned to Diadectidae with confidence, and both are from the Lower Rotliegend of Germany. On the basis of a string of six vertebrae that includes the last four presacral and two sacrals from the Leukersdorf Formation of the Erzgebirge Basin near Zwickau, Meyer (1860) described Phanerosaurus naumanni. The vertebrae are very much like those of Diadectes, having very broad, massive neural arches with far laterally placed zygapophyses. A second probable diadectid was described by Geinitz and Deichmueller (1882) from Nied- erhaeslich near Dresden in the Doehlen Basin. They described the disarticulated remains of what they believed to be two specimens representing a new species of Phanerosaurus, P. pugnax, preserved on a series of part and counterpart slabs. These remains were more thoroughly redescribed by Stappenbeck (1905), who concluded that they belonged to only one individual. On the basis of what he interpreted as differences in their vertebrae from those of P. naumanni, P. pugnax was reassigned to a new genus, Stephanospondylus, and both genera were placed in the new family Stephanospondylidae. Stappenbeck’s description of S. pugnax, which included a reconstruction of both the skull and postcranium, was strongly criticized by Romer (1925). Romer convincingly demonstrated that the remains assigned to S. pugnax belong to two distinct forms, a diadectid closely allied to Diadectes and a labyrinthodont amphibian, possibly Onchiodon. Those elements recognized as belonging to S. pugnax include upper and lower tooth-bearing jaw elements, a squamosal and articulated parietals and frontals, presacral vertebrae, expanded ribs of the subscapular region, and elements of the pectoral girdle and forelimb. If, according to Romer (1925:458), Stephanospondylus is defined by the above reduced list of elements, it “emerges as a typical diadectid, differing markedly in no known character from the well-known American forms, although a more com- plete knowledge of the animal might show differences to be present.” Stappen- beck (1905) presented a list of seven vertebral characters for distinguishing be- 86 Annals of Carnegie Museum VOL. 67 tween Phanerosaurus and Stephanospondylus and on this basis established the latter genus for P. pugnax. Romer (1925) regarded these characters, however, as either trivial, erroneous, or of uncertain value and, therefore, as providing little basis for recognizing two genera. Again, he cautioned that the discovery of ad- ditional remains of Phanerosaurus may indicate its distinctiveness. Therefore, Romer recommended that the genus Stephanospondylus be retained for the present and, although difficult to define, offered the following characterization (p. 459): . .a small diadectid, probably about 1 meter in length, lacking a hyposphene- hypantrum articulation; the subscapular ribs, although very broad, lack the V- shaped posterior angles found in Diadectes; the molar teeth lack lateral cusps.” The potential usefulness of this diagnosis would possibly be even further dimin- ished if it were determined that the holotype was a juvenile specimen. The sec- ondary vertebral articulations and the greater development of the subscapular ribs may only appear in larger, more adult individuals, where structural support is more critical; both these structures also appear to be absent in the smaller Bro- macker Diadectes. The absence of lateral cusps on the molar-like cheek teeth in Stephanospondylus could also be explained as a juvenile feature. Berman and Sumida (1995) have shown that in early juvenile stages of growth the cheek teeth in Diadectes are bulbous and lack lateral cusps. Although weakly developed, lateral cusps are, however, present in the juvenile Bromacker Diadectes skull MNG 8747. Yet, the length of this skull (ca. 100 mm) is far less than that of Stephanospondylus pugnax, estimated by Romer (1925) on the basis of the jaw elements to be about 155 mm. There is one other piece of evidence to suggest that Stephanospondylus and the Bromacker diadectid are distinct forms. In addition to the tooth-bearing jaw elements, Geinitz and Deichmueller (1882) also figured the squamosal and the articulated parietals and frontals, which were used by Stappenbeck (1905) in his reconstruction of the skull of S. pugnax. These elements were also suspected by Romer (1925) to belong to S. pugnax. The surface ornamentation on each of the parietals and frontals, however, consists of narrow ridges radiating out from a central growth region. This pattern is most like that seen in typical labyrinthodonts and quite distinct from the network of pronounced, deep, smooth, U-shaped grooves superimposed on a coarsely granular or sponge-like textured sculpturing exhibited by the skull roofing bones in Diadectes (Olson, 1950; Berman et aL, 1992), including the Bromacker skulls. In the absence of more complete materials Phanerosaurus and Stephanospondylus should be considered as very poorly de- fined, although they obviously share certain features with Diadectes. In fact, it is suspected that further examination of the known specimens would indicate that both genera should be considered as nomina dubia. North American Diadectes Although there can be no doubt of the generic assignment of the Bromacker diadectid to Diadectes, certain problems arise when attempting to compare it to North American representatives of the genus. In a study of the systematics of Diadectidae, Olson (1947) reduced significantly the number of recognized Dia- dectes species to three, possibly four, even after synonymizing several genera with this genus. Despite a marked reduction in the number of recognized Diadectes species, the basis for distinguishing between the remaining North American forms relies almost entirely on the extremely questionable grounds of size, proportions, 1998 Berman et al. — Early Permian Diadectes from Germany 87 and stratigraphic position. Further complicating the systematics of this genus are Olson’s (1947, 1950) observations that individual variations within a species are often greater than those between species, the tracing of sutures of the skull roofing bones is often difficult or impossible, and proportional differences are probably often the result of differential growth rates. The lack of detailed descriptions of early growth stages of both the skull and postcranium is particularly troublesome in recognizing differences between North American species of Diadectes (Olson, 1947; Berman et al., 1992; Berman and Sumida, 1995). Undoubtedly due in part to these problems, the numerous descriptions and reconstructions of the skulls of North American specimens (Huene, 1913; Gregory, 1946; Olson, 1947, 1950; Watson, 1954; Lewis and Vaughn, 1965) differ strikingly and contain serious errors in interpretation (Olson, 1947; Berman et al., 1992). Despite the above difficulties, several autapomorphic and plesiomorphic cranial characters can be recognized that distinguish Diadectes ab situs from all North American members of the genus. Polarities have been determined using the sey- mouriamorph Seymouria and anthracosaurs as the reference outgroups for com- parisons, which follows most recent studies that recognize them as the closest outgroups to the diadectomorphs (Gauthier et al., 1988; Panchen and Smithson, 1988; Berman et al., 1992; Laurin and Reisz, 1995). Most recently, however, Carroll (1995) and Laurin and Reisz (1997) have presented a radically new scheme of tetrapod relationships which recognizes the lepospondyls as the nearest outgroup of the diadectomorphs and amniotes. Because of the narrow scope of relationship being tested and of the charcters being analyzed here, the alternative choice of lepospondyls as the nearest outgroup to diadectomorphs produces either ambiguous, the same, or no results. Because the Late Pennsylvanian diadectid Desmatodon is regarded to be a closely related predecessor of Diadectes (Vaughn, 1969, 1972; Berman and Sumida, 1995), it has also been included as a reference outgroup. Dorsal Process of Premaxilla. — ^In D. absitus the dorsal process of the pre- maxilla is long and narrow, and extends a considerable distance onto the dorsal surface of the skull, reaching a level just beyond the posterior margin of the external naris. In all the reconstructions of North American Diadectes but that presented by Gregory (1946), the premaxilla is depicted as lacking a pronounced dorsal process. The presence of a well-developed dorsal process in the more primitive Desmatodon (Berman and Sumida, 1995), as well as in Seymouria, suggests that this is a plesiomorphic feature of D. absitus. Anterior Extent of Prefrontal. — The prefrontal in D. absitus extends well be- yond the level of the anterior margin of the frontal, whereas in North American Diadectes the prefrontal and frontal end anteriorly at approximately the same level. Inasmuch as a prefrontal extending beyond the anterior margin of the frontal is also seen in Seymouria and anthracosaurs, this feature is considered as ple- siomorphic in D. absitus. Shape of Postfrontal.— Tho, postfrontal of D. absitus is chevron-shaped, with its posterior apex penetrating deeply into the anterior margin of the parietal. This configuration is due in part to a greater lateral extension of the postfrontal along the posterior margin of the orbit. In contrast, this element in North American Diadectes is generally subrectangular except for a lateral incisment by the orbital rim and has a straight or slightly posteriorly convex line of contact with the parietal. As the shape of the postfrontal in D. absitus is not seen in other late Paleozoic tetrapods, its presence in D. absitus is judged an autapomorphy. 88 Annals of Carnegie Museum VOL. 67 Shape of P ostorbitaL— TypicsMy the postorbital in North American Diadectes is shown as having a subrectangular outline, with a broad entrance into the orbit and a broad contact with the supratemporal. In D. absitus, however, the postorbital has a very narrow entrance into the orbit (due partly to the encroachment of the postfrontal) and forms for most of its posterior extent a tapering, triangular process that ends in a narrow contact with the supratemporal. By way of comparison with other late Paleozoic primitive tetrapods, the restricted entrance of the postorbital is considered an autapomorphic feature in D. absitus, whereas its posteriorly nar- rowing, triangular shape is judged as plesiomorphic. In view of the fact that early tetrapods lack a postorbital-supratemporal contact, the narrow contact of these two elements in D. absitus is judged a plesiomorphic character relative to the broad contact in North American representatives of this genus. Ventral Margin of Jugal. — The abrupt, dorsalward, step-like retreat of the an- terior portion of the jugal’s contribution to the ventral margin of the skull in D. absitus is considered an autapomorphic feature. Among late Paleozoic tetrapods the ventral margin of the jugal either arches smoothly anterodorsally, as in North American species of Diadectes, or is straight. Basicranial Joint. — The basicranial joint in anthracosaurs, Seymouria, Des- matodon, and immature specimens of North American Diadectes is obviously open and mobile, whereas in mature North American specimens of Diadectes the joint is firmly fused (Olson, 1947; Vaughn, 1972; Berman and Sumida, 1995). Although it is not unexpected that the basicranial joint is open and mobile in the juvenile specimen of D. absitus (MNG 8747), the retention of this feature in the adult Bromacker specimen (MNG 8853) can only be considered as a plesiom- orphic state. Cross-sectional Shape of Lower Jaw. — In D. absitus the lower jaw is subcir- cular in cross section for much of its length. At the level of the coronoid eminence in MNG 8747 the mediolateral width of the lower jaw is slightly greater than the dorsoventral depth, whereas in MNG 8853 the reverse is true. In contrast, the lower jaws of North American Diadectes are extraordinary for their exaggerated vertical expansion not only at the level of the coronoid eminence, but throughout almost their entire length. As examples, the lower jaws of D. sanmiguelensis (MCZ 2989), D. lentus (FMNH UC 675), D. lentus (UCMP 33903), and Dia- dectes sp. (UCMP 59023), which have lengths of 8.8, 13.2, 16.0, and 25.5 cm, respectively, the depth exceeds the width by about 2.7 times at the level of the coronoid eminence. The value for this ratio for the lower jaw of Desmatodon hesperis (CM 47670), measuring 12.0 cm in length, is about 1:2.2 and, therefore, more closely approaches the value for North American Diadectes than D. absitus. In anthracosaurs (Panchen, 1970) the depth-to-width ratio at the level of the cor- onoid eminence undoubtedly approaches that in North American species of Dia- dectes, although rough estimates of the value of this ratio in Seymouria sanju- anensis specimens CM 28596, 28597, and 38022 is only about 2.0. This survey suggests that the relatively low, approximately 1:1 depth-to-width ratio of the lower jaw in D. absitus should be judged as an autapomorphic character. iMbial Parapet in Lower Jaw.-\n North American species of Diadectes, in- cluding early growth stages, and at least mature specimens of Desmatodon, there is lateral to the bases of the cheek teeth a shallow groove whose outer wall is formed by a vertical extension of the lateral surface of the dentary into a low, thin ridge or labial parapet (Welles, 1941; Lewis and Vaughn, 1965; Berman and Sumida, 1995). Among Paleozoic tetrapods a labial parapet of the dentary is 1998 Berman et al. — Early Permian Diadectes from Germany 89 unique to the North American diadectids Diadectes and Desmatodon. Although the lower jaws of both the juvenile and adult D. absitus specimens MNG 8747 and 8853 lack a labial parapet of the dentary, they possess in its place the equally unique feature of a wide, flat, dorsally facing platform lateral to the cheek teeth. Because of the likelihood that these two features are functionally related, the combined characters of an absence of a labial parapet and the presence of a wide, dorsally facing labial platform is judged an autapomorphy of D. absitus. The above combination of autapomorphic and plesiomorphic characters of Dia~ tectes absitus provides a much broader basis for defining this species than is available for distinguishing between those of North America. The greater unique- ness of D. absitus may reflect its wide geographic separation from the North American species. On the other hand, its greater primitiveness reinforces the as- sessment, reached previously on the basis of the character of the entire Bromacker assemblage (Berman and Martens, 1993; Sumida et al., 1996), that the biostrati- graphic position and age of the Tambach Formation, lowermost formational unit of the Upper Rotliegend, should be considered earliest Permian Wolfcampian. Contradictory Descriptions There exist at least two sources of important, misleading information about the anatomy of the North American species of Diadectes that contradict the descrip- tion given here for D. absitus. In a recent redescription by Berman et al. (1992) of the skull of North American Diadectes, based almost entirely on a single specimen (CM 25741), the temporal region was misinterpreted. Most importantly, in their description the supratemporal was mistakenly interpreted to have an ex- tensive, medially directed occipital expansion that contacted broadly the lateral and posterior margins of the postparietal as it extended to within a short distance of the skull midline. Further, whereas the supratemporal was correctly shown as forming almost the entire posterolateral, horn-like extension of the skull table, the tabular was incorrectly shown as a very small element occupying a position me- dial to the distal end of the skull table extension and as incorporated into the occipital plate. This sutural pattern, considered an autapomorphy of Diadectes (Berman et al., 1992), falsely depicted the supratemporal as not only separating widely the parietal and tabular, but also the postparietal and tabular. This error was revealed not only by examination of the D. absitus skulls, but the skull of the small, juvenile, holotypic skeleton of D. sanmiquelensis (MCZ 2989) de- scribed by Lewis and Vaughn (1965) from the Early Permian Cutler Formation of Colorado (not available until this study). The temporal region in the North American Diadectes is reinterpreted here as identical to that in the German spec- imens. This revision, however, does not alter the analysis of the interrelationships of the diadectomorphs presented by Berman et al. (1992). There are also several features of the postaxial presacral vertebrae of D. absitus that may appear to differentiate it from North American species. These differences are most likely the result of proportional growth changes, as almost all the illus- trations and descriptions of North American species are based on much larger and, therefore, undoubtedly more fully mature specimens than those known from Germany. The most noticeable among these differences include: 1) in D. absitus the neural spines are much lower; 2) the crests of the posterior zygapophyseal buttresses occupy a nearly horizontal plane in D. absitus, whereas in North Amer- ican species the buttresses slope ventrally at about a 25° angle from the horizontal; 90 Annals of Carnegie Museum VOL. 67 and 3) hypantram and hyposphene accessory articulations are, as far as can be determined, absent in D. absitus. That the above vertebral features of D. absitus may reflect an early adult stage of growth is suggested by their occurrence in a comparably sized juvenile or subadult specimen of Diadectes (CM 38036) from the Lower Permian (Wolfcampian) Cutler Formation of north-central New Mex- ico. Using the greatest transverse width of the presacral vertebrae (distance be- tween ends of the posterior zygapophyses) as a means of comparison, the 5 -cm measurement of the North American specimen is approximately only one centi- meter narrower than those in the German specimens MNG 8853 and 7721, The North American specimen (CM 38036), however, possesses one prominent feature of the presacral column not seen in the German specimens, the alternation in height and structure of the neural spines (Sumida, 1990). This feature is common among late Paleozoic tetrapods, but in at least one form, the large captorhinid Labidosaurus, both morphotypes occur (Sumida, 1987, 1990). It was suggested (Sumida, 1987) that the variability of this feature in this genus might represent either a sexual dimorphic or a specific difference. New Autapomorphies of Diadectes In the course of describing the Bromacker Diadectes two additional cranial autapomorphies of this genus have been identified in addition to those recognized by Berman et al. (1992) that appear to set it apart from other diadectomorphs: 1) the palatal ramus of the pterygoid not only makes a substantial contribution to the posterior medial border of the internal naris, but also prevents a palatine- vomer contact; and 2) the supratemporal has a broad contact with the dorsal margin of the paroccipital process of the opisthotic of the braincase. Inadequate materials prevent determining the presence or absence of the above autapomor- phies in the very closely related, nearly identical Late Pennsylvanian Desmatodon. These autapomorphies further strengthen the conclusion of Berman et al. (1992) that the highly derived nature of the genus Diadectes makes it the least desirable member of the diadectomorphs as a potential outgroup in phylogenetic analyses of the interrelationships of amniotes. Acknowledgments Research for this project was supported by National Geographic Society grant 5182-94 (to SSS and DSB); NATO grant CRG. 940779 and California State University San Bernardino Minigrant (to SSS); Edward O’Neil Endowment Fund and M. Graham Netting Research Fund, Carnegie Museum of Nat- ural History (to DSB); and Paleontology grants 1991-1993, Museum der Natur, Gotha. We are greatly indebted to Ms. Amy Henrici, Carnegie Museum of Natural History, Pittsburgh, Pennsylvania, and to Mr. Torts ten Krause, Museum der Natur Gotha, Gotha, Germany, for the difficult task of preparing the specimens discussed herein. We also would like to acknowledge Dr. Elizabeth Regan for her assistance in the field. Thanks are also due the Field Museum of Natural History, Chicago, Illinois; Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts; and Museum of Paleontology, University of California, Berkeley, California; for the loan of specimens. Literature Cited Berman, D. S., and T. Martens, 1993. First occurrence of Seymouria (Amphibia: Batrachosauria) in the Lower Permian Rotliegend of central Germany. Annals of Carnegie Museum, 62:63-79, Berman, D. S., and S. S. Sumida. 1990, A new species of Limnoscelis (Amphibia, Diadectomorpha) from the Late Pennsylvanian Sangre de Cristo Formation of central Colorado. Annals of Carnegie Museum, 59:303-341. — . 1995. New cranial material of the rare diadectid Desmatodon hesperis (Diadectomorpha) from the Late Pennsylvanian of central Colorado. Annals of Carnegie Museum, 64 :'3 15-336. 1998 Berman et al. — Early Permian Diadectes from Germany 91 Berman, D. S., S. S. Sumida, and R. E. Lombard. 1992. 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PaleoBios, 17:1-12. 1998 Berman et al, — Early Permian Diadectes from Germany 93 Vaughn, P. P. 1969. Upper Pennsylvanian vertebrates from the Sangre de Cristo Formation of central Colorado. Contributions in Science, Los Angeles County Museum of Natural History, 164:1-28. . 1972. More vertebrates, including a new microsaur, from the Upper Pennsylvanian of central Colorado. Contributions in Science, Los Angeles County Museum of Natural History, 223:1-30. Watson, D. M. S, 1954. On Bolosaurus and the origin and classification of reptiles. Museum of Comparative Zoology (Harvard University) Bulletin, 111:297-449. Welles, S. P. 1941. The mandible of a diadectid cotylosaur. University of California Publications in Geological Sciences, 25:423-432. WiBLE, J. R., D. Miao, and J. a. Hopson. 1990. The septomaxilla of fossil and Recent synapsids and the problem of the septomaxilla of monotremes and armadillos. Zoological Journal of the Linnean Society, 98:203-228. 3 *< rUi r'-v • ■ A. -;|(--’' •’•'i* '.X ' i*. 't 'll fcfllAcV''!* t-,,. *^..,.,u.V*);V ' ■ 'J*^ ' * -.-h--.- ■'.■ ' I -* ^ •ifl,.. ■:.*- ‘ w*‘ I.;> '■ • '’kfi f ' --- ’ ' V..*’ ' -••-. ' W .mv' ^:%v^-;.-- • ...-K'M^ ■ SlW* •' ^''' '<41 '• ’•'.■•■'♦ ■, , ,11 b t •“<■ iSlfev*i' ' •' '* ■ - -If ■• :S '^1 NUlli^- •’5Jf S' . < ■ ,a^ vV; . ■ .v,> ; ■— ' ^Ji - I : 'VI INSTRUCTIONS FOR AUTHORS ANNALS OF CARNEGIE MUSEUM consist of contributions to the earth sciences (including paleontol- ogy), life sciences, and anthropology, in 30 by 46 picas format (127 by 195 mm or 5 by 7% inches). Submit all manuscripts to the Office of Scientific Publications. Au- thors should give particular attention to scientific con- tent, format, and general style for the ANNALS. Manu- scripts that do not conform to the style of the AN- NALS will be returned to the author immediately. Every manuscript will be subjected to external peer re- view. 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Poletaev 105 Taxonomic status of the Early Permian Helodectes paridens Cope (Diadectidae) with discussion of occlusion of diadectid marginal dentitions David S Berman, Amy C. Henrici, and Stuart S. Sumida 181 Editors, ANNALS, BULLETIN and SPECIAL PUBLICATIONS; J. L. Carter, Ph.D B. C. Livezey, Ph.D D. R. Watters, Ph.D M. A. Schmidt, ELS, Assistant Editor Manuscripts, subscriptions, orders for individual numbers, and changes of address should be sent to: Office of Scientific Publications Carnegie Museum of Natural History 4400 Forbes Avenue Pittsburgh, PA 15213-4080 Phone (412) 622-3287 Fax (412) 622-8837 E-mail: scipubs@clpgh.org ANNALS OF CARNEGIE MUSEUM is published quarterly by The Carnegie Museum of Natural History, 4400 Forbes Avenue, Pittsburgh, Pennsylvania 15213-4080, by the authority of the Board of Trustees of Carnegie Institute. © 1998 Carnegie Institute. @ This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper). ANNALS OF CARNEGIE MUSEUM VoL. 67, Number 2, Pp. 95-104 12 May 1998 NEW SPECIES OF CERNOTINA ROSS (INSECTA: TRICHOPTERA: POLYCENTROPODIDAE) FROM THE AMAZON BASIN IN NORTHEASTERN PERU AND NORTHERN BRAZIL Jan L. Sykora^ Research Associate, Section of Invertebrate Zoology Abstract Four new species of Cernotina — C. harrisi, C. nigridentata, C. aestheticella, and C ecotura — are described from light-trap samples collected in the upper Amazon region in Peru and Brazil. Cernotina bibranchiata Flint and C cygnea Flint are reported as new records for Peru. In addition, new figures of male genitalia of C spinigera Flint are provided. Key words: Cernotina, Trichoptera, microcaddisflies, Amazon, systematics Introduction This contribution is based on two collections in the upper Amazon River basin in Peru and Brazil by Dr. L. J. Davenport from Sanford University, Birmingham, Alabama, and by Dr. Jan Wilt and George Henrych from Boa Vista, Brazil, and Pittsburgh, Pennsylvania, respectively. The microcaddisflies from the Peruvian collection were studied previously by Harris and Davenport (1992). The samples available to us were collected in the vicinity of the Explorama Lodge located approximately 50 km NE of Iquitos on the Rio Yanomono, just upstream from its confluence with the Amazon or Maranon (3°23'S, 72°52'W). The second lo- cality in Peru was the Explomapo Camp located about 70 km NNE of Iquitos (3°10'S, 72°54'W) on the Rio Socusari, The area could be described as tropical floodplain forest with water levels fluctuating up to 10 m per year (Harris and Davenport, 1992). Additional light- trap samples were collected in Brazil by Dr. Jan Wilt and Mr. George Hendry ch in Estado Roraima in the vicinity of Ecotur Park on Rfo Aqua Boa do Univini and in Boa Vista at Rio Branco, This publication deals with the genus Cernotina originally described from North America but recorded also from Central America, West Indies, and South America including Argentina (Flint, 1972, 1974, 1983). A high diversity of close- ly related Cernotina species has been reported from the Amazon Basin indicating that this region is the center of evolution of this genus (Flint, 1971). Therefore, it is not surprising that Cernotina was the dominant macrocaddisfly genus in the samples from Peru and Brazil. The collections contained a total of eight species including four previously described by Flint (1971). The type material of the four new species described in this paper is deposited in the Carnegie Museum of Natural History, Pittsburgh, Pennsylvania. ' University of Pittsburgh Graduate School of Public Health, Department of Environmental and Oc- cupational Health, 260 Kappa Drive, Pittsburgh, Pennsylvania 15238. Submitted 11 March 1996. 95 96 Annals of Carnegie Museum VOL. 67 Systematic Entomology Cernotina karris i, new species (Fig. 1-3) Diagnosis.-— This species is characterized by a combination of features found in male genitalia. These include wide aeterodorsal aspect of segment X tapering to a split posterior section with obliquely truncate apices; short, stout dorsolateral lobe of preanal appendages with robust black, pointed apex directed ventromesad, and a second similarly shaped process located in the center of mesal margin and extended anteromesad. Description. — Male: length 3.9 mm. Legs and antennal segments yellowish, rest of the body yellow brown. Wings venation and general structure typical for genus. Ninth abdominal segment produced anterolaterally, lateral view triangular, dorsal aspect broad, lateral margins with slight indentations, posteroventral margins deeply incised with rounded posterolateral sections. Segment X partially in- serted into segments IX and VIII, in dorsal view triangular with lightly sclerotized, anterodorsal section deeply incised forming blade-iike anterolateral processes, broad and bulbous midsection tapering pos- teriorly to a long apical lobe divided by a mesal split into obliquely truncate apices bearing several spines on mesal margins; lateral view semicircular with slightly undulate ventral margins. Dorsolateral lobe of preanal appendages stout, roughly triangular, shorter than segment X, with broad midsection tapering into pointed, black apex angled sharply ventromesad, center of mesal margin produced into distinct, large, black, pointed process extended anteromesad; lateral view of ventromesal lobe triangular and produced posteriorly into rounded apex with a row of spines on posterior margin. Inferior ap- pendages elongate, rectangular, with rounded apex slightly curved ventrad, anterodorsal lobe long, in lateral view sinuous and sickle-shaped, ventral view of apical section curved laterad with a row of setae mesatly; posteromesal lobe short and round with dark, apically pointed process directed mesad. Phallus slender with internal structure obscure but with a pair of dark internal spines. Remarks .—This species seems to be closely related to C nigridentata. Both species have the short dorsolateral lobe of the preanal appendages produced into a dark apical tooth and a ventromesal, black, pointed projection centered on the ventral margin. Both species can be separated easily by the shape and size of the posteromesal lobe of the inferior appendages which in C. nigridentata is well developed and longer than the main body of the inferior appendages. Based on the available material, this species is the most common Cernotina in the vicinity of Explorama Lodge and Explomapo Camp. Type Specimens.— Tiololypc, male, Peru: Departemeeto Loreto, bank of Rio Yanomono just below Explorama Lodge, 10 January 1993, L. J. Davenport. Para- types, same data as holotype, 12 males; Peru: Loreto, Edge of quiet backwater adjoining Explomapo Camp, 15 January 1993, L. J. Davenport, 27 males. Etymology. — Named for Dr. Steve Harris who provided the specimens for the species described in this paper. Cernotina nigridentata, new species (Fig. 4-6) DiagHOj-w.— This species is characterized by the dorsal lobe of preanal ap- pendages with prominent, black, apical and ventromesal projections; broad, un- divided segment X, and by a long blade-like apicomesal process of the inferior appendages extending beyond apex of the main lobe. Description. — Male: length 3.5 mm. Legs and antennae yellow; thorax, abdomen, and wings brown- ish. Wing venation and general structure typical for genus. Ninth segment in lateral view triangular with rounded anterior margin, central section of posterior margin produced in a rounded lobe providing support for the inferior appendages. Dorsal aspect of segment X bolbose with broad and inflated anterior part tapering to short and truncate apex composed of ventral, small, bilobed and membranous 1998 Sykora— New Amazonian Cernotina 97 Fig. 1-3. — Cernotina harrisi, new species, male genitalia; 1, lateral view; 2, dorsal view; 3, ventral view. 98 Annals of Carnegie Museum VOL. 67 Fig. 4-6. — Cernotina nigridentata, new species, male genitalia; 4, lateral view; 5, dorsal view; 6, ventral view. lobe flanked laterally by lightly sclerotized posterolateral extension of dorsum; lateral aspect halfmoon- shaped with blunt posterior margin bearing long setae. Preanal appendages with dorsolateral lobe elongated, gradually tapering to a single finger-shaped, blackened posterodorsal apex, center of ven- tromesal margin expanded into broad, black, pointed process directed ventromesad; ventromesal lobe produced posteriorly with several setae on truncate apex. Inferior appendages long, in lateral view with rounded apex curved slightly ventrad, ventral aspect of the main body with truncate and broad apex; anterodorsal lobe almost as long as main body, slightly sinuous with a row of large setae mesally; posteromesal lobe bladelike, elongated, well separated from the main lobe, longer than main body of 1998 Sykora — "New Amazonian Cernotina 99 inferior appendages and gradually tapering to sharp apex curved laterad. Phallus tubular with two internal spines and several membranous folds at the apex. This species is closely related to C harrisL It differs most notice- ably in the shape of the inferior appendages with a distinct posteromesal lobe which is well defined and longer than the main body, and segment X which is broad and not divided into two distinct lobes. Type Holotype, male, Peru: Departmento Loreto, banks of Yan- omono Creek just below Explorama Lodge, 10 January 1993, J. L. Davenport. Paratype, same data as holotype, one male. Etymology. — Latin for black-toothed, referring to the dark processes on the preanal appendages. Cernotina aesthetic ella, new species (Fig. 7^9) This species may be recognized by a combination of characters including the divided ventromesal lobe and bifid dorsolateral lobe of the preanal appendages, fused anterodorsal aspect of segment X, and short, free apical section of the anterodorsal lobe of inferior appendages. Description.— yidilQ: length 3.5 mm. Legs and antennae yellowish, rest of body yellowish brown, wings pale brown. Wing venation and general structure typical for genus. Lateral view of ninth ab- dominal segment triangular with rounded anterior and posteroventral section, ventral aspect broad, anterior margin with round, deep incision, lateral margins indented in middle. Dorsal aspect of segment X membranous, partially divided with narrow groove along midline and shallow, triangular incision in apical margin; lateral aspect sinuate, curved ventrad with anteroventral margin covered by leaf-like, pointed scales and with elongated apices slightly directed dorsad. Preanal appendages fused ventro- laterally with segment X, dorsolateral lobe bifid with apex produced into beak-shaped structure with two long, black spines, ventromesal lobe divided into two sections; main ventromesal body with elongated, slender posterior section and narrow anterolateral process covered by small dents with shallow incision at apex. Inferior appendages with anterodorsal lobe partially fused with and shorter than main body of inferior appendages and produced into club-shaped, free apical section; postero- mesal lobe not separated from main body and forming darkened, hook-shaped, short process. Phallus membranous, indistinct internally with two dark spines. Remarks species is related to C. declinata Flint from which it differs by the shape and structure of the preanal appendages, the rounded apex of the inferior appendages, and segment X with a triangular incision in the apical margin. The most unusual characteristic which clearly separates this species from any known Cernotina is the additional lateral process from the ventromesal lobe of the preanal appendages. Additional material of this species is needed to determine the morphology and taxonomical significance of this character. Type Specimen.— Tlololy pc, male, Peru: Departemento Loreto, bank of Yano- mono Creek just below Explorama Lodge, 10 January 1993, L. J. Davenport, Etymology. — Named for its aesthetically pleasing morphological structures. Cernotina ecotura, new species (Fig. 10-14) This species is characterized by specific morphological features found in male genitalia that include the dorsolateral lobe of the preanal append- ages with broad base tapering to a long, slender apical section with pointed apex directed dorsomesad, cup- shaped main body of the inferior appendages with fused and complex posteromesal lobe and miniature anterodorsal lobe. 100 Annals of Carnegie Museum VOL. 67 Fig. 7-9. — Cernotina aestheticella, new species, male genitalia; 7, lateral view; 8, dorsal view; 9, ventral view. 1998 Sykora- — New Amazonian Cernotina 101 Fig. 10“14.- — Cernotina ecotura, new species, male genitalia; 10, lateral view; 11, dorsal view; 12, ventral view; 13, phallus, dorsal view; 14, posteromesal lobe of inferior appendages, ventral view. 102 Annals of Carnegie Museum VOL. 67 Description. — Male: length 4.0 mm. Body, legs, and antennae yellowish brown in alcohol, wings colorless and transparent. Wing venation and general structure typical for genus. Ninth segment in lateral view almost triangular with rounded posterior and anteroventral margins; ventral aspect broad with exoskeleton forming “sunglass frame” structure. Segment X lightly sclerotized, deeply incised middorsally forming broad lateral lobes with pointed apices sparsely covered with long setae and directed posteromesad, in lateral view roughly triangular with rounded anterodorsal portion and straight ventral margin. Preanal appendages bilobed; dorsolateral lobe with rounded anterodorsal margin and broad base tapering to long, narrow apical portion slightly bent dorsad with pointed apex directed dorsomesad; ventromesal lobe short, broad, and truncate with eight setae on posterior margin and two tubercles on posterolateral corners. Inferior appendages with rudimentary, short, and verrucous antero- dorsal lobe with several small setae on dorsal and posterior margins; posteromesal lobe fused with main lobe forming complex dark structure with posteromesal corners bearing two black, hook-shaped teeth and two lightly colored spines; main body of inferior appendages in lateral view almost qua- drangular with rounded apex, ventral aspect cup-shaped with verrucous mesal margin. Phallus extended into relatively short, tubular apical section and broad posterior portion; pair of large straight spines posteriorly and two slightly curved, shorter spines apically. Remarks. — This species is unique and does not seem to be related to any of the known species of Cernotina. The structure of the preanal and inferior ap- pendages is unlike any other species of Cernotina. Type Specimen. — Holotype, male, Brazil: Estado Roraima, Boa Vista, Rio Branco, December 14, 1995, Dr. Jan Wilt and George Hendrych. Paratype, same data as holotype, one male. Entomology. — Named for Ecotur Park, the base camp of the expedition, Cernotina bibranchiata Flint Cernotina bibranchiata Flint, 1971:37. Record. — This species was reported from the Amazon basin in Brazil (Flint, 1971). This is a new record for Peru. Material. — Peru: Departemento Loreto, blackwater creek outlet of Lake Shi- migay 2 km upstream of Napo River from mouth of Sucusari River, 15 January 1993, L. J. Davenport, one male. Cernotina cygnea Flint Cernotina cygnea Flint, 1971:37. Record. — Flint (1971) described this species from the Amazon River basin in Brazil. It is a new record for Peru. Material. — Peru: Departemento Loreto, edge of quiet backwater adjoining Ex- plomapo Camp, 15 January 1993, L. J. Davenport, one male. Cernotina decembens Flint Cernotina decembens Flint, 1971:37. Cernotina decumbens Flint, 1971:62. Report. — Flint (1971) reported this species from the Amazon River basin in Brazil. Material. — Brazil: Estado Roraima, Rio Aqua Boa, December 15, 1995, Dr. Jan Wilt and George Hendrych, one male. Cernotina spinigera Flint (Fig. 15-19) Cernotina spinigera Flint, 1971:38. Record. — This species is widely distributed in the Amazon River basin. In order 1998 Fig. 15-19. — Cernotina spinigera Flint, male genitalia; 15, lateral view; 16, tenth segment, dorsal view; 17, ventral view; 18, anterodorsal lobe of inferior appendages, dorsal view; 19, phallus, dorsal view. 104 Annals of Carnegie Museum VOL. 67 to facilitate identification of this species, new figures of male genitalia are in- cluded in this paper. Material. — Brazil: Estado Roraima, Boa Vista, December 15, 1995, George Hendry ch and Dr. Jan Wilt, one male. Acknowledgments The help of Dr. Oliver S. Flint, who compared the specimen of C. spinigera Flint with the paratype, is gratefully appreciated. Literature Cited Flint, O. S. Jr. 1971. Studies of Neotropical caddisflies, XII: Rhyacophilidae, Glossosomatidae, Philopotamidae, and Psychomyiidae from the Amazon basin (Trichoptera). Amazoniana, 3:1-67. . 1972. Studies of Neotropical caddisflies, XIV: On a collection from northern Argentina. Proceedings of the Biological Society of Washington, 85:223-248. . 1974. Studies of Neotropical caddisflies, XV: The Trichoptera of Suriname. Studies on the Fauna of Suriname and the Other Guianas, 169:1-30. . 1983. Studies of Neotropical caddisflies, XXXIII: New species from austral South America (Trichoptera). Smithsonian Contributions to Zoology, 377:1-100. Harris, S. C, and L. J. Davenport. 1992. New species of microcaddisflies from the Amazon region, with special reference to northeastern Peru (Trichoptera: Hydroptilidae). Proceedings of the En- tomological Society of Washington, 94:454-470. ANNALS OF CARNEGIE MUSEUM VoL. 67, Number 2, Pp. 105-180 12 May 1998 ATOKAN (LATE BASHKIRIAN OR EARLY MOSCOVIAN) BRACHIOPODS FROM THE HARE FIORD FORMATION OF ELLESMERE ISLAND, CANADIAN ARCTIC ARCHIPELAGO John L. Carter Curator, Section of Invertebrate Paleontology Vladislav L Poletaev' Visiting Research Scientist, Section of Invertebrate Paleontology Abstract The Atokan (late Bashkirian or early Moscovian) brachiopod fauna from the lowermost Hare Fiord Formation in northern Ellesmere Island consists of at least 62 species, of which 1 8 are new, in at least 53 genera, of which three are new. New genera are the productoid Lazarevia, type species L. stepa- nowensis n. sp., the stenoscismatoid Careoseptum, type species C. septentrionalis n. sp., and an ex= ternally similar rhynchonelloid, Exlaminella, type species E. insolita n. sp. Other new species include Orthotichia dorsistrigis, Plicatiferina kalashnikovi, Rugivestis pristina, Maemia gelida, lEimbrinia borealis, Liraria paucispina, lElassonia sverdrupensis, Antronaria annosa, Camarium nuperum, Nu- cleospira aquilonaris, Tiramnia walteri, Tiramnia grunti, IHeteraria canadiensis, Elinoria ellesmer- ensis, Parachoristites tellevakensis, and Cranaena nassichuki. This diverse fauna bears clear northern European Russian affinities, as do similar-age faunas from the northern Yukon Territory and south- eastern Alaska. This suggests that there was an open seaway which allowed free faunal dispersion and communication between northern European Russia to the east and also with the northern Yukon Territory and the southeastern Alaska Alexander Terrane to the southwest. Key words: brachiopods, Carboniferous, Arctic, Ellesmere Island, Hare Fiord Formation Introduction The brachiopod fauna that is the subject of this paper was recovered from fenestellid bryozoan reefs at the base of the Hare Fiord Formation on both the north and south sides of Hare Fiord, northern Ellesmere Island. Although Carter (in Nassichuk, 1975) identified Carboniferous brachiopods from the Otto Fiord Formation from directly beneath the Hare Fiord Formation, the present paper constitutes the first description of Carboniferous brachiopods from Ellesmere Is- land and the first description of Upper Carboniferous brachiopods from the Ca- nadian Arctic Archipelago. The Hare Fiord Formation was named by Thorsteinsson (1970) for a succession of limestone, siltstone, and shale up to 732 m (2400 ft) thick within the Sverdrup Basin. This basin was formed by rifting of the Precambrian and Lower Paleozoic rocks of the Franklinian Geosyncline and contains approximately 14 km of upper Paleozoic through Cenozoic siliciclastic, carbonate, and evaporitic sediment. The Sverdrup Basin sequence rests unconformably on lower Paleozoic basement rocks of the Franklinian Geosyncline. The Hare Fiord Formation was deposited near ' Chernobyl Science and Technical Center for International Research (CHECIR), Shkolnaya 6, Cher- nobyl 255620, Ukraine. Submitted 26 November 1996. 105 106 Annals of Carnegie Museum VOL. 67 the center of the basin along its northeast-southwest axis that passes through northern Ellesmere Island. It is overlain by black shale, siltstone, and bedded chert of the Permian Van Hauen Formation and underlain by medium-bedded anhydrite, limestone, and shale of the Bashkirian Otto Fiord Formation. In much of its outcrop region the Hare Fiord Formation consists of two dis= tinctive parts, a lower biogenic limestone that ranges in thickness from a few tens of meters up to 345 m and an upper unit of limestone, siltstone, and shale. The lower reefoid part was informally designated the “Tellevak Limestone” in the Blue Mountains area south of Hare Fiord by Bonham-Carter (1966). Although the brachiopod fauna within this lower biohermal unit is rich and diverse, bra- chiopods are rare above it and appear to be of limited biostratigraphic value. Carboniferous brachiopods in the vicinity of or north of the Arctic Circle in North America are poorly known. Waterhouse, in Bamber and Waterhouse (1971), listed and illustrated Carboniferous and Permian brachiopod faunas from the northern Yukon Territory. Gorveatt and Nelson (1975) later described several Carboniferous and Permian spiriferids from this same region. Waterhouse and Waddington (1982) described both Carboniferous and Permian spiriferellid bra- chiopods from the Yukon Territory and the Canadian Arctic Archipelago. Carter (1975fl) described two small faunules believed to be of late Early Carboniferous age from Melville and Axel Heiberg islands. Carter (1975Z?) listed but did not illustrate brachiopods from the Otto Fiord Formation of Melville and Ellesmere islands. Bowsher and Dutro (1957) and Dutro (1987) listed Carboniferous bra- chiopods from the Lisburne Group of northern Alaska but did not illustrate them. Because there are many genera in the lower Hare Fiord Formation that range into the Permian and because one of the described Permian faunas was originally ascribed to the Carboniferous, citations of published reports dealing with Permian brachiopod faunas follow. These include Holtedahl (1917), Axel Heiberg Island; Chernyshev and Stepanov (1916), Ellesmere Island (originally described as Car- boniferous); Harker and Thorsteinsson (1960), Devon Island; Stehli and Grant (1971), Axel Heiberg Island; and Waterhouse and Waddington (1982), various Arctic islands. All of these described Permian faunas are relatively small with moderate diversity. Faunal Composition Late Paleozoic carbonate buildups commonly bear diverse invertebrate faunas, especially rich in lophophorates, and this is true for the “Tellevak Limestone” in the Blue Mountains and other smaller bioherms elsewhere in the lower part of the Hare Fiord Formation. Several of the collections studied here were made from the flanks of bryozoan mounds or “reefs” as described in Davies et al. (1989) and Nassichuk and Davies (1992). From essentially three rich collections we have discriminated over 60 species of articulate brachiopods, many of which are rep- resented by very few specimens making certain identification difficult or impos- sible. The bulk of the species occur in a single collection from which Nassichuk (1975) described 15 species of Atokan ammonoids (GSC locality 56430). The level of brachiopod diversity in this collection approaches that in many of the individual collections seen in Cooper and Grant’s (1972-1976) monumental monograph on the rich pre-Tethyan Permian silicified faunas of west Texas, which was based on massive samples. Sampling efficiency is usually vastly greater for large silicified samples than for most “crackout” collections. 1998 Carter and Poletaev — Upper Carboniferous Arctic Brachiopods 107 Table 1 . — Summary table showing the taxonomic composition of the brachiopod fauna of the lower Hare Fiord Formation. Group Families Genera Species Orthids 2 2 3 Orthotetids 1 1 1 Chonetids 2 2 2 Productids 8 20 21 Rhynchonellids 5 7 7 Pentamerids 2 2 2 Athyridids 2 2 2 Spiriferids 8 15 19 Retziids 1 1 1 Terebratulids 2 2 4 Because the genus is the basic taxonomic unit used in comparing brachiopods from widely separated basins or continents, only generic diversity of the Hare Fiord brachiopods is discussed below. Productids, rhynchonellids, and spiriferids make up the bulk of this fauna, as can be seen in Table 1, with several notable omissions or poor representations. For example, we found no inarticulates, few chonetids, only scraps of orthotetids, and but two unusual athyridid species. For some of these genera the level of confidence in the generic-level identifi- cation is low because of the lack of interiors. This should be borne in mind when assessing Tables 1 and 2. Even if several of the identifications are incorrect, there is clearly an impressive number of both older and younger anachronisms in this biohermal fauna. Orthids.~Two orthid genera, Rhipidomella and Orthotichia, are found in the lower part of the Hare Fiord Formation. The former, a very long-ranging genus, is not recognizable to the species level and cannot be compared with other North American species. The genus Orthotichia is represented in the lower Hare Fiord Formation by two species, including a new one, O. dorsistrigis. This genus is confined to Upper Carboniferous and Permian strata, first appearing in the late Bashkirian or early Moscovian. Only three species are known from the North American Upper Carboniferous: O. schuchertensis Girty, from the Belden Shale of Colorado and probably of Morrowan (Bashkirian) age according to Langen- Table 2. — Ranges of lower Hare Fiord Formation anachronistic brachiopod genera. Possible misi- dentifications are marked with a (?). Genus Prior range New range Camarium Silurian-Tournaisian Silurian-Lower Moscovian Retimarginifera (?) Sakmarian-Kungurian Lower Moscovian-Kungurian Maemia Late Moscovian Lower Bashkirian-Upper Moscovian Eomarginifera Visean Visean-Lower Moscovian Eomarginiferina Visean Visean-Lower Moscovian Fimbrinia (?) Kasimovian Lower Moscovian-Kasimovian Pustula Tournaisian-Serpukhovian Tournaisian-Lower Moscovian Liraria Kungurian Lower Moscovian-Kungurian Fluctuaria Visean-Serpukhovian Visean-Lower Moscovian Cenorhynchia Asselian-Kazanian Lower Moscovian-Kazanian Hemileurus (?) Asselian Lower Moscovian-Asselian Antronaria Sakmarian-Artinskian Lower Moscovian-Artinskian 108 Annals of Carnegie Museum VOL. 67 heim (1991); O. morganiana Derby, from Desmoinesian beds in Nevada as re- ported by Langenheim (1991); and O. nawtawaketensis Watkins, 1974, from the Baird Formation of California and of Bashkirian or Moscovian age. Several Rus- sian species have been described from the Bashkirian through lower Permian and their morphology and distribution are summarized by Lazarev (1976). Orthotetids.— This group, common in the upper Carboniferous (Pennnsylvan- ian) of the midcontinental portion of North America, is represented in the lower Hare Fiord fauna only by unidentifiable fragments. Chonetids.—This is another poorly represented group in the lower Hare Fiord fauna. Only two genera were identified, Chonetinella and Sokolskya. The former ranges from Bashkirian to Kungurian and the latter ranges from Bashkirian to Kasimovian. In most of North America the two most common Atokan and Des- moinesian chonetoid genera are Mesolobus and Neochonetes, both of which are missing from Ellesmere Island collections. Productids.^ — ^The productids are the most diverse group in this fauna with 21 species in 20 genera. They, along with the spiriferids and rhynchonelloids, make up the bulk of the Hare Fiord brachiopod fauna. The most common productids in the North American midcontinent are cosmopolitan genera such as Antiqua- tonia, Linoproductus, Cancrinella, Kozlowskia, and Echinaria, all represented in the Hare Fiord fauna. The suborder Strophalosiidina is represented by only one genus, Plicatiferina. This genus also occurs in the Upper Bashkirian to Lower Moscovian of south- eastern Alaska. In Russia, it ranges from the upper Moscovian into the Lower Permian. Within the superfamily Productoidea the family Productellidae includes in the lower Hare Fiord the genera Rugivestis, Krotovia, Lazarevia, Maemia, and pos- sibly Eomarginiferina, Fimbrinia, Hystriculina, and Retimarginifera. Rugivestis was described from the Permian of Oregon and also is known from the lower Moscovian through Lower Permian of Russia and Lower Permian of China. The Hare Fiord species is also found in the late Bashkirian or early Moscovian Lad- rones Limestone of southeastern Alaska. Krotovia is most commonly reported from Visean through Bashkirian strata but is rarely reported up through the Lower Permian. Lazarevia is a new monotypic genus. Maemia is otherwise known only from the late Moscovian of Cape Chaika, northern Russia. Lacking interiors the identity of the specimens identified as Eomarginiferina is not certain. That genus is previously known only from the Upper Visean of Ireland. Fimbrinia previously ranged from the late Kazimovian into the middle Permian. Hystriculina normally ranges from the Kazimovian through the Sakmarian. Retimarginifera is a Permian genus. The family Productidae includes the genera Reticulatia, Eomarginifera, Kutorginella, and possibly Kozlowskia. Reticulatia is a cosmopolitan genus rang- ing from the Bashkirian through the Lower Permian. Eomarginifera is a Visean European genus. The genus Kutorginella was described from the Moscovian of the Russian platform but it is also known from several species in Russia and North America ranging in age from Sepukhovian through Kungurian. In western Europe Antiquatonia is known only from the Lower Carboniferous but has been reported from the Upper Carboniferous as well as the Lower Carboniferous of Kazakhstan. In North America this genus first appears in the latest Mississippian and ranges upward into the Lower Permian. Kozlowskia is another cosmopolitan genus ranging from the Bashkirian through the Lower Permian. The superfamily Echinoconchoidea is represented by the genera Pustula and 1998 Carter and Poletaev — Upper Carboniferous Arctic Brachiopods 109 Echinaria. Pus tula is used here in the broad sense because no interiors are known for these Arctic specimens. True Pus tula is probably restricted to the Lower Car- boniferous but the name is applied here to pustolose nonrugose specimens with rounded outlines. Their actual affinities are unknown. Echinaria ranges through- out most of the Upper Carboniferous. Winkler Prins (1968) reported this genus from the lower Moscovian of northern Spain. The superfamily Linoproductoidea is represented by three genera. Liraria was previously known only from the Kungurian of Texas. An undescribed species also occurs in the upper Bashkirian or lower Moscovian of southeastern Alaska. Flue- tuaria was previously restricted to the Visean and possibly Serpukhovian of Eur- asia. Kos’ko et al. (1993) reported it from the middle Bashkirian of Wrangel Island, Winkler Prins (1968) reported it from the upper Moscovian of northern Spain, and Kalashnikov (1980) reported it from the late Upper Carboniferous of Novaya Zemlya. Cancrinella ranges throughout most of the Upper Carboniferous and Permian. Reports from the Bashkirian and Moscovian are less common than from, the Kasimovian and above. These small, fragile shells never seem to occur in abundance but have a remarkably wide distribution. Rhynchonellids.- — There are seven genera of rhynchonelloids in this fauna in- eluding Cenorhynchia, lElassonia, IPhrenophoria, IHemileurus, Septacamera, and Antronaria plus the new genus Exlaminella. All of these genera except the latter have Permian type species and only Septacamera occurs in the Carbonif- erous, where it ranges down into the Bashkirian. Rhynchonellids in the Upper Carboniferous of the midcontinent are not generically diverse, mostly belonging to the genera Wellerella, Leiorhynchoidea, and the punctate genus Rhynchopora, none of which occur in this fauna. The undescribed late Bashkirian or early Mos- covian rhynchonelloids from southeastern Alaska are also generically diverse judging from exteriors. -Steeoscismatoids are represented by two genera, Stenoscis- ma and a new genus, Careoseptum. The former is cosmopolitan and long ranging. The latter is unique to the Hare Fiord Formation. Athyridids .—Th&m are two unusual athyridoids in this fauna, Camarium and Nucleospira. Whereas one would expect to find the cosmopolitan genera Com- posita and Cleiothyridina in any late Paleozoic fauna, they are absent here. In- stead, the superfamily is represented by the above-named rare genera. Camarium, heretofore unknown above the Tournaisian (see Campbell and Engel, 1963, and Carter, 1967), greatly extends its stratigraphic range. Nucleospira is rare above the Visean. Only two species of this genus are described from above the Visean, one from the basal Morrowan and one from the Permian of west Texas. Spiriferids 19 species in 14 or 15 genera the spiriferids are nearly as diverse as the productids. There are two species of the ambocoelioid genus Crur- ithyris, five martinioids in three or four genera, seven species of spiriferoids in seven genera, (including four choristitids in four genera), two species of pae- ckelmanelloids of the genus Cantabriella, two species of the brachythyridoid Mer- istorygma, and one reticularioid. The choristitids and martinioids indicate strong affinities with northern Russian faunas. Few of these spiriferid genera have a narrowly restricted stratigraphic distribution. Most of them range throughout the Upper Carboniferous and Lower Permian. However, the paeckelmanelloid genus Cantabriella is previously known only from the Bashkirian and Moscovian of northern. Spain. Crurithyris ranges throughout the entire Carboniferous and most of the Perm- 10 Annals of Carnegie Museum VOL. 67 ian. The martinioid genera Tiramnia, Hetemria, and Jilinmartinia occur in this fauna. Specimens of Tiramnia occur abundantly in most of the collections. This genus was originally described from the Upper Carboniferous and Permian of Taimyr, Novaya Zemlya, Timan, Ural Mountains, and Greenland. Besides this new occurrence in the Canadian Arctic Archipelago, this genus occurs in rocks of the same age in southeastern Alaska. Heteraria was described from the Permian of west Texas. It also occurs in the lower Moscovian of southeastern Alaska. Jilinmartinia is based on a Late Carboniferous species from northern China and also occurs in the Kasimovian of Bashkiria. A single small martinioid specimen of unusual proportions probably represents a new genus and species. The spiriferoids are the most diverse superfamily of this order and include the genus Anthracospirifer of the family Spiriferidae. The family Choristitidae in- cludes Elinoria and Brachythyrina of the subfamily Angiospiriferinae, Parachor- istites and Trautscholdia of the subfamily Choristitinae, and Tangshanella of the subfamily Tangshanellinae. The trigonotretids are represented only by the genus Gypospirifer of the subfamily Neospiriferinae. Anthracospirifer is a widespread North American Carboniferous spiriferid rang- ing from the lower Visean (Keokuk) through the Moscovian (Desmoinesian). This genus is rare in Russia and its presence in the Hare Fiord Formation indicates North American faunal affinities. Elinoria is found in the upper Moscovian, Late Carboniferous, and Lower Permian of European Russia and the Donets Basin. It also is present in the lower Moscovian of southeastern Alaska as well as the Hare Fiord Formation. These North American occurrences extend the range down to the early Moscovian. Brachythyrina is rare in North America being reported from the very late Mississippian of Wyoming and the Bashkirian or Moscovian (Ettrain Eormation) of Yukon Territory. The genus ranges throughout most of the late Paleozoic of Eurasia. Parachoristites is characteristic of the Bashkirian of Russia but ranges up into the Lower Permian. This is the first report of the genus from North America. Trautscholdia ranges throughout the entire Upper Carboniferous of Russia. Again, this is the first report of this genus in North America. Tang- shanella was first described from the Upper Carboniferous of northeastern China and has subsequently been reported from the Tournaisian of Australia and the Upper Carboniferous of northwestern China and Japan. The neospiriferid Gypos- pirifer ranges from the Bashkirian into the Lower Permian and appears to have cosmopolitan distribution. Retziids. — The retzioid genus Hustedia ranges throughout the late Paleozoic and is cosmopolitan. Terebratulids. — Terebratulids generally are of little use in late Paleozoic age determination, the genera usually being long ranging. Beecheria appears in the Tournaisian and ranges into the Lower Permian. Cranaena is rarely reported in the Upper Carboniferous. It has been reported from the Devonian to the Lower Permian, although many of these reports have not been confirmed with interior morphological details. Biostratigraphy The Atokan (late Bashkirian or early Moscovian) age assigned to this brachio- pod fauna is based on direct and indirect ammonoid, conodont, and foraminifer evidence. Nassichuk (1975) described a large ammonoid fauna from a reef on the north side of Hare Fiord (GSC locality 56430) ascribing an Atokan (or early 1998 Carter and Poletaev — Upper Carboniferous Arctic Brachiopods 111 Moscovian) age to it. Cooodont data published by Bender (1980) supported the early Moscovian age. Nassichuk and Davies (1992) suggested that the Atokan included late Bashkirian as well as early Moscovian strata. Newly discovered conodonts from GSC locality 56430 are being studied by T Nemirovskaya and N. Sobolev who also have ascribed a late Bashkirian to early Moscovian age to the Hare Fiord. Diagnostic foraminifers have been recovered from the lower Hare Fiord For- mation by Mamet (in Nassichuk, 1975) who suggested an early Moscovian age. Groves et al. (1994) also correlated the Atokan with the late Bashkirian and early Moscovian and recovered both late Bashkirian and early Moscovian fusulinaceans from the coeval Nansen Formation. The brachiopods, including taxa formerly ascribed to Visean through Lower Permian stratigraphic ranges, are of much less utility in age determination. The presence of numerous genera which also have been reported in early Moscovian beds in the European portion of Russia, however, generally supports this age determination. Anachronisms Table 2 lists the genera which previously had substantially different stratigraph- ic ranges. Note that there are 12 genera of possible anachronisms, six younger and six older. Misidentifications might reduce this number but it is clear that the Hare Fiord brachiopod fauna represents an unusual upper Bashkirian to lower Moscovian biohermal biofacies unknown elsewhere. These anachronisms origi- nally led us to assign a lower Permian age to the fauna on the assumption that the youngest faunal elements gave some indication of the true age of the fauna. The overwhelming evidence from the ammonoids and microfossils of an upper Bashkirian to lower Moscovian age of the lower Hare Fiord Formation forced us to reexamine and revise our preliminary taxonomic and biostratigraphic deter- minations. Correlations and Paleobiogeography Although the lower Hare Fiord Formation fauna described herein cannot be correlated directly with previously described brachiopod faunas of similar age from North America, it bears many similarities with the undescribed allochtho- nous faunas from the Ladrones Limestone and Klawak Formation of southeastern Alaska, Alexander Terrane. These coeval formations bear rich brachiopod faunas containing a number of distinctive Hare Fiord brachiopod genera that do not occur elsewhere in Bashkirian or Moscovian strata in North America, with the possible exception of the Yukon Territory. These include Plicatiferina, Rugivestis, Fluc- tuaria, Liraria, Elinoria, Tiramnia, Gypospirifer, Meristorygma, B r achy thy rina, and Heteraria. These diverse assemblages certainly provide a strong biogeographic link to the Ellesmere Island fauna of approximately the same age and suggest that they were connected by an unrestricted, if indirect, seaway. The exact location of this seaway is speculative but could have passed through the present Yukon Territory or per- haps through northeastern Alaska, both of which are known to bear largely un- described early Moscovian (Atokan) faunas (Bamber and Waterhouse, 1971; Du- tro, 1987). Correlation with the remainder of North America, however, is much more dif- 112 Annals of Carnegie Museum VOL. 67 ficult. Considering the present fauna, only Kozlowskia splendens (Norwood and Pratten), Chonetinella cf. C jejfordsi Stevens, Reticulatia cf. R. americana Dun- bar and Condra, Antiquatonia cf. A. hermosana (Girty), and Anthracospirifer cf. A. occiduus Sadlick suggest a Pennsylvanian age, ranging from Atokan to Vir- gilian. Several Moscovian species were illustrated or listed from the Upper Carbon- iferous of northern Yukon Territory by Bamber and Waterhouse (1971) and Gor- veatt and Nelson (1975). Most of these species were later assigned a late Mos- covian age by Waterhouse and Waddington (1982). Two small faunules designated as the ''Martiniopsis'" Zone and the Buxtonia Zone, however, were later assigned to the early Moscovian by Waterhouse and Waddington (1982). None of the spe- cies listed or illustrated show much similarity to those of the Ellesmere Island fauna but the presence of so-called Choristites and ''Martiniopsis'" in the Yukon collections suggest Russian affinities. In the late Moscovian faunas the presence of Waagenoconcha and abundant Choristites suggest the same affinities. The spi- riferids described by Gorveatt and Nelson (1975) were dated as late Moscovian and also clearly indicate Russian affinities. In summary, the faunas from limestone reefs in the lower part of the Hare Fiord Formation, from the Ettrain Formation of northern Yukon Territory, and from the Ladrones Limestone and Klawak Formation of southeastern Alaska all have strong affinities with those of northern European Russia and with each other. The faunas from the Alexander Terrane of southeastern Alaska may be alloch- thonous but the close similarity between those faunas and those from the lower Hare Fiord clearly suggest an open marine communication between the two regions. Correlations with upper Bashkirian or lower Moscovian strata in northern Eu- ropean Russia is difficult, being dependent mainly on faunal lists. Kalashnikov (1980) reviewed late Paleozoic brachiopod faunas from this region and listed species for numerous localities and horizons in the general area encompassing Novaya Zemlya, Cape Chaika, Timan, and the northern Urals. However, most of his illustrations are of specimens from younger beds. The following genera from the lower Hare Fiord Formation are reported by Kalashnikov from Bashkirian or Lower Moscovian beds in northern Russia: Rhipidomella, Eomarginifera, Rugi- vestis, Kozlowskia, Antiquatonia, Krotovia, Kutorginella, Reticulatia, Fimbrinia, Linoproductus, Fluctuaria, Septacamera, Parachoristites, Purdonella, Brachy- thyrina, Flinoria, Meristorygma, Phricodothyris, Cranaena, and Tiramnia (Ka- lashnikov includes numerous references to various species of Martinia, some of which probably belong in Tiramnia). This assemblage is unique to northern Russia and Ellesmere Island but also includes many cognate genera in southeastern Alaska. GSC Localities Figure 1 shows Geological Survey of Canada (GSC) localities where late Bash- kirian to early Moscovian brachiopods were found in the Hare Fiord Formation. Descriptions of these localities follow. GSC Locality C~4083 .—In the saddle of a northeast-southwest trending ridge about eight km northwest of Mount Barrell and immediately west of Wood Glacier (80°54'N, 84°11'W). The locality is in thin-bedded, dark grey limestone, about 46 m above the base of the Hare Fiord Formation. It is in the nonreef facies of 1998 Carter and Poletaev — Upper Carboniferous Arctic Brachiopods 113 Fig. 1. — Index map showing Geological Survey of Canada (GSC) localities for the Upper Carbonif- erous brachiopods described in this report, from both the north and south sides of Hare Fiord, northern Ellesmere Island. Most of the fossils are from GSC locality 56430 in a small bryozoan bioherm on the north side of Hare Fiord. GSC locality 56430A is from the same stratigraphic level as GSC locality 56430 but is 3 m farther to the west. Similarly, GSC locality C-5202 is 1.5 m stratigraphically below locality 56430. GSC localities C-4083 and C-4084 are in thin argillaceous limestone some 45.72 m (150 ft) above the base of the Hare Fiord Formation west of Wood Glacier; GSC locality C-4084 is 60.96 m (200 ft) along strike to the west of GSC locality C-4083. GSC localities C-4087, C-4085, and 60194 in the southern Blue Mountains occur in the upper part of a bryozoan reef complex that Bonham-Carter (1966) informally named the “Tellevak Limestone” in the lower Hare Fiord Forma- tion. Dashed lines indicate boundary of permanent ice. the formation and was discovered originally by W. W. Nassichuk and C. Spinosa (Boise State University) in 1969. GSC Locality C-4084. — This locality is in the same stratigraphic position 61 m along strike to the west of GSC locality C-4083. This locality is in the nonreef, basinal facies of the Hare Fiord Formation. GSC Locality C-40S5.— This locality occurs in light grey skeletal limestone in the upper three m of the informally designated “Televak Limestone” in the Hare Fiord Formation. It is on the southwestern side of the Blue Mountains, 5.8 km northeast of the northern tip of Hare Fiord Diapir (80°44'N, 85°40'54"W). The locality is in a creek bottom near the terminus of the southernmost large westerly directed glacial tongue flowing from a small ice field in the southern Blue Moun- tains. This locality was discovered by W. W. Nassichuk and C. Spinosa (Boise State University) in 1969. GSC Locality C-4087. — This locality is in the upper part of the “Tellevak 114 Annals of Carnegie Museum VOL. 67 Limestone” in the lower Hare Fiord Formation on the northwest side of the Blue Mountains, 1.3 km east-northeast of GSC locality C-4085. GSC Locality 56430. — Principal brachiopod locality. Near the top of a small bryozoan mound about 30 m above the base of the Hare Fiord Formation. This locality is less than 300 m north of the fiord and 3.2 km west of the midpoint of the delta formed by Stepanow Creek (81°07'30"N, 84°20'W). Collected by W. W. Nassichuk and various field associates in 1963, 1969, and 1971. GSC Locality 56430A. — In the same stratigraphic position as GSC locality 56430 and within the same small bryozoan mound but three m farther to the west. GSC Locality C-5202. — This is the same general location within the same bryo- zoan mound as GSC locality 56430, only 1.5 m stratigraphically lower. GSC Locality 60194. — Upper part of the reefoid “Tellevak Limestone” in the lower Hare Fiord Formation. The locality is 174 m above the top of the underlying Otto Fiord Formation (Bashkirian) which contains mainly anhydrite, shale, and limestone. It is ten km east of eastern shore of Hare Fiord, southern Blue Moun- tains, northern Ellesmere Island (80°44'11"N, 85°45'40"W). Collected by G. F. Bonham-Carter (Geological Survey of Canada) in 1963. Systematic Paleontology In the following section we have used the relatively recent classifications of Brunton et al. (1995) for the productoids and Afanas’eva (1988) for the chone- toids. For the spiriferids we have used the revised classification of Carter et al. (1994). The rhynchonellid classification follows that of Savage (1996). For the other groups we have generally followed the classification in the original Treatise on Invertebrate Paleontology (Williams et al., 1965). All types are deposited in the type collections of the Geological Survey of Canada, Ottawa. The remainder of the specimens are in the collections of the Institute of Sedimentary and Petroleum Geology, Calgary. Phylum Brachiopoda Dumeril, 1806 Class Articulata Huxley, 1869 Order Orthida Schuchert and Cooper, 1932 Suborder Orthidina Schuchert and Cooper, 1932 Superfamily Enteletoidea Waagen, 1884 Family Rhipidomellidae Schuchert, 1913 Genus Rhipidomella Oehlert, 1890 Rhipidomella sp. (Fig. 2.1, 2.2) Discussion.— Thoie are two specimens of this genus in the collection from GSC locality 56430, plus one ventral valve from GSC locality 56430A. The large incomplete specimen is broad and thin bodied, similar in some respects to Rhip- idomella oweni Hall and Clarke from the Lower Visean of Kentucky or Rhipi- Fig. 2. — Orthids. 2.1, 2.2, Rhipidomella sp., GSC 115535; dorsal and ventral views, X 1.5. 23-2.1 , Orthotechia sp., GSC 115536; ventral, dorsal, lateral, anterior, and posterior views, X 1. 2.8-2.37, growth series of Orthotechia dorsistrigis n. sp., including the holotype (Fig. 2.8-2.12, GSC 115537) and five paratypes, GSC 115538-115542, respectively, X 1. 1998 Carter and Poletaev — Upper Carboniferous Arctic Brachiopods 115 116 Annals of Carnegie Museum VOL. 67 Table 3. — Measurements (in millimeters) of the types o/ Orthotichia dorsistrigis n. sp. GSC number Locality Length Width Thickness 115537 56430 19.5 24.1 13.0 115538 56430 17.2 21.2 11.4 115539 56430 16.9 19.0 10.4 115540 56430A 13.6 19.5 8.7 115541 56430 13.7 16.7 8.0 115542 56430 11.4 12.6 7.8 domella pressula Lazarev from the Moscovian of the polar Urals. The other spec- imens are much smaller but of similar proportions. Family Enteletidae Waagen, 1884 Subfamily Schizophoriinae Schuchert, 1929 Genus Orthotichia Hall and Clarke, 1892 Orthotichia dorsistrigis, new species (Fig. 2.8-2.37) Holotype. — GSC 115537, Figures 2.8-2.12, from GSC locality 56430. Paratypes. —GSC 115538, 115539, 115541, and 115542, Figures 2.13-2.22, 2.28-2.37, from GSC locality 56430; GSC 115540, Figures 2.23-2.27, from GSC locality 56430A. Description. — Medium size, unequally biconvex; outline transversely subovate; dorsal valve equal to or slightly longer than ventral valve; maximum width attained slightly anterior to midlength; hinge- line about equal to half of maximum width; anterior margin slightly emarginate and weakly uniplicate; ventral sulcus weakly developed; dorsum with distinct secondary sulcus on fold in anterior half or two-thirds of valve; lateral profile moderately thin, guttate; cardinal extremities of both valves slightly compressed; entire surfaces of valve regularly capillate. Ventral valve slightly thinner than opposite valve, moderately to weakly convex with slightly inflated umbonal region; sulcus broad, shallow, confined to anterior half of valve, with low median fold in rare specimens; beak small, overhanging nearly catacline low, widely triangular interarea; delthyrium about as wide as high; ventral interior with long, high dental plates and low, thick median ridge; other details unknown. Dorsal valve slightly to moderately thicker than ventral valve, most convex in umbonal region, flanks sloping evenly to lateral margins; umbonal region slightly inflated and projecting posterior to hingeline with small incurved beak; dorsum broadly depressed in most mature specimens forming shallow sulcus; dorsal interarea low, orthocline; dorsal interior with moderately thickened cardinal process and long, high, diverging brachiophore plates; other details unknown. Measurements of Types. — See Table 3. Diagnosis. — This species is characterized by its moderate size, transverse out- line, thin profile, shallow broad ventral sulcus, and dorsal sulcus. Comments. — Orthotichia pentagona (Eichwald, 1860), from the Bashkirian of Ukraine and Russia, is much larger than O. dorsistrigis n. sp. with a subpentagonal outline and stronger dorsal sulcus and ventral fold. A dorsal sulcus is also found in the specimen described as Orthotichial sp. by Yanagida (1975), from the upper Moscovian of Thailand. Distribution. — GSC locality 56430 (36 specimens); GSC locality C-5202 (five specimens). Orthotichia sp. (Fig. 2.3-2.7) Comments. — There are two large orthotichias from GSC locality 60194 that are clearly distinct from the new species described above as O. dorsosulcata n. 1998 Carter and Poletaev — Upper Carboniferous Arctic Brachiopods 17 sp. These large specimens lack the dorsal sulcus seen in that species. They appear to be similar to Orthotichia schuchertensis Girty, 1903, from the Hermosa For= mation of Colorado, in having discontinuous and disproportionately deep inter- spaces defining some of the irregularly strong costellae. Distribution.— GSC locality 60941 (two specimens). Order Strophomenida Opik, 1934 Suborder Orthotetidina Waagen, 1884 Unidentifiable orthotetoid Comments. — There are seven fragmentary specimens of orthotetids in the Hare Fiord collections. They seem to belong to an unusually thin, flattened form of unknown generic affinities. The largest fragment, which is approximately 53 mm wide by 36 mm long, is apparently a flattened dorsal valve. There is no indication of plates in either valve of these fragments and preservation is imperfect. Distribution.— All seven specimens are from GSC locality 56430. Suborder Chonetidina Muir- Wood, 1955 Superfamily Chonetoidea Bronn, 1862 Family Rugosochonetidae Muir- Wood, 1962 Subfamily Rugosochonetinae Muir- Wood, 1962 Genus Chonetinella Ramsbottom, 1952 Chonetinella cf. C. jejfordsi Stevens, 1962 (Fig. 3.6) *1962 Chonetinella jejfordsi Stevens, p. 627, pi. 93, fig. 18, 19. Description. — Medium size; moderately thick; ventral umbo moderately inflated and extending well posterior to hingeline; cardinal extremities defined by concave flexures; venter moderately broad with narrow, shallow, rounded sulcus that originates near beak; ornament of about 20 to 21 capillae per five mm near anterior margin; dorsal valve and interiors unknown. Comments. — This single specimen is similar to specimens of Chonetinella jef- fordsi Stevens, 1962, described and illustrated by both Stevens and Sutherland and Harlow (1973:34). Distribution. — GSC locality 56430 (one specimen). Subfamily Chalimochonetinae Afanas’eva, 1988 Genus Sokolskya Aizenverg, 1980 ISokolskya sp. (Fig. 3.5) Description. — A single large chonetid ventral valve is tentatively assigned here to the genus So- kolskya Aizenverg on the basis of its large size, weakly convex profile with flattened umbonal region, fine ribbing, and short thin ventral septum. Distribution.^ — -GSC locality 56430 (one specimen). Suborder Strophalosiidina Waagen, 1883 Superfamily Strophalosioidea Schuchert, 1913 Family Araksalosiidae Lazarev, 1989 Subfamily Quadratiinae Lazarev, 1989 Genus Plicatiferina Kalashnikov, 1980 Plicatiferina kalashnikovi, new species (Fig. 3. 1-3.4) Holotype. — An incomplete ventral valve showing the hingeline spines, GSC 115544, from GSC locality C-5202, Figure 3.2. 118 Annals of Carnegie Museum VOL. 67 1998 Carter and Poletaev — Upper Carboniferous Arctic Brachiopods 19 Table 4. — Measurements (in millimeters) of the types o/ Plicatiferina kalshnikovi n. sp. GSC number Locality Length Width 115543 56430A 15.0+ 18.6 + 115544 C-5202 19.1 + 24.3 + 115545 56430A 14.1 21.3 + 115546 C-5202 16.6 25.7 Paratypes. — A ventral valve with a disarticulated dorsal valve impressed against it, GSC 115543, from GSC locality 56430A, Figure 3.1; a small ventral valve, GSC 115545, from GSC locality 56430A, Figure 3.3; a natural mold of a dorsal valve exterior, GSC 115546, from GSC locality C- 5202, Figure 3.4. Description. — Medium size; outline transversely subquadrate; maximum width at hingeline; very thin profile; both valves flattened, nearly planoconvex with very thin body cavity; both valves entirely covered with slightly irregular concentric rugae, numbering about nine to 12 per cm in largest spec- imens. Ventral valve weakly convex with weakly concave lateral extremities; beak very small, projecting slightly posterior to hingeline; umbonal region very slightly inflated, remainder of valve flattened; interarea flattened or slightly concave, low, apsacline; delthyrium apparently open; five or more pairs of regularly spaced spines at hingeline directed posterolaterally at angle of 30 to 45 degrees to hinge- line; fine low spines on body of shell rare, scattered, on small nodes or occasionally short, elongate ridges; teeth small; other interior details unknown. Dorsal valve nearly flat; dorsal interarea absent or not observed; spines absent; rugae as in opposite valve; interior with low bilobed cardinal process; other internal details not observed. (Note: a dorsal interior from the Ladrones Limestone of southeastern Alaska has an alveolus in front of a small bilobed cardinal process which is flanked by small transverse tooth sockets. The specimen is spalled anteriorly and no muscle scars or other internal details are indicated.) Diagnosis. — -Medium size Plicatiferina with nearly regular concentric rugae and a row of hingeline spines directed posterolaterally at a moderately low angle to the hingeline. Measurements.— -See Table 4. Comparisons. — -This species externally resembles the holotype of Plicatiferina pseudoplicatilis (Stepanov, 1948), from the Carboniferous (Moscovian or Kasi- movian?) of Bashkiria, differing mainly in its more regular rugae. A row of hingeline spines has never been reported in this species. Stepanov’s paratypes have a more rounded outline with irregular bifurcating rugae than this Hare Fiord species. Kalashnikov (1980) and Lazarev (1990:pl. 40, fig. 8--11) illustrated spec- imens of P. neoplicatilis (Stepanov, 1939) from Cape Chaika showing this irreg- ular rugation, as does P. borealica Kalshnikov, 1980. All of these Russian species Fig. 3. — Chonetoids, strophalosioids, and productoids. 3. 1-3.4, Plicatiferina kalashnikovi n. sp.; 3.1, dorsal view showing ventral interarea and teeth, GSC 115543, X 4; 3.2, the holotype, a ventral valve showing spine bases at hingeline, GSC 115544, X 2; 3.3, 3.4, two ventral valves, GSC 115545, 115546, X 1. 3.5, ventral valve of ISokolskya sp., GSC 1 15547, X 1. 3.6, ventral valve of Chonetinella cf. C. jeffordsi Stevens, 1962, GSC 115548, X 1. 3.7-3.13, Hystriculina cf. H. wabashensis (Norwood and Pratten, 1855); 3.7, 3.8, ventral and dorsal views of an articulated specimen, GSC 115549; 3.9, natural mold of dorsal exterior, GSC 115550; 3.10-3.13, ventral, anterior, posterior, and lateral views of ventral valve, GSC 115551, all X 1.5. 3.14-3.29, Lazarevia stepanowensis n. sp.; 3.14-3.19, ventral, dorsal, dorsal mold, anterior, posterior, and lateral views of the holotype, GSC 1 15552; 3.20- 3.23, lateral, ventral, anterior, and posterior views of a large ventral valve paratype, GSC 115553; 3.24-3.27, ventral, anterior, posterior, and lateral views of a nearly complete paratype with a portion of the dorsal mold exposed, GSC 115554; 3.28, 3.29, ventral and posterior views of a natural mold paratype of the ventral interior showing muscle field, GSC 115555; all X 1. 120 Annals of Carnegie Museum VOL. 67 are significantly younger than P. kalashnikovi, ranging from Late Moscovian to Kasimovian in age. Branton et al. (1994) recently redescribed the type species of the chonopectinid genus Semenewia Paeckelmann, 1931. This genus is similar to Plicatiferina in having a flattened, regularly rugose “bauplan” with well-developed interareas in both valves and a row of spines at the hingeline. The surface is finely striate or capillate. Internally, the bilobed cardinal process is supported by a median septum. No radial micro-ornament has been observed in either the Hare Fiord Formation or Ladrones Limestone specimens which are of similar age. If the dorsal interior of the Ladrones specimen described above can be assumed to also typify the Hare Fiord Formation, it is safe to say that these late Bashkirian or early Moscovian specimens do not belong in Semenewia. Distribution. — GSC locality C-5202 (34 specimens); GSC locality 56430A (two specimens). This species, or one very similar to it, occurs in collections of the Ladrones Limestone of southeastern Alaska. This material was collected in 1918 by G. H. Girty of the U. S. Geological Survey. Suborder Productidina Waagen, 1883 Superfamily Productoidea Gray, 1840 Family Productellidae Schuchert, 1929 Subfamily Productininae Muir-Wood and Cooper, 1960 Tribe Paramarginiferini Lazarev, 1986 Genus Eomarginiferina Brunton, 1966 lEomarginiferina sp. (Fig. 6.11-6.15) Comments. — These two specimens differ from the species described below as Eomarginifera in their narrower subtrigonal outline, similar to that seen in the genus Eomarginiferina Brunton. The mold of the dorsal exterior has the charac- teristic concave grooves delineating the ears that indicate the presence of a strong ridge wrapping around the ears of the dorsal valve interior. Unfortunately, one ear of the ventral valve is missing and the other is spalled making it impossible to determine whether or not there were halteroid spines on the ears. For this reason the assignment to Eomarginiferina is queried. Furthermore, this genus is not known above the Visean. Distribution. — GSC locality C-5202 (one ventral valve, one dorsal valve mold). Fig. 4. — Productoids. 4. 1-4.6, IRetimarginifera sp., ventral, dorsal, mold of dorsal exterior, lateral, anterior, and posterior views, GSC 115556, X 1.5. 4.7-4.23, Rugivestis pristina n. sp.; 4.7, mold of dorsal exterior paratype, GSC 115557; 4,8-4.19, lateral, ventral, anterior, and posterior views of three ventral valves, GSC 115558-115560, respectively, including the holotype (GSC 115558) and two paratypes; 4.20-4.23, ventral, dorsal, partial dorsal exterior mold, and lateral views of a nearly com- plete specimen, GSC 115561; all X 1.5. 4.24-4.43, Maemia gelida n. sp.; 4.24-4.28, ventral, dorsal, mold of dorsal exterior, posterior, and lateral views of the large holotype, GSC 115562; 4.29-4.31, ventral, dorsal, and mold of dorsal exterior of a small paratype, GSC 115563; 4.32-4.43, ventral, anterior, posterior, and lateral views of three ventral valve paratypes, GSC 115564-115566, respec- tively; all X 2. 1998 Carter and Poletaev — Upper Carboniferous Arctic Brachiopods 121 122 Annals of Carnegie Museum VOL. 67 Table 5. — Measurements (in millimeters) of the types o/Rugivestis pristina n. sp. from GSC localities 56430 and 56430A. GSC number Length Width Height Surface length 115558 14.9 18.8 8.5 21.8 115559 14.8 18.4 6.7 21.3 115560 12.6 15.0 7.1 19.9 115561 14.0 — 6.9 20.2 Genus Rugivestis Muir- Wood and Cooper, 1960 Rugivestis pristina, new species (Fig. 4.7--4.23) Holotype. — GSC 115558, Figures 4.8-4.11, from GSC locality 56430. Paratypes.— GSC 115557, Figure 4.7, GSC locality 56430A; 115559, Figures 4.12-4.15, GSC locality 56430A; 115560, Figures 4.16-4.19, from GSC locality 56430; 115561, Figures 4.20-4.23, from GSC locality 56430A. Description. — Medium size for genus, transversely trigonal in outline; maximum width at hingeline; both valves strongly geniculate; body cavity thin; lateral profile moderately inflated, subangular. Ventral valve with visceral disc wider than long, evenly and moderately convex, abruptly delimited from trail by strong cincture; ears large, flattened, weakly costellate, lacking rugae, defined by abrupt straight posterolateral vertical cincture of visceral disc; cardinal extremities formed as vertical flanges at 90-degree angle to ventral surface of ears; trail long and commonly anteriorly nasute; trail may be rounded, flattened, or have shallow sulcus posteriorly; visceral disc, excluding ears, reticulate, orna- mented by moderately coarse rugae and moderately strong costellae, about seven to ten in five mm, commonly seven or eight; rugae abruptly terminate at ear ridges; trail with costellae only; spines rare, commonly with one or two pairs symmetrically arranged on visceral disc and one or more others scattered on visceral disc; spines on trail not observed; interior unknown. Dorsal valve moderately concave; ears defined by straight posterolateral ridge as in opposite valve; trail short; anterior portion of visceral disc and trail with low fold; small protegulal node at hingeline; ornament of visceral disc similar to opposite valve; cincture absent on exterior molds; spines on dorsal visceral disc not observed; interior poorly known, with lateral ridges diverging from hingeline and presumably following cincture; other details not observed. Measurements Table 5. Diagnosis. — This species is characterized by moderate size, coarse ribbing, nonrugose ears, and moderately nasute trail. Comments. — Rugivestis pristina, n. sp., is similar in most respects to both Ru- givestis carinata (Muir- Wood and Cooper, 1957), the type species, and R. kutor- gae (Chernyshev, 1902). The former, of Middle Permian age, is smaller and has a ventral sulcus on the ventral visceral disc, a narrowly nasute trail, and weakly rugose ears. The latter, of Asselian or Early Permian age, is considerably larger and most specimens are not nasute. In addition the ears are proportionately smaller and bear good rugae. The genus is quite rare and the dorsal interior is still un- known so far as we know. Distribution. — GSC locality 56430 (four ventral valves); GSC locality 56430A (four ventral valves, one mold of dorsal exterior, two partially bivalved speci- mens); GSC locality C-4087 (one ventral valve). This species, or one very similar to it, also occurs in the Ladrones Limestone of southeastern Alaska. 1998 Carter and Poletaev — Upper Carboniferous Arctic Brachiopods 123 Table 6. — Measurements (in millimeters) o/ ?Fimbrinia borealis n. sp. from GSC locality 56430. GSC number Valve Length Width Height Surface length 115582 ventral 11.1 10.6 5.4 17.0 115583 ventral 10.0 10.9 4.8 14.3 115584 dorsal 8.5 11.2 4.6 — Subfamily Overtoniinae Muir- Wood and Cooper, 1960 Tribe Overtoniini Muir- Wood and Cooper, 1960 Genus Fimbrinia Cooper, 1972 IFimbrinia borealis, new species (Fig. 6.33-6.43) Holotype. — GSC 115582, Figures 6.33-6.36, a ventral valve, from GSC local- ity 56430. Paratypes. — GSC 115583 and 115584, Figures 6.39-6.43, a ventral valve and a natural mold of the dorsal exterior, from same locality as holotype. Description. — Small; strongly concavoconvex, body cavity thin; outline subovate; greatest width usually at hingeline, rarely at midlength; lateral profile subguttate to subovate; anterior profile bell- shaped. Ventral valve strongly inflated posteriorly, most convex in umbonal region; beak small, scarcely overhanging hingeline; venter well rounded in both views, flanks sloping steeply to lateral margins; ears of moderate size, subangular, well defined by concave flexures; trail moderately convex; entire surface strongly rugose and thickly lamellose with strong, moderately broad, irregular rugae; spine bases nearly regularly spaced, arranged in concentric rows along crests of rugae; interior unknown. Dorsal valve strongly concave; most concave posteriorly; ears delimited by convex flexures; very small protegulal node present at midhingeline; trail absent; ornament consisting of strong lamellose rugae only; spines and dimples apparently absent; interior unknown. Measurements. — See Table 6. Diagnosis. — This species is characterized by its small size, rounded outline, strongly convex venter, tumid profile, and coarse rugose ornament on both valves. Comments. — The dorsal valve of this species is poorly known. The two spec- imens at hand are spalled dorsal interiors and do not give any indication of the presence of spines or dimples on the exterior surface as one would expect in the genus Fimbrinia. However, the state of preservation of these specimens is far from perfect and we cannot discount the possibility that the true ornament is not fully expressed in these specimens. This species cannot be assigned to a genus or even family with confidence. The strong rugae on both valves, especially the dorsal valve, are unusual for the genus Fimbrinia or, for that matter, Overtonia. The apparent absence of dorsal spines is even more troubling. Assignment to the genus Fimbrinia is a name choice of convenience, for this species must have a genus name. Its distinctive shape, profile, and ornament are reason enough to propose a new species but we have little idea as to its true congeners. Distribution. — GSC locality 56430 (five ventral valves, two dorsal valves). Tribe Krotoviini Brunton, Lazarev and Grant, 1995 Genus Krotovia Frederiks, 1928 Krotovia cf. K. spinulosa (J. Sowerby, 1814) (Fig. 6.31, 6.32) 1814 Productus spinulosus J. Sowerby, p. 155, pi. 68, fig. 3. Description. — Medium size; outline subcircular to slightly transversely subovate; ventral valve mod- 124 Annals of Carnegie Museum VOL. 67 erately and almost evenly convex, most convex in umbonal region; umbonal region moderately in- flated, narrow, moderately incurved; beak small, slightly overhanging hingeline; ornament of weak, closely spaced rugae, strongest near sides of umbo, and concentric rows of round, small, tubercular spine bases crudely arranged in quincunx; widely spaced growth lamellae rarely preserved; spines apparently erect; other details unknown. Comments. — This Arctic species conforms fairly closely to Brunton’s (1966) redescription of Krotovia spinulosa (J. Sowerby) from the Upper Visean of Ire- land. It differs mainly in having stronger rugation and a slightly narrower umbonal region. Distribution. — GSC locality 56430 (three ventral valves). Krotovia cf. K. lamellosa Brunton, 1966 (Fig. 6.28-6.30) 1966 Krotovia lamellosa Brunton, p. 225, pi. 13, fig. 8-16; pi. 14, fig. 1-19. Description. — Small, outline subcircular; ventral umbonal region strongly inflated, narrow, strongly incurved; beak small, overhanging hingeline; ornament consisting only of lamellose growth lamellae; fine spines of variable size arranged in concentric rows on growth lamellae; other details not observed. Comments. — This solitary specimen agrees in most respects with Brunton’s (1966) description and illustrations of Krotovia lamellosa from the Upper Visean of Ireland. It differs only in having fewer spines on the growth lamellae. Distribution. — GSC locality 56430 (one ventral valve). Subfamily Marginiferinae Shehli, 1954 Tribe Paucispiniferini Muir- Wood and Cooper, 1960 Genus Hystriculina Muir- Wood and Cooper, 1960 IHystriculina cf. H. wabashensis (Norwood and Pratten, 1855) (Fig. 3.7-3.13) 1855 Productus wabashensis Norwood and Pratten, p. 13, pi. 1, fig. 6a-d. Description of Hare Fiord Specimens. — Small, strongly concavoconvex; outline of visceral disc subquadrate; greatest width at hingeline; both valves moderately geniculate; cardinal extremities sub- angular; lateral profile nearly semicircular; anterior profile subquadrate to subtrapezoidal; shell sub- stance thin. Ventral valve strongly inflated, most convex at point of geniculation; visceral disc only moderately convex; flanks dropping steeply to lateral margins; cardinal extremities well defined by concave flex- ures; beak small, projecting posteriorly slightly beyond hingeline; umbonal region not much inflated, subtending an angle of about 90 to 100 degrees; sulcus well developed, originating on anterior half of visceral disc, becoming deeper and extending to anterior margin; ornament on visceral disc con- sisting of moderately strong, nearly regular, low rugae on whole visceral disc and weak costae ante- riorly; flanks and venter with numerous low, rounded, variably strong costellae, about seven to eight per five mm near point of geniculation; erect spine bases sparsely scattered on visceral disc and trail, several finer spines in row along hingeline; interior not observed. Dorsal valve weakly concave in visceral disc; cardinal extremities well set off by convex flexures; sulcus originating in anterior third of visceral disc; ornament consisting of weak rugae and scattered, rounded pits on visceral disc; ribbing on trail complementary to opposite valve; spines apparently absent; interior not observed. Comments. — This genus has heretofore been restricted to much younger hori- zons and the generic identification should be viewed with caution. These Hare Fiord specimens differ little externally from the specimens of H. wabashensis illustrated by Sturgeon and Hoare (1968) from the Ames Limestone (Upper Mis- sourian) of Ohio. The ears of this Arctic species are considerably smaller than on those illustrated by Dunbar and Condra (1932) from the Missourian and Virgilian of Nebraska. 1998 Carter and Poletaev — Upper Carboniferous Arctic Brachiopods 125 Distribution, — GSC locality 56430 (16 specimens); GSC locality 56430A (one ventral valve). Genus Retimarginifera Waterhouse, 1970 IRetimarginifera sp. (Fig. 4. 1-4.6) Description. — Small to medium size; strongly concavoconvex; body cavity thin; outline subquad- rate; maximum width at hingeline; ears medium size, not large; visceral disc strongly geniculated; trail moderately long. Ventral valve strongly inflated; sulcus well developed, originating in umbonal region, becoming uniformly deeper and wider over much of valve; angular ears defined by concave flexures; visceral disc weakly convex, most convex near beak, weakly reticulate with few irregularly spaced rugae intersected by numerous simple or bifurcating fine costae on anterior portion; trail with moderately strong costae, about eight to ten per five mm; spines sparsely distributed, with single strong spine on ears, pair of strong spines on sides of umbo, and few fine spines on umbo and near hingeline; interior unknown. Dorsal valve strongly concave; fold originating in umbonal region; ears well delimited by abrupt convex flexures; ornament complementary to opposite valve but with few rounded pits on visceral disc; spines absent; interior unknown. Comments, — This species is represented in the Hare Fiord by only one nearly complete specimen. Another poorly preserved ventral valve may belong here as well. The lack of a dorsal marginal ridge or multiple trails eliminates it from the Marginiferinae. There is superficial similarity to the Late Carboniferous genus Jiguliconcha Lazarev, 1990, but that genus has weak, fine ribbing, and no haL teroid spines on the ears or umbo, but it does have a row of spines on the lateral slopes unlike the Hare Fiord specimen. The modest size, thin body cavity, weakly reticulate visceral disc, paired halteroid spines at the ears and umbo suggest place- ment in the subfamily Hystriculininae. Assignment within this subfamily is dif- ficult. The Permian genus Retimarginifera Waterhouse is similar in most respects but has a much more strongly reticulate visceral disc. Therefore we assign it here with considerable uncertainty. Distribution.— -GSC locality C-5202 (one ventral valve); GSC locality 56430 (one ventral valve). Subfamily Plicatiferinae Muir- Wood and Cooper, 1960 Tribe Plicatiferini Muir- Wood and Cooper, 1960 Genus Lazarevia, new genus Type Species. — Lazarevia stepanowensis, n. sp. Derivation of Name. — ^Named in honor of the productid brachiopod specialist S. S. Lazarev. Diagnosis,— for tribe, outline transverse; both valves with vertical flanges at cardinal extremities; ornament consisting of weak, irregularly spaced rugae on visceral disc, and numerous uniform costae on trail and anterior portion of visceral disc, forming weak reticulation on anterior portion of disc; fine spine bases scat- tered sparsely over surface of ventral valve; row of several fine spines close to hingeline on each side of umbo; row of erect spine bases wrapping around ears to posterolateral margins; dorsal spines absent; ventral interior with low rim of thickened shell matter extending around auriculations and near anterior and lateral margins of valve. Comments,- — This new genus is most similar to, and possibly derived from, the Visean genus Plicatifera Chao, 1927. It differs in having much weaker, more 126 Annals of Carnegie Museum VOL. 67 irregular rugae, a shorter trail, more numerous spines on the ventral valve with a row along the hingeline and another row wrapping around the ears to the pos- terolateral margins. Internally, there is a low rim extending around the whole ventral valve. Plicatifera has strong, regular rugae on the visceral disc and a moderately long, well-developed trail. The ventral interior seems to lack the thick- ened rim seen in Lazarevia and the spine distribution is much different. Other members of this tribe, such as Ferganoproductus Galitzkaya, 1977 and Rugocon= cha Jin and Sun, 1981, can be differentiated by the following characters. Fer- ganoproductus has irregular wavy rugae on the visceral disc, lacks costae, and has numerous elongated spine tubercles crudely arranged in quincunx on the ven- tral valve. It also may have strong, erect spines on the dorsal valve. Rugoconcha is based on Plicatifera chaoi Grabau, 1936, but the few poor photographs avail- able show specimens that seem more similar to Plicatiferina Kalashnikov than Plicatifera Chao. It has strong, regular, numerous rugae over the entire surface, lacks ribs, seems to lack a trail, and has scattered spine bases. The dorsal valve appears to be unknown as does ventral valve morphology as seen from a dorsal view. Distribution. — Late Bashkirian to early Moscovian of Ellesmere Island, Ca- nadian Arctic Archipelago. Species Assigned. — Monotypic. Lazarevia stepanowensis, new species (Fig. 3.14-3.29) Holotype. — GSC 115552, Figures 3.14-3.19, a nearly complete specimen from GSC locality 56430. Paratypes.— GSC 115553, Figures 3.20-3.23, a ventral valve; GSC 115554, Figures 3.24-3.27, both valves partially preserved; GSC 1 15555, Figures 3.28 and 3.29, natural mold of ventral interior; all from GSC locality 56430. Description. — Outline transversely semicircular to subovate; maximum width at or slightly anterior to hingeline; strongly concavoconvex with thin body cavity; lateral profile subsemicircular, evenly convex. Ventral valve strongly and almost evenly convex in lateral profile; venter flattened or with weak sulcus; ears of moderate size, subangular, defined by concave flexures ventrally and nearly vertical gutterlike flanges laterally; lateral slopes moderately convex; umbonal region broad, weakly inflated; beak small; ornament consisting of numerous low, fine, irregularly spaced rugae on visceral disc and numerous fine, mostly regular costae or coarse costellae that originate on anterior portion of visceral disc and extend to anterior margin, about six or seven in five mm at 20-mm surface measure from beak; fine spine bases scattered sparsely over surface; row of several (four or more) fine spines very close to hingeline on each side of umbo; row of seven or eight erect spine bases wrapping around ears to posterolateral margins; interior smooth, lacking shagreen, in umbonal region; with low rim, strongest at auriculations, that extends around entire visceral disc except for umbo, producing weak cincture in spalled specimens; adductor scars chordate, posteriorly placed in umbonal region, moder- ately impressed; diductors not impressed, poorly delimited, weakly striate. Dorsal valve slightly less convex than opposite valve; gutterlike flanges at cardinal extremities similar to those of opposite valve; small, prominent protegulal node at posteromedial margin; fold, if present, weak and originating on anterior portion of visceral disc; ornament similar and complementary to ventral valve but weaker; spine bases not present; interior with small, sessile, bilobed cardinal process, short, thin median septum that extends forward from near base of cardinal process to about half length of visceral disc, moderately developed lateral ridges that diverge from hingeline at low angle extending about halfway toward auriculations, and numerous strong endospines anteriorly; other internal details unobserved. Measurements. — See Table 7. Comments. — This Arctic species bears some superficial resemblance to species 1998 Carter and Poletaev — Upper Carboniferous Arctic Brachiopods 127 Table 7. — Measurements (in millimeters) o/Lazarevia stepanowensis n. sp. from GSC locality 56430. GSC number Length Width Height Surface length 115552 22.3 32.6 14.2 35.3 155553 25.3 40.0 16.5 39.0 115554 24.3 39.0 15.8 36.2 of the genus Desmoinesia Hoare but can readily be differentiated from that genus by its larger size, more transverse outline, lack of dorsal spines, the presence of lateral flanges, and a weak cincture on spalled surfaces of the ventral valve that reflects a low thickening or rim that extends around the interior of the ventral valve. Plicatifera pseudopUcatilis (Muir- Wood), as illustrated in Brunton et al. (1993), is similar in size, transverse outline, and general ornament but differs in having much stronger and more regular rugae, a longer trail, and an almost completely reticulate visceral disc. Distribution.=--T\ii^ species is common at GSC locality 56430 (90 specimens). Tribe Semicostellini Nalivkin, 1979 Genus Maemia Lazarev, 1997 Maemia gelida, new species (Fig. 4.24-=-4.43) Hoiotype.^GSC 115562, Figures 4.24-4.28, from GSC locality C-5202. Paratypes. — -GSC 1 15563--1 15566, Figures 4.29-4.43, from GSC locality C--5202. Description. — Small, nearly planoconvex; outline transversely subovate; greatest width at about midleegth, rarely at hingeline; ears small, compressed, subangular; anterior profile subsemicircular to rounded subtrapezoidal; lateral profile subsemicircular. Ventral valve strongly inflated, evenly convex except for more convex umbonal region; visceral disc not delineated; venter weakly convex, flanks dropping steeply to lateral margins; trail very short, not well defined; ears set off by deeply concave flexures; umbonal region weakly inflated and of moderate width; beak small, slightly overhanging hingeline; entire valve weakly and irregularly rugose and slightly lamellose, each strong concentric stricture anterior to strong rugae indicating former position of dorsal visceral disc; coarse, slightly elongate nodular spine bases or tubercles arranged crudely in quincunx over most of valve; finer, more numerous spines on ears, on sides of umbo, and near hingeline; interior not observed. Dorsal valve almost flat or very weakly concave, except for very short, sharply geniculated trail; small medial protegulal node present; ornament consisting of weaker rugae than those of opposite valve and scattered shallow dimples; numerous fine, scattered spine bases present; interior not ob- served. Measurements.- — See Table 8. Table %. -—Measurements (in millimeters) of the types o/ Maemia gelida n. sp. from GSC locality C- 5252. GSC number Length Width Height Surface length 115562 10.2 13.0 6.1 16.9 115563 8.5 10.6 5.4 13.4 115564 9.9 12.3 6.6 17.2 155565 9.5 11.5 6.7 16.1 115566 10.1 11.5 6.5 17.5 128 Annals of Carnegie Museum VOL. 67 1998 Carter and Poletaev — Upper Carboniferous Arctic Brachiopods 129 Diagnosis. — The combined characters of a rugose, spinose, noncostate oma- ment and flat, spinose dorsal valve with very short trail characterize this species. Comments. — Despite the small size of this new species its general morphology suggests that it is closely related to the recently described genus Maemia Lazarev. The type species of that genus, Maemia chaykensis Lazarev, 1997 (in Brunton and Lazarev, 1997), however, is much larger and bears a shallow ventral sulcus and coarse costae on the trail. Another species from the same horizon at Cape Chaika, Maemia nana Lazarev, however, is similar to this Canadian species in size and ornament. It differs in having much weaker spine base tubercles and weak, coarse ribbing on the ventral valve. Distribution.— GSC locality C~5202 (25 specimens); GSC locality 56430 (six specimens). Family Productidae Gray, 1840 Subfamily Dictyoclostinae Stehli, 1954 Genus Reticulatia Muir- Wood and Cooper, 1960 Reticulatia cf. R. americanus (Dunbar and Condra, 1932) (Fig. 5. 1-5.8) 1932 Dictyoclostus americanus Dunbar and Condra, p. 218, pi. 34, fig. 3-6. Description. — Large, moderately concavoconvex; outline transversely subovate to subquadrate; maximum width probably near hingeline (ears incomplete in all specimens); lateral profile subsemi- circular; anterior profile subtrapezoidal; fold and sulcus moderately developed; body cavity moderately thick. Ventral valve moderately inflated, almost evenly convex, most convex near beak; umbo broad, moderately produced, subtending an angle well in excess of 90 degrees; beak small, slightly over- hanging hingeline; flanks moderately convex, sloping steeply to lateral margins; rounded sulcus orig- inating in umbonal region, remaining shallow, sometimes obscure, throughout entire length; ears de- limited by concave flexures, not fully preserved; visceral disc delineated by weak geniculation and loss of reticulate ornament; ornament of visceral disc strongly and regularly reticulate with numerous, almost regular rugae intersecting regular coarse costae, about four or five per five mm on anterior portion of visceral disc; spine bases found in row along hingeline, around ears, and scattered elsewhere; interior unknown. Dorsal valve moderately concave, geniculate; fold low throughout; ears delimited by convex flex- ures; trail not well preserved; ornament of visceral disc complementary to opposite valve; no spines observed; interior unknown. Comments .-—Th&sc incomplete Arctic specimens agree in most observable de- tails with typical Reticulatia americanus (Dunbar and Condra), of Missourian or Kazimovian age of Nebraska and surrounding environs. The latter differs little from the Hare Fiord specimens, mainly in having numerous spines scattered on the trail. Muir- Wood and Cooper (1960) emphasized the well-developed gingly- mus in this genus, a character not preserved in our specimens. The trail also is broken in all of our specimens. Therefore, identification is tentative. Fig. 5. — Productoids. 5. 1-5.8, Reticulatia cf. R. americanus (Dunbar and Condra, 1932), ventral, lateral, posterior, and anterior views of two ventral valves, GSC 115567 and 115568, X 1. 5.9-5.16, Kutorginella cf. K. mosquensis Ivanova, 1951; 5.9, 5.10, ventral and oblique views of ventral valve with siphon, GSC 115569; 5.11-5.16, ventral, dorsal, mold of a dorsal exterior, posterior, lateral, and anterior views of a small complete specimen, GSC 115570; both X 1. 5.17-5.19, Antiquatonia cf. A. hermosana (Girty, 1903), ventral, posterior, and lateral views of a ventral valve showing the spine ridge on the right ear, GSC 115571, X 1. 130 Annals of Carnegie Museum VOL, 67 1998 Carter and Poletaev — Upper Carboniferous Arctic Brachiopods 131 Distribution. — GSC locality 60194 (two ventral valves and two articulated specimens). Tribe Kozlowskiini Genus Eomarginifera Muir- Wood, 1930 Eomarginifera sp. (Fig. 6.1—6.10) Comments. — This species is assigned to Eomarginifera on the basis of having three pairs of halteroid spines on the ears, umbo, and trail of the ventral valve. Deep concave grooves defining the ears indicate that there are strong lateral ridges curving around the ears in the dorsal valve as in Eomarginifera. A large fragment of conjoined valves shows the trails of both valves touching each other at the anterior margin, obviating the possibility that this species could be assigned to Kozlowskia which has numerous short trails bunched at the anterior margin. The transverse outline and widely spaced pair of halteroid spines on the trail dem- onstrate that it cannot be assigned to Eomarginiferina Brunton, 1966. Assignment to a species of Eomarginifera is more difficult. The coarse ribs and rounded venter of this Arctic species are not usual for the genus Eomargin- ifera. Externally, the ventral valves resemble the species described as Kozlowskia sp. by Lazarev (1990:pL 21, fig. 24b, v), of Moscovian age from the Moscow Basin, Distribution. — GSC locality 56430 (five ventral valves, two dorsal valve molds, one fragment of conjoined valves); GSC locality C-5202 (one ventral valve). Genus Kozlowskia Frederiks, 1933 1 Kozlowskia splendens (Norwood and Pratten, 1855) (Fig. 6.16-6.27) 1855 Productus splendens Norwood and Pratten, p. 11, pi. 1, fig. 5a-d. Description. — Small; outline transversely subsemicircular to subtrapezoidal; greatest width at hinge- line; lateral profile subtrigonal to subsemicircular; anterior profile subtrapezoidal to subsemicircular. Ventral valve strongly inflated with moderately convex visceral disc and less convex geniculated trail; ears large, slightly mucronate or compressed, well delineated by concave flexures; venter rounded to flattened or weakly sulcate; flanks steeply sloping to lateral margins; radial ornament of low, weak costellae, confined to trail and that portion of visceral disc anterior to umbonal region; visceral disc with weak rugae, forming weakly reticulate pattern near point of geniculation; three pairs of halteroid spines at ears, sides of visceral disc, and on trail; few finer spine bases scattered near hingeline and on umbo; interior unknown. Fig. 6. — Productoids. 6.1-6.10, Eomarginifera sp.; 6. 1-6.4, 6.7-6.10, ventral, anterior, posterior, and lateral views of two ventral valves, GSC 115572 and 115573; 6.5, 6.6, ventral and anterior views of natural mold of dorsal exterior, GSC 115574; all X 1.5. 6.11-6.15, 1 Eomarginiferina sp.; 6.11-6.14, ventral, anterior, posterior, and lateral views of ventral valve, GSC 115575; 6.15, ventral view of natural mold of dorsal exterior, GSC 115576; all X 1.5. 6.16-6.27, ^Kozlowskia splendens (Norwood and Pratten, 1855), ventral, anterior, posterior, and lateral views of three ventral valves, GSC 115577- 115579, respectively, X 1.5. 6.28-6.30, ventral, lateral, and posterior views of Krotovia cf. K. lamel- losa Brunton, 1966, GSC 115580, X 3. 6.31, 6.32, ventral and posterior views of Krotovia cf. K. spinulosa (J. Sowerby, 1814), GSC 115581, X 3. 6.33-6.43, IFimbrinia borealis n. sp.; 6.33-6.40, ventral, anterior, posterior, and lateral views of two ventral valves, including the holotype (6.33-6.36), GSC 115582 and 115583; 6.41-6.43, ventral, lateral, and anterior views of a mold of the dorsal exterior, GSC 115584; all X 2. 132 Annals of Carnegie Museum VOL. 67 Dorsal valve known only from externa! molds. Trail sharply geniculated; ears smaller than opposite valve; visceral disc flattened, weakly convex; ornament as in opposite valve but weaker; spines absent; evidence of numerous trails lacking; interior unknown. Comments. — The large ears, variably developed fold, and obsolescent costellae on the trail characterize these Hare Fiord specimens. In this respect these Arctic specimens are virtually indistinguishable from typical specimens of Koziowskia splendens from Desmoinesian and Missourian beds in the midcontinent, an ad- mittedly highly variable species according to Dunbar and Condra (1932), The absence of specimens that indicate the presence of a sheaf of closely packed short trails at the dorsal margin is unfortunate. If there is no dorsal rim of numerous trails this species is not a Koziowskia and might be reassigned to another genus such as Eomarginiferina Brunton, 1968. However, the weak retic- ulation and anterior ribbing in this Arctic species are more suggestive of Koziows- kia than Eomarginiferina. Distribution. — GSC locality 56430 (six ventral valves); GSC locality 56430A (five ventral valves). Tribe Retariini Muir- Wood and Cooper, 1960 Genus Kutorginella Ivanova, 1951 Kutorginella cf. K. mosquensis Ivanova, 1951 (Fig, 5.9-5.16) 1951 Kutorginella mosquensis Ivanova, p. 329. Description. — Medium size; outline subquadrate; lateral profile strongly inflated, subtrapezoidal; anterior profile subquadrate to subtrapezoidal; point of maximum width apparently anterior to hinge- line; ears small to medium; fold and sulcus moderately well developed, originating in umbonal region, becoming broader and deeper anteriorly; visceral disc reticulate with numerous rugae intersecting moderately strong costae; costae numbering about five or six in five mm on trail, extending forward over entire trail. Ventral valve moderately geniculated; ears defined by concave flexures; lateral slopes dropping steeply to lateral margins; trail long with medial tube-like extension formed by distinctive stricture in largest specimen; spine bases scattered over umbonal region and trail with regular row wrapping around each ear; spines along hingeline not observed due to poor preservation; interior not observed. Dorsal valve with flattened visceral disc and subquadrate outline; ears delimited by low convex flexures; dorsum with low fold; trail at least eight to ten mm long; ornament reticulate; spines absent; irregular rows of rounded pits on ears; interior unknown. Comments. —ThQ genus Kutorginella Ivanova ranges from the uppermost Ser- pukhovian to the Kungurian and many species have been described. These Hare Fiord specimens resemble K. mosquensis Ivanova from the Kazimovian of the Moscow Basin in ornament and development of an incomplete siphonal or tube- like trail extension in large, fully mature specimens. They also fall within the size range of specimens illustrated by Sarycheva (1971:pL 5, fig. la, b) and Sarycheva and Sokolskaya (1952:pL 37, fig. 230). However, they differ from the specimens illustrated by Sarycheva and Sokolskaya (1952) and Lazarev (1990:pL 19, fig. 1) in being less transverse and the ears appear to be smaller, although these Hare Fiord specimens are not complete. Distribution. — -GSC locality 56430 (one ventral valve); GSC loc. 4085 (one specimen); GSC locality C-5202 (one ventral valve). Genus Antiquatonia Miloradovich, 1945 Antiquatonia cf. A. hermosana (Girty, 1903) (Fig. 5.17-5.19) 1903 Productus semireticulatus var. hermosanus Girty, p. 358, pi. 2, fig. 1-4. Description. — Large, strongly inflated; outline subovate to subquadrate; widest at hingeline; ears 1998 Carter and Poletaev — Upper Carboniferous Arctic Brachiopods 133 large, well defined; ventral sulcus narrow, shallow; visceral disc strongly reticulate, slightly geniculated from long trail; trail costate with about seven to nine costae per cm near anterior margin; ears set off by conspicuous ridge that wraps around ears; coarse spine bases sparsely scattered on trail and visceral disc; spines on ears and umbonal region obscured by poor preservation; other details not observed. Comments. — The umbonal region of this single ventral valve is slightly crushed. Otherwise this specimen closely resembles the large ventral valve illus- trated by Girty (1903:pL 3, fig. 1-lc). Sutherland and Harlow (1973:pL 11, fig. 6a-=c) illustrated a smaller narrower specimen that otherwise is similar to this Arctic specimen. Distribution. — ^GSC locality 56430 (one ventral valve). Superfamily Echinoconchoidea Family Echinoconchidae Stehli, 1954 Subfamily Pustulinae Waterhouse, 1981 Genus Pustula Thomas, 1914 IPustula sp. (Fig. 7.2) Comments. — This single incomplete, slightly distorted ventral valve has mostly slightly elongate spine bases arranged in quincunx, with distinctive scattered, rounded pits between some of the spine bases. There are a few weak, irregular rugae, most noticeable around the ears. The low, rounded profile and vaguely rounded outline combined with this peculiar ornament suggest assignment to the genus Pustula, although that genus is more rugose and does not normally have rounded pits between the spine bases of the ventral valve. Distribution.— ~GSC locality C-5202 (one ventral valve). Subfamily Echinoconchinae Stehli, 1954 Tribe Echinoconchini Stehli, 1954 Genus Echinaria Muir- Wood and Cooper, 1960 lEchinaria sp. (Fig. 7.1) Comments. — A single large, partial echinoconchoid ventral valve seems likely to belong in the genus Echinaria. Although spine bases cannot be discerned, the shape and size of this specimen suggest assignment here, although a dorsal interior is needed for certain identification. Distribution.— GSC locality 56430 (one partial ventral valve). Superfamily Linoproductoidea Stehli, 1954 Family Monticuliferidae Muir- Wood and Cooper, 1960 Subfamily Auriculispinae Waterhouse, 1986 Genus Liraria Cooper and Grant, 1975 Liraria paucispina, new species (Fig. 7.3-7.18) Holotype. — GSC 115587, Figures 7.3, 7.11-7.14, a ventral valve, from GSC locality C-5202. Paratypes. — GSC 115588, Figures 7.4, 7.15-7.18, a ventral valve; GSC 115589, Figures 7.5 and 7.6, a complete specimen with the visceral disc removed showing the mold of the dorsal exterior; GSC 115590, Figures 7.7-7.10, a large ventral valve; all from GSC locality C-5202. Description. — Small to medi-um size; strongly concavoconvex; body cavity thin; outline subovate; 134 Annals of Carnegie Museum VOL. 67 23 24 2 5 2 6 2 7 28 Fig. 7. — Productoids. 7.1, lEchinaria, ventral valve, GSC 115585, X 1. 7.2, IPustula sp., ventral valve, GSC 115586, X 2. 7.3-7.18, Liraria paucispina n. sp.; 7.3, 7.11-7.14, ventral posterior view showing attachment scars and spine bases at hinge, X 4, and ventral, posterior, anterior, and lateral views of the holotype, X 1.5, GSC 115587; 7.4, 7.15-7.18, ventral posterior of a paratype showing attachment scars and spine bases at hinge, X 4, and ventral, posterior, anterior, and lateral views of paratype GSC 115588, X 1.5; 7.5, 7.6, ventral and posterior views of complete paratype with visceral disc removed, GSC 115589, X 1.5; 7.7-7.10, ventral, posterior, anterior, and lateral views of a large ventral valve paratype, GSC 115590, X 1.5. 7.19-7.22, Fluctuaria cf. F. undata (Defrance, 1826), ventral, anterior, posterior, and lateral views of ventral valve, GSC 115591, X 1. 7.23-7.26, IFino- productus sp.; 7.23-7.25, ventral, posterior, and lateral views of ventral valve, GSC 115592, X 1; 7.26, ventral view of mold of dorsal exterior, GSC 115593; X 1. 7.27, 7.28, Cancrinella sp., ventral exterior and mold of dorsal exterior, GSC 115594 and 115595, X 2. 1998 Carter and Poletaev — Upper Carboniferous Arctic Brachiopods 135 Table 9. — Measurements (in millimeters) o/Liraria paucispina n. sp. from GSC locality C-5202. GSC number Length Width Height Surface length 115589 17.5 17.8 + 8.8 27.7 115590 16.9 19.3 7.2 25.6 115587 15.2 17.0 8.7 23.4 115588 12.7 17.7 6.5 20.5 maximum width anterior to hingeline in most specimens; lateral profile strongly convex, most convex in umbonal region; anterior profile and venter almost evenly rounded; trails not produced or differ- entiated. Ventral valve strongly inflated posteriorly, most convex in umbonal region; umbo broad; beak small, scarcely overhanging hingeline, with up to three pairs of fine, vertically oriented attachment spines; ears small, flattened, delineated by concave flexures; ornament of fine, rounded costellae, about 1 1 to 13 per five mm at a surface distance of 15 mm from beak, which increase by intercalation; few rugae limited to sides of umbo and lateral slopes; one or two pairs of prostrate, elongate, laterally or an- terolaterally directed spine bases on ears just anterior to hingeline; other spines rare; spines along hingeline usually absent, rarely with one or two pairs of erect spines at hingeline; growth lines very fine, regularly spaced; interior unknown. Dorsal valve with moderately concave visceral region; prominent rounded protegulal node at hinge- line; ears small, defined by convex flexures; fine costellae and rugae as in opposite valve; spines not observed, apparently absent; interior unknown. Measurements Table 9. Diagnosis. —This species is characterized by modest size, subovate outline, and one pair (rarely two pairs) of elongated, laterally directed, fine spine bases just anterior to the hingeline. Comments. — Liraria paucispina, n. sp., can be differentiated from the type and only other described species, Liraria lirata Cooper and Grant, 1975, from the basal Bone Spring Formation (Leonardian-Artinskian) of west Texas, by its mod- est size, subovate outline, and pair (rarely two pairs) of elongated, laterally di- rected, fine spine bases near the hingeline. This species seems to have been attached throughout life normal to its substrate. All observed attachment areas on the ventral beak bear fine, elongated, vertical impressions such as might be made by productid spines. Nearly all of the spec- imens from GSC locality C-5202 were recovered from a single small lump of limestone crowded with these shells. It is possible that the spat of this species tended to attach to spines of adults of the same species. Distribution. — GSC locality C-5202 (100+ specimens, mostly ventral valves); GSC locality 56430 (three ventral valves, one dorsal valve mold); GSC locality 56430 A (50 ventral valves). Family Linoproductidae Stehli, 1954 Subfamily Linoproductinae Stehli, 1954 Genus Linoproductus Chao, 1927 ILinoproductus sp. (Fig. 7.23-7.26) Description. — Medium size; moderately concavoconvex; outline transversely subovate; maximum width near midlength; ears small, slightly compressed; body cavity estimated to be moderately thick; lateral profile rounded, most convex in umbonal region; anterior profile subtrapezoidal. Ventral valve strongly inflated posteriorly; umbonal region broad; beak small, not appreciably over- hanging hingeline; venter flattened or weakly sulcate; flanks weakly convex, sloping moderately steep- ly to lateral margins; trail, if present, short; entire surface with fine costellae, about nine to 1 1 per 136 Annals of Carnegie Museum VOL. 67 five mm near front margin, which increase by intercalation; three to four rugae on sides of umbo; spine bases not observed, probably because of poor preservation; interior unknown. Dorsal valve moderately concave; ears delineated by weakly convex flexures; rugae strongest near hingeline, one or two of which may extend around visceral disc; ribbing as in opposite valve; spines absent; interior unknown. Comments.-— ¥oot preservation of the ventral valve and lack of a dorsal interior makes generic identification difficult. In general aspect this species is most similar to Linoproductus Chao but this assignment is far from certain. Distribution.— GSC locality 56430 (two ventral valves; one mold of dorsal valve exterior). Genus Fluctuaria Muir- Wood and Cooper, 1960 Fluctuaria cf. F. undata (Defrance, 1826) (Fig. 7.19-7.22) 1826 Productus undatus Defrance, p. 354. Description. — This description is based on a single, nearly complete ventral valve. Medium size; strongly convex; outline transversely subovate; greatest width anterior to midlength; lateral profile almost evenly convex; umbonal region most convex, moderately broad; beak small, slightly overhanging hingeline; venter more weakly convex or slightly flattened; flanks moderately spreading, sloping steeply to lateral margins; ears small, defined by slightly concave flexures; entire surface finely costellate with about 11 to 13 costellae per five mm at surface measure of 20 mm from beak; costellae increase by intercalation; entire valve strongly rugose; rugae irregular, sometimes dis- continuous, strongest at sides of umbo; spine bases rare on trail, not observed near hingeline because of poor preservation; ventral interior and dorsal valve unknown. Comments. — This specimen is similar to the broader specimens of Fluctuaria undata figured by Koninck (1843;pL 12, fig. 2; 1847:pL 5, fig. 3) from the Visean of Belgium, and Sarycheva (1937:pL 7, fig. 2a; Sarycheva et al., 1963:pL 37, fig. 9) from the Serpukhovian of European Russia and the Kuznets. Distribution.— GSC locality 56430 (one good ventral valve, five questionable ventral valves). A similar form occurs in the Ladrones Limestone of southeastern Alaska. Subfamily Grandaurispininae Lazarev, 1986 Genus Cancrinella Frederiks, 1928 Cancrinella sp. (Fig. 7.27, 7.28) Comments. — -There are three imperfectly preserved specimens of this genus in the Hare Fiord collections, all from GSC locality 56430, The distinctive fine radial ornament of this species with elongate quincuntially arranged spine bases on Fig. 8. — Stenoscismatoids and rhynchonelloids. 8. 1-8,4, Stenoscisma sp., ventral, dorsal, anterior, and lateral views (beak removed by grinding), GSC 115596, X 1. 8.5-8.16, Careoseptum septentrionalis n. gen. n. sp., ventral, dorsal, anterior, and lateral views of three specimens, including the holotype (8.9-8.12) and two paratypes, GSC 115597-115599, respectively, X 2. 8.17-8.20, Cenorhynchia sp., ventral, dorsal, anterior, and lateral views, GSC 115600, X 2, 8.21-8.28, Exlaminella insoiita n. gen. n. sp., ventral, dorsal, anterior, and lateral views of two specimens, including the holotype (8.25-8.28, GSC 1 15601), and the paratype, GSC 1 15602, X 2. 8.29-8.40, lElassonia sverdrupensis n. sp., ventral, dorsal, anterior, and lateral views of three specimens, including the holotype (8.29-8.32) and two paratypes, GSC 115603-115605, respectively, X 2. 8.41-8,48, IHemileurus sp., ventral, dorsal, an- terior, and lateral views of two specimens, GSC 115606 and 115607, X 2. 1998 Carter and Poletaev — Upper Carboniferous Arctic Brachiopods 137 138 Annals of Carnegie Museum VOL. 67 Fig. 9. — Transverse serial sections of Stenoscisma sp., GSC 1 15608, X 2.5. Numbers refer to distance in millimeters from ventral beak. swollen costellae clearly attest to its generic affinities. However, these few poor, incomplete specimens do not allow specific assignment. Distribution.— GSC locality 56430 (two ventral valves and one dorsal valve). Order Pentamerida Schuchert and Cooper, 1931 Superfamily Stenoscismatoidea Oehlert, 1887 Family Stenoscismatidae Oehlert, 1887 Genus Stenoscisma Conrad, 1839 Stenoscisma sp. (Fig. 8.1--8.4, 9) Description. — This description is based on four specimens, including two complete shells, one ventral valve, and one dorsal valve. One complete specimen was sectioned and is illustrated in Figure 9. The largest complete shell is illustrated in Figure 8. The beak of this weathered specimen was ground down to confirm the presence of a spondylium. Medium size, moderately to strongly transverse, strongly inequivalved; ventral flanks concave, marked by about three coarse, subangular plicae and possibly a fourth much weaker one; ventral umbonal region smooth; fold and sulcus well developed and marked by two or three strong, angular plicae and up to two much weaker parietal plicae; dorsal valve strongly inflated with convex flanks and ornament complementary to opposite valve. Ventral interior with deep spondylium elevated well above floor; teeth small, blunt; septum duplex extending anterior to spondylium. Dorsal interior with short, wide-set sockets; fimbriate cardinal process supported by short, slightly convex hingeplate and long, high median septum; camarophorium narrow, concave, moderately long, extending forward well anterior to dorsally attached portion of septum, with short, low intercamaro- phorial plate; crura not observed. Comments. — The few poor specimens of this species do not allow precise iden- tification. We have not been able to find mature individuals of another species of this genus similar in size and proportions to this Canadian species. Stenoscisma mutabilis (Chernyshev, 1902) from the Lower Permian of Russia is a highly 1998 Carter and Poletaev — Upper Carboniferous Arctic Brachiopods 139 variable species, some smaller unusually transverse individuals of which might be generally similar in outline and ornament to this Arctic species. Distribution, — GSC locality 60194 (three specimens); GSC locality 56430A (one poor ventral valve). Family Atriboniidae Grant, 1965 Subfamily Psilocamarinae Grant, 1965 Genus Careoseptum, new genus Type Species.— Careoseptum septentrionalis, n. sp. Derivation of Name. — From the Latin careo, without or lacking; and septum, partition. Diagnosis. — Small, strongly biconvex; flanks smooth; fold and sulcus rounded, smooth, or with weak median rib; median septa lacking or vestigial in both valves; intercamarophorial plate lacking; hingeplate complete in mature shell, forming septalium with camarophorium. Comments. — The lack of a dorsal septum makes this genus unique for the superfamily Stenoscismatoidea. The absence of an intercamarophorial plate places it in the family Atriboniidae, subfamily Psilocamarinae. Within this subfamily Careoseptum is externally similar only to the genus Psilocamara Grant, 1965, described from the Moscovian of Texas and Missouri. Internally, the two genera are greatly different. Distribution. — Lower Moscovian of Arctic Canada. Species Assigned.— KsLOwn only from the type species. Careoseptum septentrionalis, new species (Fig. 8.5-8.16, 10) Holotype. — Figures 8,9-8.12, GSC 115598, from GSC locality C-5202. Paratypes. — Figures 8. 5-8. 8, GSC 115597, from GSC locality 56430; Figures 8.13-8.16, GSC 115599, from GSC locality 56430; Figure lOA, GSC 115609, lOB, GSC 115610. Description. — Medium size, outline subovate, strongly inequivalved; fold and sulcus absent poste- riorly, strongly produced anteriorly, rounded, smooth, or with weak median rib; lateral slopes smooth; stolidium not apparent; shell matter thickened posteriorly. Ventral valve weakly convex; umbonal region subtending an angle greater than 90 degrees; beak small, subangular, projecting slightly posterior to dorsal valve; foramen small, rounded; delthyrium apparently open; spondylium supported by thick callus posteriorly, free anteriorly; teeth broad, blunt. Dorsal valve thick, evenly convex on flanks; fold originating in posterior third to half of valve; umbonal region slightly swollen, compressed laterally; septum lacking; camarophorium unsupported except at posterior margin, covered posteriorly; intercamarophorial plate lacking; crura not observed. Measurements. — See Table 10. Diagnosis. as for the genus. Comments.- — The authors know of no other species internally similar to Car- eoseptum septentrionalis. Table 10. — Measurements (in millimeters) of the types o/ Careospetum septentrionalis, n. sp. GSC number Locality Length Width Thickness 115597 56430 10.6 10.7 8.1 115598 C-5202 9.6 9.0 6.4 115599 56430 9.5 8.5 6.5 140 Annals of Carnegie Museum VOL. 67 SS8888 1.5 1.6 ^ ^ ^ ouOoo 2.4 2.6 2.8 3.0 3.2 0.6 ^ © © © 1.0 1.1 1 2 O o r> v_y viy 'ey u/ B 1.8 1.6 1.4 1.3 Fig. 10. — Transverse serial sections of Careoseptum septentrionalis n. gen. n. sp. Numbers refer to distance in millimeters from ventral beak. A. Large mature specimen, GSC 115609, X 4. B. Small specimen, GSC 115610, X 5. 1998 Carter and Poletaev — Upper Carboniferous Arctic Brachiopods 141 Fig. 1 1. — Transverse serial sections of Cenorhynchia sp., GSC 115611, X 5. Numbers refer to distance in millimeters from ventral beak. Distribution. — GSC locality 56430 (eight specimens); GSC locality C-5202 (two specimens). Order Rhynchonellida Kuhn, 1949 Superfamily Rhynchonelloidea Gray, 1848 Family Wellerellidae Likharev, 1956 Genus Cenorhynchia Cooper and Grant, 1976 Cenorhynchia sp. (Fig. 8.17-8.20, 11) Description. — Medium size for genus, unequally biconvex; outline rounded subtrigonal, lateral pro- file subelliptical; anterior or posterior profile subelliptical; umbonal region moderately broad, subtend- ing an umbonal angle of about 90 degrees; fold and sulcus moderately developed and moderately wide; flanks of both valves smooth; fold and sulcus with few strong plicae only in anterior third of valves. Ventral valve moderately convex in lateral profile; beak slightly incurved, small; deltidial plates not observed; foramen not observed; sulcus originating anterior to midlength; sulcus with two strong, sounded plicae on tongue; interior with stout, short, medially concave dental plates. Dorsal valve with flattened umbonal region; flanks moderately convex, sloping steeply to lateral margins; fold originating anterior to midlength, rising little above lateral slopes, with three strong, rounded plicae confined to anterior third of valve; interior with complete hingeplate covering short septalium; median septum long, very high, nearly in plane of lateral commissure; sockets wide, defined by high socket ridges; crura not observed. Comments .—This, late Bashkirian or early Moscovian Arctic species is similar in its proportions and ornament to Cenorhynchia saginata Cooper and Grant, 1976, and Cenorhynchia triangulata Cooper and Grant, 1976, from the Permian of west Texas. It differs from the former in its larger size and complete lack of lateral ribbing. Cenorhynchia triangulata is less similar to this Arctic species with its inflated dorsal umbo, obscure lateral ribbing, and laterally compressed ventral beak. Distribution. — GSC locality 56430 (three specimens, one sectioned). 142 Annals of Carnegie Museum VOL. 67 Genus Phrenophoria Cooper and Grant, 1969 IPhrenophoria sp. (Fig. 15.20-15.24) Comments. — single complete specimen externally similar to the genus Phren- ophoria Cooper and Grant was not sectioned. Dental plates and a strong dorsal median septum are clearly present which supports this generic assignment but the lack of other internal details prevents certain generic placement. A single disar- ticulated dorsal valve is nearly identical to that of the complete specimen but adds no additional information concerning morphology. Distribution. — GSC locality 60194 (two specimens). Family uncertain Genus Exlaminella, new genus Type Species. — Exlaminella insolita, n. sp. Derivation of Name. — From the Latin ex, without; lamina, plate; and ella, feminine diminutive. Diagnosis. — Small, unequally biconvex; outline rounded subtrigonal to subov- ate; flanks smooth; fold and sulcus with plicae confined to anterior half of valves; interior lacking dental plates or dorsal septum; hingeplate divided, crura falcifer; shell posteriorly thickened in both valves. Comments. — This genus is unusual in its lack of lateral ribbing and internal plates. The latter feature is found in the Permian genera lotina Cooper and Grant, 1976, and Ptilotorhynchus Cooper and Grant, 1976, where the dental plates are fused or suppressed. Both of these genera have strong ribbing on the flanks and much different growth form. Hemileurus Cooper and Grant, 1976, also Permian, possesses a similar dorsal interior and ribbing confined to the anterior half of the valves. The ventral interior is conventional with long, slender dental plates. Nev- ertheless, Hemileurus is more likely related to Exlaminella than the other Permian genera if growth form, ornament, and dorsal interior are of paramount importance in the phytogeny of this group. In the meantime, we cannot definitely assign this new genus to any known family. Species Assigned. — Monotypic. Stratigraphic Range.— Bashkirian or early Moscovian of Arctic Canada. Exlaminella insolita, new species (Fig. 8.21-8.28, 12) Holotype. — Figures 8.25-8.28, GSC 115602, from GSC locality 56430. Paratypes.—¥\guvQS 8.21-8.24, GSC 1 15601, from GSC locality C-5202; Figure 12A, GSC 1 15612, from GSC locality C-5202; Figure 12B, GSC 115613, from GSC locality C-5202. Description. — Ventral valve weakly convex; sulcus originating at about midlength, with two strong subangular plicae anteriorly; umbonal region moderately broad, subtending an angle of 90 degrees or more; beak small, slightly incurved; foramen not observed; delthyrium not observed; ventral interior with thick shell matter but with no indication of dental plates. Dorsal valve moderately convex in umbonal region, sloping steeply to lateral margins; fold origi- nating near or posterior to midlength, rising moderately to high, marked by three short, strong, rounded plicae anteriorly; dorsal interior with thick callus deposits posteriorly; hingeplate divided; crura falcifer, remaining in plane of lateral commissure; low, thick median ridge variably developed. Measurements. — See Table 1 1 . Diagnosis. — Same as for genus. Comments. — Externally this species is similar in size and growth form to the 1998 Carter and Poletaev — Upper Carboniferous Arctic Brachiopods 143 Fig. 12. — Transverse serial sections of Exlaminella insolita n. gen. n. sp. Numbers refer to distance in millimeters from ventral beak. A. Large mature paratype, GSC 115612, X 4. B. Juvenile paratype, GSC 115613, X 4. 144 Annals of Carnegie Museum VOL. 67 Table 1 1. — Measurements (in millimeters) o/ Exlaminelia insolita, gen. n. sp. from the lower Hare Fiord Formation. GSC number Locality Length Width Thickness 115601 C-5202 8.4 9.3 6.5 115602 56430 9.9 9.7 8.0 stenoscismatoid genus Careoseptum n. gen. and differs externally only in the possession of the short anterior plicae in the fold and sulcus. Careoseptum sep- tentrionalis n. gen. n. sp., the type and only species, may have a weak single plica in the sulcus but never has two as does Exiaminella insolita. Internally the two species are very different. Cenorhynchia sp. is also externally similar to Exiaminella insolita in growth form, size, and ornament. The former differs in= temally in its high dorsal septum and stout dental plates. Distribution.— GSC locality 56430 (two specimens); GSC locality C“5202 (three specimens, two sectioned). Family Petasmatheridae Cooper and Grant, 1976 Genus Elassonia Cooper and Grant, 1976 lElassonia sverdrupensis, new species (Fig. 8.29-8.40, 13) Holotype. —Pigums 8.29-8.32, GSC 115603, from GSC locality 56430. Paratypes. ~¥\gnms 833-8.40, GSC 115604, 115605, and 115614, from GSC locality 56430. Description. — Small, flattened, nearly equally biconvex, outline rounded subtrigonal to guttate; lat- eral and anterior profiles lenticular; fold and sulcus lacking; anterior commissure rectimarginate; both valves completely costate to beaks with numerous bifurcating, intercalating, or simple costae; umbonal regions evenly elongated, usually subtending an angle of less than 90 degrees, margins slightly com- pressed. Ventral valve slightly thicker than dorsal valve, weakly and evenly convex in lateral profile; venter weakly convex or flattened; flanks sloping nearly normal to lateral margins, forming subangular beak ridges; beak small, undifferentiated from umbo, suberect; foramen not observed; interior with short, medially concave, thin dental plates. Dorsal valve more evenly rounded and slightly compressed laterally in umbonal region; dorsum anteriorly flattened or with faint sulcus; otherwise similar in convexity to opposite valve; interior with very short septalium supported by stout, moderately long median septum; hingeplate divided or un- divided; crura falcifer, thin, curving ventrad. Measurements. See Table 12. This species can be differentiated by small size, thin profile, tri- angular outline, the lack of a fold and sulcus, and bifurcating costae. Comments. — These specimens, lacking a fold and sulcus, resemble juvenile specimens of much larger species. However, we are convinced that these 14 spec- imens are mature individuals because of the uniformity in size of the specimens. Assignment of this unusual species to the Permian genus Elassonia Cooper and Grant is arbitrary. We have little doubt that it represents an undescribed genus but the indifferent preservation of this collection and lack of pristine specimens inhibits us from proposing one. Distribution.— GSC locality 56430 (14 specimens, eight complete or nearly complete, two sectioned). 1998 Carter and Poletaev — Upper Carboniferous Arctic Brachiopods 145 O Q o 09 nn 0-8 0.9 VI/ 1.0 1.1 Fig. 13. — Transverse serial sections of lElassonia sverdrupensis n. sp., paratype, GSC 115614, X 5. Numbers refer to distance in millimeters from ventral beak. Family Allorhynchidae Cooper and Grant, 1976 Genus Hemileurus Cooper and Grant, 1976 IHemileurus sp. (Fig. 8.41-8.48, 14) Description. — Small, almost equally biconvex, outline longitudinally subovate; lateral and anterior profiles thinly lenticular; fold and sulcus lacking; anterior commissure rectimarginate; umbonal regions smooth, anterolateral margins with weak costellae; beak ridges rounded. Ventral valve slightly thicker than dorsal valve, evenly convex; umbonal region slightly extended, subtending an angle of about 90 degrees; beak small, nearly straight, inconspicuous; posterolateral margins slightly compressed in some specimens; interior with short, thin, straight dental plates; muscle field deeply incised. Dorsal valve evenly convex, with slightly flattened dorsum anteriorly; umbonal region slightly swollen, weakly compressed laterally; interior with divided hingeplate; median septum lacking; crura falcifer, not rising but remaining in plane of lateral commissure. Comments. — =These small specimens, lacking any indication of a fold and sul- cus, are probably juveniles of the genus Hemileurus Cooper and Grant, agreeing Table 12. — Measurements (in millimeters) o/?Elassonia sverdrupensis, n. sp. from GSC locality 56430 of the lower Hare Fiord Formation. GSC number Length Width Thickness 115603 10.5 9.5 5.3 115604 10.1 9.8 5.0 115605 9.5 9.9 4.4 146 Annals of Carnegie Museum VOL. 67 Fig. 14. — Transverse serial sections of IHemileurus sp., GSC 115615, X 8. Numbers refer to distance in millimeters from ventral beak. in most respects with the type species of that genus, except for their lack of a fold and sulcus and having more numerous costellae near the margins. They are not similar to any other species from the Hare Fiord Formation. Distribution. — GSC locality 56430 (five specimens, one sectioned). Family Tetracameridae Likharev, 1958 Genus Septacamera Stepanov, 1937 Septacamera sp. (Fig. 15.1-15.4) Description. — This description is based on a single, nearly complete specimen. Medium size for genus, strongly inequivalved; outline and lateral profile rounded subtrigonal, an- terior profile subquadrate; anterior surface flattened; anterior commissure uniplicate with large ventral tongue; fold and sulcus moderately well developed, originating in umbonal region of both valves; plicae simple, subangular, numbering three in sulcus, four on fold, and five or six on each lateral slope, lateral two or three very weak. Ventral valve gently convex with sharply geniculated tongue in sulcus; beak and foramen not preserved. Dorsal valve thick, strongly inflated with weakly convex dorsum; lateral slopes bent sharply ventrad with flattened sides. Interior not observed. Comments. — This species is similar in size, lateral profile, and ornament to Septacamera kutorgae (Chernyshev, 1902) from the Lower Permian of Russia. It differs in having a more triangular outline, a more quadrate anterior profile, and a flatter fold. Distribution. — -GSC locality 60194 (one specimen). Family Pontisiidae Cooper and Grant, 1976 Genus Antronaria Cooper and Grant, 1976 Antronaria annosa, new species (Fig. 15.5-15.19, 16) Holotype. —Figures 15.5-15.9, GSC 115617, from GSC locality 56430. Paratypes.— Figures 15.10-15.14, GSC 115618, from GSC locality 56430; Figures 15.15-15.19 and 16, GSC 115619, from GSC locality 60194. 1998 Carter and Poletaev — Upper Carboniferous Arctic Brachiopods 147 Fig. 15. — Rhynchonelloids. 15.1-15.4, Septacamera sp., ventral, dorsal, anterior, and lateral views, CSC 115616, X 1. 15.5—15.19, Antronaria annosa n. sp., ventral, dorsal, anterior, posterior, and lateral views of three specimens, including the holotype (15.5-15.9), GSC 115617-115619, respectively, X 1. 15.20-15.24, IPhrenophoria sp., ventral, dorsal, anterior, posterior, and lateral views, GSC 115620, X 1. Description. — Larger than average for genus, unequally biconvex; outline subelliptical; umbonal region very broad, subtending an angle of 135 degrees or more; both valve margins near posterior commissure compressed; fold and sulcus wide, well developed, originating in posterior half or third of valves; anterior commissure strongly uniplicate; beak short, inconspicuous, scarcely extending pos- terior to opposite valve; deltidial plates not observed; umbonal regions smooth, remainder of valves costate; flanks with five to seven subangular costae, originating well anterior to umbonal region; fold and sulcus with five to seven subangular costae. Ventral valve weakly to moderately convex or flattened in lateral profile, flat or concave in anterior profile; umbonal region slightly swollen, flanks flattened or weakly concave; sulcus originating pos- terior to midvalve, in anterior portion of umbonal region; sulcus rounded, evenly curving dorsad, not geniculate; dental plates strong, diverging anteriorly, of moderate length. Dorsal valve rounded subtrigonal in lateral profile, strongly convex in anterior profile; umbonal region laterally defined by concave flexures; flanks convex, curving steeply to lateral margins; fold originating near midlength, well defined and elevated anteriorly, flattened or weakly convex in anterior profile; costae on fold not depressed as in type species; interior with flattened, complete hingeplate; 148 Annals of Carnegie Museum VOL. 67 Fig. 16. — Transverse serial sections of Antronaria annosa n. sp., paratype GSC 115619, X 2.5. Num- bers refer to distance in millimeters from ventral beak. crural bases attached to dorsal surface of hingeplate, slightly convergent, extending forward in plane of commissure as long, slender, medially concave crura. Measurements. — See Table 13. Diagnosis.- — This species is characterized by evenly rounded lateral margins, inconspicuous ventral beak, smooth umbonal regions, and lack of depressed costae on the fold. Comments. — These specimens are similar to those illustrated by Cooper and Grant (1976) of Antronaria transversa (King) from the Wolfcampian of Texas. They differ in minor details, having more evenly rounded lateral margins, a small- er ventral beak, the ribbing originates more anteriorly, and they lack depressed medial ribs on the fold. This Arctic species is the first reported in the literature from strata older than the early Permian. Table 13. — Measurements (in millimeters) o/ Antronaria annosa n. sp. (King). GSC number Locality Length Width Thickness 115617 56430 16.2 23.5 14.9 115618 56430 14.4 20.6 13.2 115619 60194 15.5 20.8 13.7 1998 Carter and Poletaev — Upper Carboniferous Arctic Brachiopods 149 17 18 19 20 Fig. 17. — Athyridoids, 17.1-17.8, Camarium nuperum n. sp., ventral, dorsal, anterior, and lateral views of two specimens, including the holotype (17.5-17.8), GSC 115621 and 115622, X 3. 17.9-17.20, Nucleospira aquilonaris n. sp., ventral, dorsal, anterior, and lateral views of three specimens, including the holotype (17.9-17.12), GSC 115623-115625, respectively, X 2. Distribution. — -GSC locality 56430 (four specimens); GSC locality 60194 (two specimens, one sectioned). Order Athyridida Boucot, Johnson, and Staton, 1965 Suborder Athyridina Boucot, Johnson, and Staton, 1965 Superfamily Athyridoidea Davidson, 1881 Family Meristellidae Waagen, 1883 Subfamily Meristinae Hall and Clarke, 1895 Genus Camarium Hall, 1859 Camarium nuperum, new species (Fig. 17.1-17.8, 18) Holotype. — Figures 17.5—17.8, GSC 115621, from GSC locality 56430. Paratypes. — Figures 17.1-17.4, GSC 115622, from GSC locality 564530; Figure 18, GSC 115626, from GSC locality 56430. Description. — Small, subequally biconvex, outline subovate; maximum width attained near mid- length; lateral and anterior profiles lenticular; fold lacking but shallow sulcus present on anterior half 150 Annals of Carnegie Museum VOL. 67 Fig. 18. — Transverse serial sections of Camarium miperum n. sp., GSC 115626, X 6. Numbers refer to distance in millimeters from ventral beak. of ventral valve producing slight emargination; posterolateral margins slightly compressed; anterior commissure nearly rectimarginate with slight medial dorsal flexure; surfaces smooth except for fine growth lines. Ventral valve with slightly swollen umbonal region with moderately broad posterolateral extremities which subtend an angle greater than 90 degrees; beak small, incurved; foramen not observed; venter and lateral slopes evenly and moderately convex; sulcus originating near midlength, becoming mod- erately deep and rounded near anterior margin; interior with medially concave dental plates and low, short, convex shoelifter process which originates in umbonal region; teeth broad, blunt. Dorsal valve with sides of umbonal region slightly compressed, flanks and dorsum evenly convex near midvalve, dorsum slightly flattened anteriorly; interior with septalium supported by high median septum which extends forward nearly as far as shoelifter process in opposite valve; crura and spiralia not observed. Measurements, Sqq Table 14. Diagnosis. — This species can be differentiated by its small size and distinct ventral sulcus. Comments. — Amsden (1968) validated the genus Camarium Hall after discov- ering that it lacks mystrochial plates, unlike the genus Merista Suess which has them. The type species of Camarium, C. typa Hall, is from the Upper Silurian of the eastern United States. Carter (1967) described an anachronistic species of Camarium as Merista maccullochensis from the Lower Mississippian of Texas. Campbell and Engel (1963) also reported Merista sp. in the Toumaisian of New South Wales. All other reports of Camarium are from Middle Devonian or older strata. Camarium nuperum n. sp. is similar to the above-mentioned species of Tour- naisian age, differing only in the presence of a distinct ventral sulcus. Distribution. — GSC locality 56430 (four specimens, one sectioned). Table 14, — Measurements (in millimeters) of the types o/Camarium nuperum, n. sp.,from GSC locality 56430. GSC number Length Width Thickness 115621 9.4+ 9.0 6.1 115622 8.2 8.1 4.9 1998 Carter and Poletaev — Upper Carboniferous Arctic Brachiopods 151 o O * « 0.5 0.6 0.7 0.8 0.9 Fig. 19. — Transverse serial sections of Nucleospira aquilonaris n. sp., GSC 115627, X 5. Numbers refer to distance in millimeters from ventral beak. Family Nucleospiridae Davidson, 1881 Genus Nucleospira Hall, 1859 Nucleospira aquilonaris, new species (Fig. 17.9^17.20, 19) Holotype, —Vigums 17.9-17.12, GSC 115623, from GSC locality 56430. Paratypes. — Figures 17.13--17.20, two shells, GSC 115624 and 115625, from GSC locality 56430; Figure 19, GSC 115627, from GSC locality 56430. Description. — Small, subequally and moderately biconvex; outline subovate to weakly subpenta- gonal; greatest width near midlength; anterior margin rounded to slightly emarginate; lateral and anterior profiles lenticular; surfaces smooth except for irregularly spaced growth varices; micro-or- nament not preserved; fold lacking; sulcus lacking or weak, shallow sulcus present on some ventral valves; anterior commissure rectimarginate to weakly uniplicate; shell substance very thick. Ventral valve moderately inflated, evenly convex in lateral profile; umbonal region broad, slightly compressed at margins; beak small, acute, straight, projecting slightly posterior to opposite valve; foramen triangular; interarea acutely triangular, flattened to weakly concave; sulcus, if present, shallow, weak, originating in anterior half of valve; interior with large, blunt, unsupported teeth and low, weak median ridge. Dorsal valve with slightly swollen umbonal region defined laterally by concave flexures and com- pressed lateral margins; dorsum evenly convex or slightly flattened anteriorly; dorsal interarea not detected; interior with high bilobed hingeplate and thick myophragm; other details not observed. Measurements.— -See Table 15. Diagnosis. — ^This species is characterized by its thickened shell, well-developed ventral beak, flattened dorsum, and moderately inflated lateral profile. Comments.— Upper Carboniferous Nucleospira are very rare. Nucleospira su- perata Easton, 1962, from the Cameron Creek Formation (Lower Morrowan) of Montana, is the only other Upper Carboniferous species we can find in the lit- erature and is of very early Late Carboniferous age, much older than the Hare Fiord species. The latter is more transverse than N. aquilonaris, with an incon- spicuous ventral beak, and the lateral slopes are more flattened in anterior profile. Brunton (1984) also failed to find other evidence of Upper Carboniferous spe- cies of this genus. Cooper and Grant (1976) state that it generally had been Table 15. — Measurements (in millimeters) o/ Nucleospira aquilonaris, n. sp., from GSC locality 56430. GSC number Length Width Thickness 115623 9.7 9.9 6.6 115624 9.6 8.7 6.3 115625 8.5 8.6 5.8 152 Annals of Carnegie Museum VOL. 67 believed that Nucleospira became extinct at the end of the Early Carboniferous but they described a typical species, N. cunctata, from the Lower Permian of west Texas. The reduced ventral median ridge found in N. aquilonaris is also seen in this Permian species. Distribution. — GSC locality 56430 (40 specimens, two sectioned). Order Spiriferida Waagen, 1883 Suborder Spiriferidina Waagen, 1883 Superfamily Ambocoelioidea George, 1931 Family Ambocoeliidae George, 1931 Genus Crurithyris George, 1931 Crurithyris cf. tschernyschewi Likharev, 1939 (Fig. 20.18-20.21) 1902 Amhocoelia planoconvexa Shumard: Chernyshev, p. 196, pi. 20, fig. 1; pi. 49, fig. 7. 1939 Amhocoelia (Crurithyris) tschernyschewi Likharev, p. 108, pi. 27, fig. 5. Description. — Hingeline somewhat shorter than width; ventral valve moderately convex; greatest thickness at or just posterior to hingeline; ventral umbo narrow and weakly incurved, beak not over- hanging hinge plane; area low and concave under beak, clearly restricted, triangular; ventral sinus absent but valve flattened along midline; dorsal valve almost flat or slightly convex with shallow groove anteriorly; area low, triangular, well delimited; surface smooth, with few irregular coarse growth varices anteriorly; dental adminicula in ventral valve absent; other internal features not ob- served. Diagnosis. — A small, transversely ovate Crurithyris with greatest width no more than ten mm. Comments. — The general shape, size, and characters of the ventral valve allow us to conclude that this species is very close to or identical with Crurithyris tschernyschewi Likharev. However, our restricted material, consisting of only one complete shell and one ventral valve, are both poorly preserved and compel us to avoid a certain definition. The other nearest known species to ours is Cruri- thyris expansa Dunbar and Condra, 1932, from the Wabaunsee Group (Permian) in Nebraska. This species differs in its much more incurved ventral umbo with the apex lying approximately in the plane of the dorsal valve. Crurithyris plan- oconvexa (Shumard, 1855), from the Pennsylvanian and Lower Permian of North America and Europe, differs in its much more convex ventral valve with more stout, incurved ventral umbo and much smaller size. Distribution. — Both specimens are from GSC locality 56430. Crurithyris sp, (Fig. 20.22-20.25) Comments.— A second species of Crurithyris was found at GSC locality 56430. This one is characterized by its small size, more rounded outline, more tumid ventral valve, and more flattened dorsal valve than the previously described spe- Fig. 20. — Ambocoelioids and martinioids. 20.1-20.17, Tiramnia walteri n. sp.; 20.1-20.10, 20.13- 20.17, ventral, dorsal, lateral, anterior, and posterior views of three specimens, including the holotype (20.1-20.5), GSC 1 15628-1 15630, respectively; 20.1 1, 20.12, ventral molds showing vascular marks, GSC 115631 and 115632; all X 1. 20.18-20.21, Crurithyris cf. C. tschernyschewi Likharev, 1939, ventral, dorsal, anterior, and lateral views of a spalled shell, GSC 115633, X 3. 20.22-20.25, Cruri- thyris sp., ventral, dorsal, anterior, and lateral views, GSC 115634, X 3. 1998 Carter and Poletaev— Upper Carboniferous Arctic Brachiopods 153 154 Annals of Carnegie Museum VOL. 67 cies. Precise identification of this species is impossible because of the paucity of specimens, only two shells, one with a broken ventral beak. Superfamily Martinioidea Waagen, 1883 Family Martiniidae Waagen, 1883 Subfamily Martiniinae Waagen, 1883 Genus Tir amnia Grunt, 1977 Tiramnia walteri, new species (Fig. 20.1-20.17) Holotype. ~GSC 115628, Figures 20.1-20.5, GSC locality 56430. Paratypes. — GSC 115629 and 115630, Figures 20.6-20.10, 20.13-20.17, same collection as the holotype; GSC 115631 and 115632, Figures 20.11 and 20.12, vascular impressions of two ventral valves; all from GSC locality 56430. Description. — Medium to large for genus, ventribiconvex, usually wider than long; outline trans- versely oval to subpentagonal, greatest width attained near or anterior to midlength; ventral valve one and one-half times thicker than dorsal valve; anterior commissure uniplicate; fold and sulcus wide, shallow, and poorly delimited, developed only anteriorly; ornamentation lacking and surface smooth except for growth lines and irregularly spaced growth varices; spalled surfaces may show faint radial striations in front part of some adult specimens. Ventral valve most convex in umbonal region; beak small, incurved; umbonal region moderately broad, only moderately extended posterior to hingeline; flanks gently convex; sulcus originates ob- scurely some distance in front of beak as narrow, weak furrow becoming wider and somewhat deeper past midway to front margin, where in some shells its sides slope gently toward faint depression along midline; some variation in shape of sulcus; often furrow is almost obscure and sulcus depression is very softly curved, producing moderate tongue at anterior commissure; hingeline usually little less than half width of shell; ventral interarea small, triangular, moderately high and curved, often obscurely defined, with apical angle of about 115 to 120 degrees; delthyrium open. Ventral muscle field narrow, fusiform, deeply impressed, located mostly within umbonal region; posterior part of adult valves thickened; vascular impressions distinct, radial, bifurcating anteriorly. Dorsal valve transversely subovate in outline, slightly thinner than opposite one, with moderately gibbous umbonal region and inconspicuous beak; fold arises obscurely in anterior part of valve and rises to its maximum height at front margin, where it is obscurely delimited; dorsal interarea orthocline, less than one-third height of ventral, with broad, open notothyrium; muscle scars very simple, radially dissected and lightly impressed; other internal details not observed. Measurements.— S&c Table 16. Diagnosis.— This species is characterized by comparatively large size (as much as 40 mm long and 50 mm wide), transversely subovate to subpentagonal outline, longer thicker ventral valve than dorsal, and fold and sulcus only clearly expressed anteriorly. Comments. — Tiramnia walteri n. sp. is similar in general outline to Martinial triquetra Gemmellaro, 1899, sensu Grunt and Dmitriev, 1973, from the Nizhne- ganskaya Subsuite, Murgabian Stage, lower part of the Upper Permian of the southeastern Pamir Mountains. The latter differs from Tiramnia walteri n. sp. in having a much longer hingeline and a deeper, more distinctly acute sulcus. Tiramnia greenlandica Dunbar, 1955, from the Permian Martiniakalk of east Table 16. — Measurements (in millimeters) o/ Tiramnia walteri, n. sp. from GSC locality 56430. GSC number Length Width Thickness Hingeline 115628 39.5 43.7 26.5 17.1 115629 35.4 37.5 23.8 23.2 115630 24.7 26.8 14.7 10.4 1998 Carter and Poletaev — Upper Carboniferous Arctic Brachiopods 155 Greenland, is very similar to young specimens of this new species. Mature forms of T. walteri n. sp. differ in being much larger, they mostly have a less stout umbo and a shorter and less curved beak; the sulcus is more obscure posteriorly and lacks angular borders anteriorly. Distribution.— Tir amnia walteri n. sp. is abundant at GSC localities 56430 (37 specimens) and C-4011 (22 specimens). It also occurs at GSC localities 56430A (seven specimens) and C-4084 (seven specimens). A similar or identical species occurs in the Ladrones Limestone of southeastern Alaska. Tiramnia grunti, new species (Fig. 21.1-~21.16) V. 1977 Tiramnia semiglobosa Chernyschev, 1902:Gmnt, p. 66, pi. 11, fig. 8, 9. V.1977 Tiramnia greenlandica Dunbar, 1955:Grunt, p. 64, pi. 11, fig. 4, 6, non 5. Holotype.- — GSC 115635, Figures 21.1-21.5, from GSC locality 56430. Paratypes. —GSC 115636-115638, Figures 21.6-21.16, all from GSC locality 56430. Description. — Small to medium size for genus, moderately to strongly subequally biconvex; outline longitudinally subovate to almost round with dorsal valve often almost same length as opposite valve; hingeline much less than greatest width, often one-third or less; maximum width attained near mid- length; fold and sulcus obscure posteriorly and weak or moderately developed anteriorly, where they produce weakly or clearly uniplicate commissure; shell substance very thin; surface almost smooth, rarely with a few visible growth varices near anterior commissure. Ventral valve moderately to considerably inflated, rounded in outline, with greatest thickness near midlength; umbo short and suppressed; small beak, not extending posteriorly to hingeline; interarea low, catacline, concave, poorly differentiated from flanks; delthyrium wide, forming equilateral triangle and occupying approximately one-half of interarea; deltidial plates narrow; sulcus obscure, shallow, appearing near midlength; some specimens with narrow furrow along midline, which begins on umbo and extends to end of moderately long tongue; ventral interior with narrow, deeply impressed fusiform muscle field surrounded by radial vascular impressions; most prominent vascular impression lies on midline and continues to midlength. Dorsal valve usually as inflated as opposite valve; outline rounded or subpentagonal with large, slightly tumid umbonal region and small beak protruding posteriorly; dorsal interarea low, narrowly triangular, orthocline with open nothothyrium; fold absent or weakly expressed near anterior margin; interior with delicate cardinalia; muscle scars weakly impressed. Measurements. — See Table 17. Diagnosis. — This species is characterized by its comparatively small size, rounded outline, almost equidimensional ventral and dorsal valves, prominent dorsal umbo, very weak fold, and moderate or weak sulcus. Comments. — Tiramnia grunti n. sp. varies in such important characteristics as relative length of the dorsal and ventral valves, depth of sulcus, and thickness. Some specimens show extreme variation in one or more of these characters but are united by a multitude of intermediate forms. Tiramnia grunti n. sp. is most similar to Martinial corculum Kutorga, 1842, from the early Permian (Asselian?) near Sterlitamak, Bashkiria, south Urals. The latter differs in being smaller and having a flatter and shorter dorsal valve with an obscure umbo. Tiramnia greenlandica (Dunbar, 1955), from the Permian Mar- tiniakalk of east Greenland, differs also in its wider, more subtriangular outline and wider sulcus with rather angular borders. Distribution.— This is the most numerous species at GSC locality 56430 (144 good specimens plus several hundred disarticulated valves). Six specimens were found at GSC locality 56430A. It is known also, but not described, from the Upper Carboniferous of Cape Chaika, Ugor Peninsula, northern Ural Mountains, Russia, and the Ladrones Limestone of southeastern Alaska. 156 Annals of Carnegie Museum VOL. 67 Fig. 21. — Martinioids. 21.1-21.16, Tiramnia grunti n. sp.; 21.1-21.15, ventral, dorsal, anterior, pos- terior, and lateral views of three specimens, including the holotype (21.1-21.5), GSC 115635-115637, respectively; 21.16, mold of ventral interior showing vascular marks, GSC 1 15638, X 1. 21.17-21.32, IHeteraria canadiensis n. sp.; 21.17, micro-ornament, GSC 115639, X 10; 21.18-21.32, ventral, dor- sal, lateral, anterior, and posterior views of three specimens, including the holotype (21.18-21.22), GSC 115640-115642, respectively, X 1.5. 1998 Carter and Poletaev — Upper Carbonieerous Arctic Brachiopods 157 Table 17. — Measurements (in millimeters) o/Tiramnia grunti, n. sp. from GSC locality 56430. GSC number Length Width Thickness Hingeline 115635 22.4 22.7 16.6 7.9 115636 20.1 19.9 14.9 8.6 115637 16.7 13.9 12.2 5.8 Genus Jilinmartinia Lee and Gu, 1980 IJilinmartinia cf. J. sokolovi (Tschernyschew, 1902) (Fig. 23.1-23.4) V. 1902 Spirifer sokolovi Chernyshev, p. 166, pi. 8, fig. 3, pi. 39, fig. 4. 1939 Spirifer (Brachythyris) sokolovi Likharev, p. 107, pi. 27, fig. 3. 1980 Martinia sokolovi Kalashnikov, p. 100, pi. 36, fig. 2, 3; textfig. 10. Discussion.— T\iqx& is only one specimen of this species from GSC locality 56430. It is a large, completely smooth martiniid closely similar to the lectotype of Spirifer sokolovi Chernyshev (1902:pl. 8, fig. 3). The Canadian specimen has almost the same size, general shape, outline, regularly curved ventral valve, and nearly flat dorsal valve as seen in the Uralian specimen from the Schwagerina beds (Asselian) of Kazarmensky Kamen Hill near the town of Asha on the Sim River, Ural Mountains. Comments. — Assignment of this species to the genus Jilinmartinia is uncertain because it is based only on external similarity with the type specimen of the type species. The internal morphology of the Chinese and Canadian specimens is un- known. Kalashnikov (1980) reported Martinia sokolovi (Chernyshev) from the Lower Moscovian (Vereyan) and Upper Carboniferous beds of Novaya Zemlya and the Moscovian of north Timan. The specimen illustrated by Kalashnikov is more similar to specimens of Jilinmartinia laevis (Likharev, 1939) from the lower Gje- lian of the Donets Basin. Subfamily Eomartiniopsinae Carter, 1994 Genus Heteraria Cooper and Grant, 1976 Comments. — Cooper and Grant (1976) did not note the similarity of their new genus Heteraria to the Lower Carboniferous genera Eomartiniopsis Sokolskaya, 1941, or Merospirifer Reed, 1949. In the case of the former this was undoubtedly because of their misunderstanding of the dorsal interior of the type species of Eomartiniopsis, which was described by Sokolskaya as having short or incipient “septal” plates. In fact, there are no dorsal plates supporting the cardinalia in Eomartiniopsis . Thus, the dorsal interiors of Eomartiniopsis, Merospirifer, and Heteraria are similar. Merospirifer Reed, 1949, was recently redescribed by Brun- ton (1984) and appears to be similar to Eomartiniopsis in the dorsal interior. As far as we can determine, these genera can be internally differentiated only by the presence of unusually wide, transverse, flattened dental ridges in the ventral valve of Heteraria with narrow, more conventional ridges in Eomartiniopsis and Mer- ospirifer. There are only three Late Carboniferous eomartiniopsinine species known, Eomartiniopsis planosinuata Poletaev, 1975, from the early Bashkirian of the Donets Basin, E.l susanae Martinez-Chacon, 1978, from the middle to late Bash- 158 Annals of Carnegie Museum VOL. 67 dD dD (H) (Jb 0.2 O 0.4 C) Ip) 0.6 0.8 n n -1 A X a (K) O.y 1,0 OD 1.1 (Jh 1.i j 5 1.4 XV 1.3 1.2 Fig. 22. — Transverse serial sections of IHeteraria canadiensis n. sp., paratype, GSC 115643, from CSC locality 56430, X 3. Numbers refer to distance in millimeters from ventral beak. kirian of the Cantabrian Mountains, northern Spain, and the new species described below from the upper Bashkirian or lower Moscovian of Ellesmere Island. Their similarities and differences are discussed below. IHeteraria canadiensis, new species (Fig. 21.17-21.32, 22) Holotype.— GSC 115640, Figures 21.18-21.22, from GSC locality 56430. Paratypes. — GSC 115641, Figures 21.23-21.27, from GSC locality C-5202; GSC 115642, Figures 21.28-21.32, from GSC locality 56430; GSC 115639, Figure 21.17, micro-ornament, from GSC lo- cality 56430A; Figure 22, GSC 115643. Description. — Small to medium size for genus, smooth, thin shelled; transversely subovate or sub- pentagonal in outline; subequally biconvex, both valves considerably inflated; beaks small; hingeline narrow, half of maximum width; maximum width attained near midlength or slightly posterior to midlength; cardinal extremities rounded in all growth stages, but slightly angular in juveniles and delineated by obscure reflexing of outer surface of ventral valve; ornament consisting of irregularly spaced growth varices and some adult specimens have faint low costae or plicae on flanks; micro- ornament consisting of very small pits. Ventral valve most convex in umbonal region; beak small and slightly incurved; sulcus originating as shallow groove posteriorly in umbonal region, becoming broader and deeper anteriorly, with angular borders and flattened floor; front margin strongly uniplicate with angular tongue; flanks gently convex or nearly flat; interarea moderately high, sharply defined, triangular; dental ridges (sensu Dunlop, 1962) very wide, flattened; deltidial or stegidial plates not observed; interior with short, slender, subparallel intrasinal dental adminicula. Dorsal valve similar in convexity to ventral valve and almost as long; prominent umbo has small beak; fold poorly defined posteriorly, forming only low, rounded ridge, becoming well expressed only anteriorly; flanks slightly concave near the ears and gently convex otherwise; dorsal interarea low, acutely triangular, inconspicuous; interior with small ctenophoridium and narrow, vertical, medially concave crural bases which become broader and convergent anteriorly; other details not preserved in sectioned specimen. Measurements. — See Table 18. Diagnosis.- — This species is characterized by its transversely subovate to sub- quadrate outline, almost equal thickness of the ventral and dorsal valves, small 1998 Carter and Poletaev — Upper Carboniferous Arctic Brachiopods 159 Table 18. — Measurements (in millimeters) o/Heteraria canadiensis, n. sp. GSC number Length Width Thickness Hingeline 115640 >17.0 -21.0 14.5 -10.5 115641 13.6 16.2 9.4 -1.5 115642 11. 1 12.3 8.1 6.2 ventral umbo with a short, narrow beak, deep flat-bottomed sulcus with anteriorly abrupt slopes, and very wide, flattened dental ridges. Comments. — Heteraria canadiensis n. sp. resembles Eomartiniopsis planosin- uata Poletaev, 1975, from the Lower Bashkirian of the Donets Basin. The Ukrain- ian species differs in its smaller size, very unequal convexity of ventral and dorsal valves, comparatively longer ventral umbo, higher interarea, and shallower sulcus than the Canadian species. Eomartiniopsis susanae Martinez-Chacon, 1978, from the San Emiliano For- mation (Bashkirian) of northern Spain, differs from H. canadiensis in having a roundly subquadrate outline with its greatest width at midlength or anterior to it, and a narrower sulcus with distinct grooves on the midline. Distribution. — GSC locality 56430, four complete shells, two ventral valves, and one dorsal valve. GSC locality C-5202, two complete shells. GSC locality C-4085, one ventral valve. GSC locality 56430A, a well-preserved large ventral valve. Indeterminate martinioid (Fig. 23.5-23.9) Description. — Small, strongly and unequally biconvex with much thicker dorsal valve; outline sub- elliptical to rounded subpentagonal; lateral profile subelliptical, almost subovate; anterior profile thick- ly lenticular; sulcus well developed but fold scarcely discernible; anterior commissure strongly uni- plicate; surfaces smooth except for very faint, widely spaced radial lirae and fine growth lines, the former possibly reflecting internal vascular markings. Ventral valve shorter and much thinner than dorsal valve; umbonal region moderately convex, defined by strongly concave lateral flexures, forming compressed posterolateral margins; beak very small, incurved; delthyrium open, small, triangular; interarea small, narrow, procline, triangular, weakly delineated by scarcely perceptible beak ridges; lateral slopes gently concave, sloping evenly to lateral margins; sulcus originating in umbonal region, becoming smoothly rounded and moderately deep and wide anteriorly; interior unknown. Dorsal valve very thick, strongly inflated, with moderately swollen umbonal region delineated by concave flexures that are less concave than those of opposite valve; dorsum strongly convex, lateral slopes more gently convex, sloping evenly to lateral margins; fold confined to very slight dorsal deflection of shell surface near anterior margin; interarea very short and narrow, about orthocline; interior unknown. Comments. — We little doubt that this shell represents an adult individual be- cause of the well-developed sulcus and rotund shape. The translucent shell ma- terial provides no suggestion of internal structures in either valve. Growth form, lack of ornamentation, and probable lack of internal plates indicate that this small shell is undoubtedly a martinioid of some kind. However, the short ventral valve with procline ventral interarea, greatly swollen dorsal valve, and unusual micro- ornament preclude assignment to a known species, genus, or even family. Distribution. — -GSC locality 56430 (one specimen). 160 Annals of Carnegie Museum VOL. 67 Fig. 23. — Martinioids. 23.1-23.4, IJilimnartinia cf. J. sokolovi (Chernyshev, 1902), ventral, dorsal, posterior, and lateral views, GSC 115644, X 1. 23.5-23.9, indeterminate martinioid, ventral, dorsal, anterior, posterior, and lateral views, GSC 1 15645, X 2. Superfamily Spiriferoidea King, 1846 Family Spiriferidae King, 1846 Subfamily Sergospiriferinae Carter, 1994 Genus Anthracospirifer Lane, 1963 Anthracospirifer cf. A. occiduus Sadlick, 1960 (Fig. 24.9-24.13) I960 Spirifer occiduus Sadlick, p. 1210. Description. — This description is based on two specimens from GSC locality 56430. An almost complete shell with incomplete cardinal extremities measures approximately 37 to 38 mm in width, 22.7 mm in length, and 15.8 mm in thickness. The remaining specimen is a small ventral valve. Fig. 24. — Spiriferoids. 24.1-24.8, Elinoria ellesmerensis n. sp.; 24.1-24.3, ventral, anterior, and pos- terior views of a large ventral valve, GSC 115646; 24.4—24.8, ventral, dorsal, anterior, posterior, and lateral views of the holotype, GSC 1 15647, X 1. 24.9-24.13, Anthracospirifer cf. A. occiduus Sadlick, 1960, anterior, lateral, ventral, dorsal, and posterior views, GSC 11^648, X 1; 24. 14-24. f6, IBrach- ythyrina sp., ventral, anterior, and posterior views oT a ventral valve, GSC 115649, X 1. 162 Annals of Carnegie Museum VOL. 67 Ventral valve with short, narrow ventral umbo, moderately high, triangular interarea, deep V-shaped sulcus with obvious median rib and three ribs on each side of sulcus, up to 14 or 15 ribs on each flank, four or five of them bifurcated. Dorsal valve with short, strongly incurved umbo, narrow acute interarea, prominent fold with groove on midline, and four or five bifurcated ribs on each flank. Remarks. — This shell is similar in general shape and ribbing to the lectotype of Anthracospirifer occiduus (Sadlick), as illustrated by Girty (1927) and Gordon (1975). The shell from Ellesmere Island differs from the lectotype in having a higher ventral interarea and concave lateral slopes on the ventral valve and gently convex flanks on the dorsal valve. Family Choristitidae Waterhouse, 1968 Subfamily Angiospiriferinae Legrand-Blain, 1985 Genus Elinoria Cooper and Muir- Wood, 1951 Elinoria ellesmerensis, new species (Fig. 24.1-24.8) Holotype. — GSC 115647, Figures 24.4-24.8, from GSC locality 56430. Paratypes. — GSC 115646, Figures 24.1-24.3, a large ventral valve from GSC locality 60194. Description. — Medium to large, subequally biconvex; outline transversely subovate to subsemicir- cular in early growth stages, becoming subtrapezoidal in adults; cardinal extremities alate to submu- cronate; sulcus moderately wide and deep, well defined; fold low, rounded, sharply delineated; anterior commissure uniplicate; flanks with few, simple, low, rounded costae, separated by narrow interspaces, and strong, irregularly spaced growth varices; width of costae varies from 0.6 to 0.8 mm near umbo, up to 3.0 mm or more near anterior margin; micro-ornament consisting of fine concentric growth lines and capillae; shell substance of moderate thickness for genus, reaching 3.0 mm thick in ventral umbo. Ventral valve moderately to strongly inflated with greatest convexity near or posterior to midvalve; surface of valve regularly convex except for sulcus and slight flexure near ears; umbo comparatively short and umbonal angle usually about 1 10 degrees; interarea flat or slightly concave and of moderate height, vertically grooved, abruptly truncated laterally, apsacline to nearly catacline; delthyrium form- ing equilateral triangle, or slightly higher than wide, open; sulcus subtending angle of about 30 degrees; sulcus with strong median costa, one pair of strong lateral sulcal costae which also originate at beak, and two or three pairs of weak, flat costae bifurcating from sulcus-bounding costae; sulcus spreading anteriorly in some large specimens to incorporate one or two pairs of costae from lateral slopes; lateral slopes with about 15 simple, rounded costae at anterior margin of each flank in adult specimens. Ventral interior lacking dental plates, bearing deep, narrow, depressed musclescar with short partial myophragm. Dorsal valve less inflated than opposite valve, forming semicircular anterior profile; dorsal umbo blunt, beak small, incurved; interarea orthocline, low, with acute extremities; fold low and clearly delineated, usually bearing four costae, flatter than those of lateral slopes; lateral costae simple, ex- cluding those nearest fold; dorsal interior unknown. Measurements. — See Table 19. Diagnosis. — The large size, transversely subtrapezoidal outline, clearly defined sulcus, coarse simple costae, and absence of dental plates distinguish this species from other Hare Fiord spiriferids. Comparisons. — Elinoria ellesmerensis n, sp. is most similar to E. rectangula (Kutorga) sensu Chernyshev (1902), from the Lower Permian (Asselian) of the Table 19. — Measurements (in millimeters) o/ Elinoria ellesmerensis, n, sp. GSC number Length Width Thickness Hinge line 1 15646 45.7 69.2 20.5 64.1 115647 41.3 57.0 34.1 55.3 1998 Carter and Poletaev- — Upper Carboniferous Arctic Brachiopods 163 western slope of the middle Urals, The Canadian species differs from the Russian one in its thicker, more stout shell, narrower width, a higher interarea in adults, mucronate cardinal extremities in juveniles, and simple or very rarely bifurcating lateral costae. Two other similar Russian species are Elinoria kremenskensis (Se- mikhatova, 1953) from the Upper Moscovian and Kasimovian of the Don^Med- veditsa district, and E, subgrandis Poletaev, 1986, from the lower Gjelian of the Donets Basin and Samarskaya Luka on the middle Volga River. Elinoria kremen- skensis is much smaller, almost half the size, lacks mucronate cardinal extremities, has a poorly delineated sulcus, and often has bifurcated costae on the lateral slopes. Elinoria subgrandis differs in its less inflated profile, semicircular to sub- quadrate outline, and has bifurcating costae on the lateral slopes. Distribution.— GSC locality 56430 (four specimens, two complete, including the holotype); GSC locality C-4083 (three more or less complete small ventral valves); GSC locality 60194 (one large ventral valve paratype and one small complete shell). A similar or identical species occurs in the Ladrones Limestone of southeastern Alaska. Genus Brachythyrina Frederiks, 1929 IBrachythyrina sp. (Fig. 24.14=24.16) Description - — This description is based on a single ventral valve from GSC locality C-4087. Medium size for genus (length 19.8 mm, width 24.4 mm, thickness 10.7 mm), moderately inflated, most convex in umbonal region; outline transversely subtrapezoidal; anterior commissure uniplicate; greatest width at hingeline; lateral extremities subangular, slightly mucronate and compressed in adult and transversely subovate in juveniles; beak small, narrow, incurved; beak ridges inconspicuous on umbo, angular and well defined near ears; interarea acutely triangular, weakly to moderately concave, of moderate height, catacline, vertically grooved; hingeline denticulate; delthyrium slightly higher than wide, apparently open; sulcus strong, well defined, moderately wide and deep, rounded; median sulcal costa very weak, originating in umbonal region; two pairs of lateral sulcal costae originating by bifurcating from sulcus-bounding costae, remaining simple for entire length; macro-ornament consist- ing of eight to ten simple, low, flattened, rounded costae on each flank and rare irregular growth varices; micro-ornament consisting of fine, regularly spaced growth lines; capillae not observed; um- bonal region thickened with callus; inner structures not observed. Comments specimen was assigned to Brachythyrina because it has the external characters of that genus and because it is externally similar to the Middle- Upper Bashkirian species Spirifer {Brachythyrina) probus Rotai, 1951, from the Donets Basin and Brachythyrina pinica Martinez-Chacon, 1978, from the Can- tabrian Mountains, northern Spain. Subfamily Choristitinae Waterhouse, 1968 Genus Parachoristites Barchatova, 1970 Parachoristites tellevakensis, new species (Fig. 25.1-25.15) Holotype.— A nearly complete shell with incomplete cardinal extremities, GSC 115650, Figures 25.1=25.5, collected by Bonham-Carter at GSC locality 60194. Paratypes. — Two almost complete shells, GSC 115651 and 115652, Figures 25.6-25.15, from the same locality as the holotype. Description. — Medium to large; outline subtriangular, transversely subrhomboidal, or subsemicir- cular; hingeline equal to or only slightly shorter than greatest width in shells with moderately large and incurved ventral beak; greatest thickness located near midlength; ventral valve slightly thicker than dorsal; ventral interarea high or moderately high for genus, with distinct vertical grooves and truncated lateral extremities; sulcus weakly developed, shallow, narrow, poorly defined; anterior com- 164 Annals of Carnegie Museum VOL. 67 1998 Carter and Poletaev — Upper Carboniferous Arctic Brachiopods 165 Table 20. — Measurements (in millimeters) of the types o/Parachoristites tellevakensis, n. sp.from GSC locality 60194. GSC number Length Width Thickness Hingeline 115650 47.4 -70.0 31.8 — 115651 >40.3 54.5 27.1 51.6 115652 31.8 39.5 21.3 -34.8 missure usually clearly unipiicate; fold very low and rounded posteriorly but clearly defined and elevated anteriorly; macro-ornament consisting of numerous flattened, rounded costae, mostly irreg- ularly bifurcated or trifurcated, with 11 to 13 costae in anterior part of sulcus, including very weak median rib, and up to 20 costae on each lateral slope; costae separated by narrow interspaces and sometimes crossed by strong, irregularly spaced growth varices; micro-ornament consisting of fine capillae and fine regular growth lines. Ventral valve most convex in umbonal region; beak of small to moderate size, incurved, overhanging hingeline; umbonal region moderately broad with umbonal angle of about 95 to 100 degrees; flanks gently convex; lateral extremities slightly compressed on ears, forming sharp beak ridges; sulcus originating at beak, flat and shallow, becoming wider and deeper anteriorly, producing moderate tongue; interarea apsacline, clearly triangular, concave in juveniles, becoming flat with nearly parallel sides at maturity, and abruptly truncated extremities; deithyrium open, triangular, higher than wide, deltidial plates not observed; interior with thick, weakly diverging, moderately long dental admieicula which reach almost one-quarter length of valve; delthyrial plates conjoin in beak, becoming free anteriorly and including between them muscle impressions; posterior inner surface of the valve with traces of bifurcating vascular impressions. Dorsal valve usually semicircular in outline, moderately convex with short incurved umbo and low subparallel interarea; fold low, medially flattened, rounded, clearly defi.ned, with shallow midline groove and six to ten obscure costae at anterior margin; lateral slopes with 18 to 20 ribs on each side, most of which bifurcate; interior with ctenophoridium composed of numerous plates. Measurements. — -See Table 20. Diagnosis.- — This species is distinguished by its large size, comparatively high ventral interarea with sharp beak ridges in young stages; weak shallow sulcus with obscure borders; low, rounded, clearly defined fold; low, flat, irregularly bifurcating costae, and moderately long, thick, subparallel dental admiriicula. Comments. — -Parachoristites tellevakensis n. sp. bears similarity to the type species, P. voiongaensis Barkhatova, 1970, from the Bashkirian of the Volonga River and north Timan (Russia). The Canadian species differs in usually being smaller with a longer more subrhomboidal outline, a higher subtriangular ventral interarea, more elevated fold anteriorly, and flatter costae. Parachoristites tellevakensis n. sp. has some similarity to two other species of large spiriferids from the Televak Limestone Member, The species we refer to here as ITrautscholdia sp., from GSC locality C-56430, differs in its transversely subovate outline, absence of mucronate cardinal extremities, low subparallel ven- tral interarea, very weak sulcus with medial groove, very weak tongue, almost flat dorsal valve, and a fold defined only by deep interspaces. Elinoria ellesmerensis n. sp. differs in having a high subparallel ventral inter- area; a strong, clearly restricted sulcus; simple coarse costae; and no dental ad- minicula. Fig. 25. — Spiriferoids. 25.1-25.15, Parachoristites tellevakensis n. sp., ventral, dorsal, lateral, anterior, and posterior views of three specimens, including- the holotype (25.1-25.5), GSC 115650-115652, respectively, X 1. 166 Annals of Carnegie Museum VOL. 67 Distribution.— Thi^ species is moderately common at GSC locality 60194 (27 complete specimens) and GSC locality 56430 (more than 12 complete specimens, plus disarticulated valves). Genus Trautschoidia Ustritsky, 1967 ITrautscholdia sp. (Fig. 26.2=26.6) Description. — This description is based on a single large specimen (length 48.4 mm, width 66.2 mm, thickness 29.5 mm), from GSC locality 56430. Large, ventribiconvex, transversely subovate in outline with moderately thick ventral and weakly convex dorsal valves; cardinal extremities rounded on all growth stages; maximum width attained near midlength; ventral interarea low, acutely triangular, apsacline, concave; hingeline slightly shorter than greatest width, denticulate; sulcus very weak, narrow, poorly defined, medially grooved; lateral slopes with up to 30 wide, low, flattened, rounded costae at anterior margin, mostly simple or irregularly bifurcating or, rarely, trifurcating; growth varices irregularly spaced; micro-ornament consisting of strong capillae, about eight to ten per mm, and very fine, regularly spaced growth lines forming cancellate pattern; both valves much thickened posteriorly; ventral interior with nearly parallel dental adminicula. Comments.— This specimen is similar to Choristites jigulensis (Stuckenberg, 1905) sensu Ivanov and Ivanova, 1937 (pL 11, fig. 1 and pL 12, fig. 1) from the Upper Moscovian (Podolsky Horizon) of Shchurovo village, Moscow Basin. However, the Upper Moscovian specimens and the specimen from Ellesmere Is- land differ from the lectotype in the outline of their shells. The lectotype of Spirifer jigulensis Stuckenberg, 1905, is from the Lower Gjelian of the Samar^ skaya Luka (Volga River near the city of Samara). This lectotype has a subquad^ rate, not a transversely subovate, outline; a shorter hingeline; and a longer ventral umbo than either the Russian Upper Moscovian specimens or the specimen from Ellesmere Island. Trautschoidia jigulensis (Stuckenberg, 1905), in the junior author’s opinion, has many synonyms, including other species described by Stuckenberg from the type locality at Samarskaya Luka. We do not include Upper Moscovian specimens in the synonomy of T. jigulensis (Stuckenberg). It is possible that the specimens of Moscovian age represent a cognate but different, perhaps undescribed, species. Subfamily Tangshanellinae Carter, 1994 Genus Tangshanella Chao, 1929 ITangshanella sp. (Fig. 26.7=26.9) Description. — This description is based on a single incomplete specimen from GSC locality 56430. Medium size (length 38.3 mm, width 44.4 mm, thickness 23.5 mm), moderately inflated, ventribi- convex; outline transversely subovate in all growth stages; ventral umbo narrow, moderately inflated, and extended; cardinal extremities and lateral margins well rounded, hingeline very narrow (15.2 mm), about one-third maximum width; sulcus shallow, poorly defined, with weak groove in midline; fold low, poorly delineated, expressed only anteriorly, producing uniplicate anterior commissure; costae wide, low, roundly flattened, mostly bifurcating but almost disappearing at posterolateral margins; growth varices coarse, rare, irregular; micro-ornament not observed. Ventral valve twice as thick as dorsal valve, moderately and regularly convex with maximum width at midlength; interarea apsacline, moderately high, triangular; delthyrium open, wider than sides of interarea; hingeline very narrow, denticulation not observed; sulcus with three pairs of primary costae, weak median costa, which begins much later than primary costae; all primary costae usually bifurcate anteriorly; dental adminicula either very short or absent (beak lost). Dorsal valve subovate, very gently convex, with small, prominent umbo, low, almost linear, dorsal interarea, and wide notothyrium; interior not observed. 1998 Carter and Poletaev — Upper Carboniferous Arctic Brachiopods 167 Fig. 26. — Spiriferoids. 26.1, 26.10, Gypospirifer sp.; 26.1, a small ventral valve, GSC 115653; 26.10, a large dorsal valve, GSC 115656. 26.2-26.6, ITrautscholdia sp., ventral, dorsal, anterior, posterior, and lateral views, GSC 1 15654. 26.7-26.9, ITangshanella sp., ventral, dorsal, and anterior views, GSC 115655. All X 1. Diagnosis.— Th& transversely subovate outline, extremely narrow hingeline, wide, roundly flattened commonly bifurcating costae, shallow, poorly delimited sulcus and weak fold, and possible lack of dental adminicula serve to differentiate this species from all other spiriferids in the Televak Limestone Member. 168 Annals of Carnegie Museum VOL. 67 Comments. — This Canadian specimen differs from the other representatives of the genus Tangshanella Chao, 1929. The type species, T. kaipingensis Chao, from the Tangshan Limestone of the Penchi Series (Upper Carboniferous) of north China, differs in having a thicker shell with a much stronger sulcus and fold, a wider hingeline, and a higher ventral interarea. Tangshanella taimyrica (Einor, 1939), from the Makarovsky Horizon (Bashkirian) of Taimyr, and T. byrangi Cherniak, 1963 (in Ustritskii and Cherniak, 1963), from the Kholodninskaya Suite (Bashkirian) of Taimyr, have wider ventral interareas, and clearly fasciculate, strong, thin costae. They may not belong in this genus. This ITangshanella sp. from the lower Hare Fiord Formation is similar in outline, growth form, ribbing, and foldsulcus to Neomunella Ozaki, 1931, from the Pen-hsi-hsien (Upper Carboniferous) of Manchuria (north China). However, the latter has short, distinct, divergent dental plates. We assign our specimen to the genus Tangshanella with a query (?) because we cannot determine whether or not there were short dental adminicula in this incomplete specimen. Family Trigonotretidae Schuchert, 1893 Subfamily Neospiriferinae Waterhouse, 1968 Genus Gypospirifer Cooper and Grant, 1976 Gypospirifer sp. (Fig. 26.1, 26.10) Description. — This description is based on two specimens. A small ventral valve, perhaps a juvenile, and a large dorsal valve were collected at GSC locality 56430. Ventral valve moderately convex, strongly transverse with maximum width at hingeline; cardinal extremities slightly mucronate, slender (length 15.2 mm, width 43.3+ mm, complete width probably close to 50.0 mm, thickness —8.0 mm); commissure uniplicate; beak short, incurved, not swollen; sulcus shallow, well delineated by bounding costae and producing angle near 25 degrees; interarea triangular, comparatively high (approximately one-third valve length), slightly concave, strongly ver- tically grooved; delthyrium as high as wide, open or partly closed by stegidial plates; costae fine, bifurcated or trifurcated only in umbonal region, anteriorly simple, numbering ten per cm near anterior commissure; sulcus with median costa originating at beak; two pairs of costae, split from bounding costae inside sulcus, one pair split outside; growth lamellae distinct, regularly spaced, slightly raised to give tegulate appearance. Dorsal valve of average size (length 33.4 mm, width >67.2 mm, probably about 85 to 90 mm when complete); outline transversely semicircular with maximum width at hingeline; dorsum moderately convex; maximum thickness in umbonal region; flanks weakly convex with slightly compressed ears; cardinal extremities acute and mucronate in all growth stages; fold gently rounded, not raised much above flanks; costae medium size (eight per cm near anterior commissure), rounded, separated by rounded interspaces, 26 to 28 on flanks, mostly trifurcating from five to six primary ribs on umbo; ten costae on fold, bifurcating from median rib; growth varices numerous, regular; interior and micro- ornament not observed. Superfamily Paeckelmanelloidea Ivanova, 1972 Family Strophopleuridae Carter, 1974 Subfamily Strophopleurinae Carter, 1974 Genus Cantabriella Martinez-Chacon and Rio Garcia, 1987 This genus was assigned originally to the Licharewiidae by Martinez-Chacon and Rio Garcia (1987). However, the type species, Orulgania schulzi Martinez- Chacon, 1978, is impunctate according to Martinez-Chacon (1978). The pits de- scribed later by Martinez-Chacon and Rio Garcia (1987:pl. 1) appear to be an artifact of chemical corrosion. The type species of Cantabriella clearly has affin- ities with the paeckelmanellids, having such characters as disproportionately strong sulcus-bounding ribs, high ventral interarea, noncostate foldsulcus; strongly 1998 Carter and Poletaev — Upper Carboniferous Arctic Brachiopods 169 denticulate hingeline, close-set, intrasinal dental adminicula, and capillate micro- ornament. The authors of this genus included three species, all the from the Upper Car- boniferous of Spain: Cantabriella schulzi (Martinez-Chacon, 1978) from Upper Bashkirian of Leon and Asturias; C palentina (Martinez-Chacon, 1978) from Upper Moscovian of Palencia, and C. lavianica Martinez-Chacon, 1987, from Lower Moscovian of Asturias. In our opinion Spirifer (Syringothyris?) bistritzae Schellwien, 1900, from the Trogkofel shales (?Lower Permian) of the Carnic Alps of northern Slavonia also can to be assigned to this genus. Cantabriella sp. A (Fig. 27.6-27.10) Description. — This description is based on a single, nearly complete, possibly slightly deformed shell collected at GSC locality 56430. Medium size (length 13.6 mm, width >20.0 mm, thickness 11.1 mm), subequally biconvex, trans- versely subpentagonal in outline; lateral extremities acute and angular in all growth stages, becoming almost rectangular in adults; maximum width attained at hingeline; sulcus well developed and defined; anterior commissure uniplicate; ornament consisting of seven to eight simple, rounded, flattened costae on flanks; sulcus and fold smooth with weak, shallow furrow on midline; micro-ornament consisting of weak, irregularly spaced growth varices, very faint regularly and closely spaced growth lines, crossed by comparatively coarse capillae. Ventral valve subpyramidal with maximum thickness in umbo; beak small, acute, incurved; ventral interarea high, procline, concave only near beak, weakly defined by rounded beak ridges, grooved by strong vertical furrows; delthyrium moderately broad but higher than wide; deltidial plates not ob- served; sulcus originating at beak as narrow groove bordered by pair of wide costae, becoming wider and deeper anteriorly, producing flattened, long tongue in anterior third of valve; interior with dental adminicula. Dorsal valve transversely semicircular to subelliptical in outline, most convex in umbonal region, nearly as thick as ventral valve, reaching maximum thickness near midlength; umbo slightly swollen; beak small, inconspicuous; fold defined only by bounding interspaces posteriorly, becoming slightly raised anteriorly from midlength; internal details not observed. Diagnosis. — The subpentagonal outline, almost equal thickness of the subpyr- amidal ventral and swollen dorsal valves, procline ventral interarea, and simple costae rapidly decreasing to size toward the ears are the most characteristic fea- tures of this specimen. Cantabriella sp. B (Fig. 27.1-27.5) Description. — This description is based on a single, almost complete shell with broken cardinal extremities, spalled surface, but well-preserved growth form, from GSC locality C-4085. Medium size (length 15.7 mm, width >23.0 mm, thickness 10.6 mm); very unequally ventribicon- vex; transversely subtriangular in outline; lateral extremities angular in all growth stages; maximum width attained at hingeline; sulcus and fold well developed and defined; anterior commissure unipli- cate; ornament consisting of from nine to 1 1 rounded, flattened costae on flanks, broad near foldsulcus, becoming much finer laterally; bounding costae bifurcate on one side only; fold and sulcus smooth; sulcus with subangular floor; micro-ornament not preserved but consisting at least of more or less regularly spaced growth varices. Ventral valve subpyramidal, much thicker than dorsal valve, with maximum thickness in umbonal region; flanks concave, sloping evenly to anterolateral margins; ears compressed; umbonal region narrowly elongated; beak small, acute, slightly incurved; ventral interarea high, catacline or nearly procline, generally flat, weakly concave near beak, defined by sharp beak ridges; surface of interarea with strong vertical grooves; delthyrium narrow, almost three times higher than wide; subdelthyrial plate long, set well below surface of interarea between close-set intrasinal dental adminicula; stegidial plates not observed; sulcus comparatively shallow, originating as narrow groove, becoming wider and deeper anteriorly, defined by strong bounding costae. 1998 Carter and Poletaev-=-=Upper Carboniferous Arctic Brachiopods 171 Dorsal valve traosversely trapezoidal in outline, with maximum thickness at midlength, much thinner than opposite valve; umbo slightly produced above beak ridges; middle part of valve slightly convex or swollen from beak to front commissure; flanks concave, ears compressed; fold originating at small beak, well defined by bounding interspaces, slightly flattened medially near anterior margin; internal details not observed. Diagnosis.— This specimen probably also represents a new species of this ge- nus, differing from other species of Cantabriella in having a transversely trian- gular outline, concave flanks, elongated and compressed ears, and nine to 1 1 ribs on each flank. Superfamily Brachythyridoidea Frederiks, 1924 Family Brachythrididae Frederiks, 1924 Genus Meristorygma Carter, 1974 Meristorygma arctica Carter, 1974 (Fig. 27.11-27.14) v*1974 Meristorygma arctica Carter, p. 690, pi. 3, fig. 21“-30; text-fig. 4. Original Diagnosis.— ^ "This species is characterized by its transversely ovate outline, compressed lateral extremities, two or four sulcal costae, and dorsally deflected fold at the anterior margin,” Comments .—This species was originally described on the basis of eight spec- imens from GSC locality 56430. Our present collection contains nearly 100 spec- imens, many complete, from the same locality. Some natural molds of the ventral valves show that the type species of Mer- istorygma has a ramose, radially arranged vascular system surrounding the muscle field with a pair of strong vascula media, which approximately correspond to the strongest pair of costae in the sulcus. The distinctive vascular system in this genus constitutes an important difference between it and Brachythyris, because the latter has a weak, simple, radial, MartiniaTik& vascular system. Distribution. — In addition to the large collection from GSC locality 56430 there is one incomplete specimen from GSC locality C“4087 and four specimens from GSC locality 56430A. A similar species occurs in the Ladrones Limestone of southeastern Alaska. Meristorygma, new species A (Fig. 27.15-27.18) Two disarticulated ventral valves of Meristorygma sp. from GSC locality 56430A preserve the general shape and most other characters of Meristorygma arctica Carter, 1974, but differ distinctly from that species in having four to five clearly expressed bifurcating costae on both flanks and a very thin, weak median Fig. 27. — Paeckelmannelloids, brachythyridoids, and reticularioids. 27.1-27.5, Cantabriella sp. B, ven- tral, dorsal, lateral, anterior, and posterior views, GSC 115657, X 1.5. 27.6-27.10, Cantabriella sp. A, ventral, dorsal, lateral, anterior, and posterior views, GSC 115658, X 1.5. 27.11-27.14, Meristor- ygma arctica Carter, 1974, ventral, dorsal, anterior, and lateral views, GSC 115659, X 1. 27.15-27.18, Meristorygma n. sp. A, ventral and oblique views of two ventral valves, GSC 115660 and 115661, X 1. 27.19-27.32, Phricodothyris asiatica (Chao, 1929), ventral, dorsal, anterior, posterior, and lateral views of three specimens, GSC 115662-115664, respectively, X 1. 172 Annals of Carnegie Museum VOL. 67 rib in the sulcus. Obviously these specimens represent a different, perhaps new, species but the lack of specimens prevents us from describing it. Suborder Delthyridina Ivanova, 1972 Superfamily Reticularioidea Waagen, 1883 Family Elythidae Frederiks, 1924 Subfamily Phricodothyridinae Caster, 1939 Genus Phricodothyris George, 1932 Phricodothyris asiatica (Chao, 1929) (Fig. 27.19^27.32) 1902 Reticularia lineata Martin; Chernyshev, p. 193, pL 20, fig. 9-13. 1929 Squamuiaria asiatica Chao, p. 91, pi. 11, fig. 12-14. 1969 Phricodothyris asiatica (Chao): Pavlova, p. 95, pi. 8, fig. 4; pi. 9, fig. 1, 2, textfig. 10-11. 1990 Phricodothyris asiatica{C\i^o): Liang, p. 285, pi. 62, fig. 1-10; pi. 63, fig. 6-11; pi. 65, fig. 18; textfig. 38. Description. — Average size for genus, moderately ventribiconvex, subcircular to suboval in outline, usually slightly wider than long, especially in early stages; cardinal extremities and lateral and front margins regularly rounded; anterior commissure straight; fold and sulcus absent or very weakly ex- pressed; hingeiine approximately half or less than half maximum width; concentric lamellae distinct but not prominent, numbering about seven to eight per five mm near or anterior to midlength. Ventral valve moderately and evenly convex; beak rather broad, incurved, acute; interarea poorly defined, smooth; delthyrium large, equilaterally triangular, bounded on each side by low stegidial plates that form flanges nearly perpendicular to interarea surface; interior with moderately deep del- thyrial ridges and deeply impressed pair of muscle scars. Dorsal valve slightly less strongly convex than opposite valve; beak broad, blunt, extending into delthyrial opening and partly blocking it; interarea low, relatively narrow, bisected by wide notothyr- ium; interior not observed. Measurements. —See Table 21. This species is characterized by its slightly transversely suboval outline, short hingeline, narrow, distinct, concentric lamella, numbering seven or eight per five mm. Distribution. — Phricodothyris asiatica (Chao, 1929) has been found at GSC locality 56430 (five complete specimens and four ventral valves), two complete and one broken shell at GSC locality 56430A, and one almost complete shell from GSC locality C-4085. This species has been described from the Upper Car- Table 21. — Measurements (in millimeters) o/ Phricodothyris asiatica Chao from GSC locality 56430. GSC number Length Width Thickness Hingeline Width/HL ratio 115662 22.1 22.5 14.2 11.1 2.0 115663 — 21.0 11.8 10.6 2.0 115664 14.0 14.3 9.9 6.7 2.1 -A Fig. 28. — Terebratuloids and retzioids. 28.1-28.20, Beecheria cf. B. itaitubensis (Derby, 1874), ventral, dorsal, lateral, and anterior views of a growth series of five specimens, GSC 115665-115669, respec- tively, X 1. 28.21-28.36, Cranaena nassichuki n. sp., ventral, dorsal, lateral, and anterior views of four specimens, including the holotype (28.21-28.24), GSC 1 15670-1 15673, respectively, X 3. 2837, 28.38, Beecheria sp., ventral and dorsal views of a crashed specimen, GSC 115674, X 1.5. 28.39, 28.40, indeterminate terebratuloid, ventral and anterior views of a ventral valve, GSC 115675, X 1.5. 28.41, Hustedia sp., ventral valve, GSC 115676, X 3. 1998 Carter and Poletaev — Upper Carboniferous Arctic Brachiopods 173 39 174 Annals of Carnegie Museum VOL. 67 Table 22. — Measurements (in millimeters) of the types o/Cranaena nassichuki, n. sp. from GSC locality 56430. GSC number Length Width Thickness 115670 9.1 7.1 5.8 115671 7.0 5.7 4.9 115672 7.1 6.5 3.5 115673 5.9 4.8 3.3 boniferous and Lower Permian of China, Malaysia, Transcaucasus, and the Ural Mountains. Order Retziidina Boucot, Johnson, and Staton, 1965 Superfamily Retzioidea Waagen, 1883 Family Retziidae Waagen, 1883 Genus Hustedia Hall and Clarke, 1893 Hustedia sp. (Fig. 28.41) Comments. — A minute ventral valve of a clearly punctate retziid can be as- signed to the genus Hustedia Hall and Clarke. It bears eight strong subangular costae. A rounded median interspace is present. This specimen falls within the range of morphological variation of several species. Distribution. — GSC locality 56430 (one ventral valve). Order Terebratulida Waagen, 1883 Suborder Terebrtatulidina Waagen, 1883 Superfamily Dielasmatoidea Schuchert, 1913 Family Cranaenidae Cloud, 1942 Genus Cranaena Hall and Clarke, 1893 Cranaena nassichuki, new species (Fig. 28.21-28.36) Holotype. —VignvQS 28.21-28.24, GSC 115670, from GSC locality 56430. Paratypes. — Figures 28.25-28.36, three juveniles, GSC 115671-115673, all from GSC locality 56430. ' Description. — Small for genus, subequally biconvex; outline rounded subpentagonal to guttate; lat- eral profile lenticular; anterior profile of large specimens subovate; fold lacking but well-developed sulcus present in anterior third of adult ventral valves; anterior commissure uniplicate in adult shells, rectimarginate in juveniles. Ventral valve with moderately inflated and narrow umbonal region; lateral slopes evenly convex to lateral margins; sulcus originating well anterior to midlength, rounded, forming emargination in adults in ventral view; beak nearly straight in adults to suberect in juveniles; foramen rounded, epithyridid to mesothyridid; interior with pedicle collar and short dental plates. Dorsal valve with narrow swollen umbonal region laterally delineated by concave flexures; lateral slopes less convex than those of opposite valve; dorsum well rounded posteriorly, becoming flattened or even slightly concave anteriorly; interior with posteriorly perforate undivided dorsally concave hingeplate; loop not observed. Measurements. — See Table 22. Diagnosis. — This species can be differentiated by its well-developed ventral sulcus and moderately narrow ventral umbonal region. Comments. — Two specimens of this species were sectioned. Although coarsely recrystallized internally, enough internal details were gleaned from these sped- 1998 Carter and Poletaev— Upper Carboniferous Arctic Brachiopods 175 mens to suggest that this species is most likely a Cranaena, although the loop was not preserved in either specimen» Cranaena is rare above the Lower Car- boniferous with species being described by Lane (1962) from the Lower Mos- covian of Nevada, Czarniecki (1969) from the Gzheliae of Spitsbergen, and Ka- lashnikov (1980) from the Bashkiriae of Novaya Zemlya and the Asselian of the northern Urals. None of these species is closely similar externally to this Hare Fiord species, all having somewhat broader ventral umbones and thicker profiles. Distribution.— GSC locality 56430 (13 specimens, two sectioned). Family Heterelasminidae Likharev, 1956 Genus Beecheria Hall and Clarke, 1893 Beecheria cf. B. itaitubensis (Derby, 1874) (Fig. 28.1--28.20) 1874 Terebratula itaitubensis Derby, p. 1, pi. 2, fig. 1, 3, 8, 16; pi. 3, fig. 24; pi. 6, fig. 15. Description. — Large, subequally biconvex, moderately compressed, much longer than wide; outline longitudinally guttate to rounded subquadrate; lateral and anterior profiles lenticular; maximum width varying from just in front of midlength to just posterior to it; venter flattened or with shallow sulcus; dorsum in anterior profile arched to evenly rounded; anterior commissure weakly uniplicate to recti- marginate; surfaces smooth except for irregularly spaced growth varices and faint fine growth lines. Ventral valve with moderately elongated umbonal region; lateral slopes and venter evenly and moderately convex; sulcus, if present, very weak and shallow, confined to anterior half of valve; beak ridges rounded but well differentiated; beak nearly erect; foramen large, subovate, permesothyridid; deltidial plates not observed, obscured by beak; interior with strong short dental plates. Dorsal valve moderately inflated, less convex than opposite valve, most convex posteriorly, with moderately to prominently arched dorsum in large specimens, more evenly convex in juveniles; lateral slopes gently convex; beak acute, small, obscured by ventral valve; interior with inner and outer hingepiates sessile and forming tent-like structures, fused medially with valve floor and forming false median septum, bearing crural bases at peaks. Comments. — This species is similar ie outline and size to Beecheria itaitubensis Derby from the Desmoinesian of the Amazon Basin, Brazil. Mendes (1957) iL lustiated a small specimen of this species with similar outline and appearance from the same general horizon and region as Derby’s specimens. Distribution.— GSC locality 60194 (36 specimens). Beecheria sp. (Fig. 28.37, 28.38) CoMifie?it^.=This small, crashed specimen bears the two tent-like structures on the floor of the dorsal valve that characterize the genus Beecheria. A narrow ventral umbonal region and strong foldsulcus preclude a close relationship with the species described above as Beecheria cf. B. itaitubensis (Tierby). The modest size and strong foldsulcus of this specimen suggest similarity to Beecheria dab tonensis Sutherland and Harlow, 1973, from Moscovian-age beds in New Mexico. Most illustrated specimens of the latter have a broader ventral umbonal region and a robust lateral profile, a feature not ascertainable ie this crashed specimen. Distribution.— A single, crushed specimen from GSC locality C“5202. Indeterminate terebratuloid (Fig. 28.39, 28.40) Comments.- — -This disarticulated ventral valve is similar in size to the specimen described above as Beecheria sp. It has a well-developed sulcus like that specimen 176 Annals of Carnegie Museum VOL. 67 but the ventral umbo is broad and well rounded laterally. This specimen is not crushed but lacking a dorsal valve cannot be assigned to a genus. Distribution. — A single ventral valve from GSC locality C-5202. Acknowledgments Walter Nassichuk, Geological Survey of Canada, collected most of the specimens that form the basis of this report and we are grateful to him for his generous support of our efforts, for the loan of his collections, and a critique of this report. Stanislav S. Lazarev assisted us greatly with advice on the identification and classification of the productids. J. T. Dutro, Jr., R. B. Blodgett, and Norman Savage apprised us of and otherwise helped us with the faunas of southeastern Alaska. Bruce Wardlaw of the U. S. 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ANNALS OF CARNEGIE MUSEUM VoL. 67, Number 2, Pp. 181-196 12 May 1998 TAXONOMIC STATUS OF THE EARLY PERMIAN HELODECTES PARIDENS COPE (DIADECTIDAE) WITH DISCUSSION OF OCCLUSION OF DIADECTID MARGINAL DENTITIONS David S Berman Curator, Section of Vertebrate Paleontology Amy C. Henrici Preparator, Section of Vertebrate Paleontology Stuart S. Sumida^ Research Associate, Section of Vertebrate Paleontology Abstract Helodectes paridens Cope (1880) is a problematic Early Permian taxon from Texas that has been considered variously as a diadectid or captorhinid, or simply indeterminate. It was based on a poorly preserved jaw and adjoining elements of the left side of a small skull that was believed to possess a double row of marginal teeth. Thorough preparation of the holotype reveals that one row represents a normal complement of rooted premaxillary and maxillary teeth and the other a row of crowns derived from the opposing teeth of the absent dentary. There are no detectable differences from the commonly encountered Diadectes of the same age, and Helodectes should be regarded as a junior synonym of that genus. As the holotype of H. paridens exhibits no features on which to base specific identity, it is referred to as Diadectes sp. The same interpretation undoubtedly applies to the double row of premaxillary-maxillary marginal teeth in the lost and only known specimen of the equally small holotype of H. isaaci Cope (1880) from the same locality as H. paridens. A second example of Helodectes-Wks, dental preservation is described in the opposing upper and lower jaw elements in a very small Diadectes specimen from the Early Permian of Texas. The double=toothed row preservation of the marginal teeth in these specimens prompts discussion of aspects of the occlusion of the cheek teeth in Diadectes and those in the closely related Late Pennsylvanian Desrnatodon. It is speculated that, in addition to occlusion between the upper and lower cheek teeth, mastication also occurred through contact between the maxillary cheek teeth and the inner surface of the parapet of the dentary, and between the cheek teeth of the dentary and the ventral surface of the secondary palatal shelf of the palatine. Changes in the pattern of attrition indicate that with increased molarization the occlusion between the upper and lower cheek teeth shifted from a strictly side=to=side contact in extremely small juveniles to a strictly dorsoventral, vertical-alignment contact between the upper surfaces of the crowns in adults. In the adult pattern of occlusion the upper and lower cheek-tooth series were only partially aligned dorsoventrally, and it is assumed that the labial margin of the upper series and the lingual margin of the lower series occluded with the inner surface of the deetary parapet and the ventral surface of the secondary palatal shelf of the palatine, respectively. These changes in the molarization and occlusion from extremely small juveniles to adults may have been accompanied by a shift in diet. Key words: Diadectidae {Diadectes), Helodectes, dentition, Early Permian ' Department of Biology, California State University-San Bernardino, 5500 University Parkway, San Bernardino, California 92407. Submitted 3 April 1997. 181 182 Annals of Carnegie Museum VOL. 67 Introduction Olson’s (1947) study of the family Diadectidae was initiated with the aim of 1) better understanding the Cotylosauria, then considered by most to include all the primitive reptiles lacking temporal openings and comprising the Diadecto- morpha, Captorhinomorpha, and Seymouriamorpha; and 2) a revision of the clas- sification of the Reptilia, now considered as a paraphyletic grouping. The dia- dectomorph family Diadectidae was considered by him as occupying a central position to these two endeavors, because it is the family on which Cotylosauria was founded and because of the availability of specimens of its member genera, particularly the type genus Diadectes, that permitted detailed anatomical study. Upon examination of most of the existing diadectid materials collected from the Late Pennsylvanian and Early Permian of North America, Olson (1947) concluded that of the then 11 recognized genera only Diadectes Cope, 1878, Desmatodon Case, 1908, and Diasparactus Case, 1910, are valid taxa. Subsequent studies of new and more complete specimens of the poorly known Desmatodon have strong- ly reaffirmed the validity of this taxon (Vaughn, 1969, 1972; Berman and Sumida, 1995). Seven of the diadectid genera {Empedocles Cope, 1878, Nothodon Marsh, 1878, Chilonyx Cope, 1883, Empedias Cope, 1883, Bolbodon Cope, 1896, Dia- dectoides Case, 1911, and Animasaurus Case and Williston, 1912) were reeval- uated by Olson (1947) as synonymous with Diadectes. He concluded that the specimens on which these genera were based either exhibited no features that could be used to differentiate them from Diadectes or exhibited features which are highly variable with age, such as thickness and rugosity of cranial bones or proportional differences in skeletal elements, and therefore could not be consid- ered to be of generic, or even specific, significance. The remaining genus to be considered and the subject of this report is Helo- decteSy which was originally described and referred to the Diadectidae by Cope (1880). Cope (1880:48-49) described Helodectes as represented by two species, each based on a single specimen of jaw fragment with teeth and collected pre- sumably from the same Early Permian, Wichita Group, site in Texas by Jacob Boll between 1878 and 1880 (Romer, 1958). The specimen on which was founded the first named species, H. paridens, was described by Cope as a left maxilla and probable adjoining premaxilla (AMNH 4346). The jaw fragment was very poorly preserved and encrusted with a very incalcitrant, oxidized iron matrix. Most un- usual and forming the basis of Cope’s (1880) generic diagnosis was the presence of both an inner and outer row of marginal teeth. The second species of Helo~ dectes described by Cope, H. isaaci, was based on a probable fragment of maxilla also possessing a double row of teeth (catalogue number not given) and exhibiting the same poor quality of preservation. According to Case (1911), the holotype and only known specimen of H. isaaci is lost and no specimen can be identified from Cope’s description. The two Helodectes species were distinguished from one another by the number, relative sizes, and arrangements of the teeth. Cope (1880) noted that occurring with the type specimens of both species of Helodectes were diadectid skeletal fragments that could belong to either of them or even to a small specimen of Empedocles, whose teeth were included in the same lot. These ma- terials were not described, however, because of the uncertainty of association. Case (1911:48) believed it was very doubtful that additional specimens of Hel- odectes could be identified with confidence because of the poor quality of the holotypic material. Continuing, he remarked that the “numerous teeth in the frag- 1998 Berman et al. — Helodectes Reassessed as Diadectes 183 ment of the jaw indicate a member of the suborder Pareiasauria, but that the fragment and the roots of the teeth indicate an animal much larger than any well- known member of the suborder.” For these reasons he retained the genus provi- sionally as a captorhinid. In Olson’s (1947) opinion, however, the unassigned skeletal fragments found with the holotype of H. paridens are so poorly preserved that only some of the postcranial fragments can be very doubtfully referred to Diadectidae. He, therefore, concluded that the jaw fragment of H. paridens does not belong to this group, and, although some of the associated postcranial frag- ments are diadectid, the genus must be rejected as indeterminate. Preparation of the holotype of Helodectes paridens reveals that it possesses only a single row of rooted, premaxillary-^maxillary teeth lacking their crowns. The second row of teeth observed by Cope (1880) consists of the crowns of the dentary dentition of the opposing lower jaw that has been lost due to weathering. Peter P Vaughn of the University of California, Los Angeles, realized the true nature and origin of the dentition of Helodectes as early as the mid-1960s. One of us (DSB), while a graduate student of Vaughn, recalls having been told by him the explanation for the double row of teeth in Helodectes. His evidence was almost certainly based initially on an at-hand collection of undescribed jaw and postcranial elements of several extremely small individuals of Diadectes from the Early Permian of Texas (collectively catalogued as MCZ 2780). Included was a set of upper and lower jaw elements that not only exhibits the identical, double- toothed row condition seen in the holotype of H. paridens, but when rejoined in their original, preserved position also reveals clearly the origin of the deceptive appearance of its dentition. He later confirmed his suspicion by examining the holotype of H. paridens. It was not until the loan of the MCZ 2780 material was transferred to the Carnegie Museum of Natural History that the authors became aware of the HelodectesAike set of jaws and realized its implications. Careful examination of the holotype of H. paridens not only resolves the riddle of its dentition, but also provides indisputable evidence that it should be assigned to Diadectidae as Diadectes, although a species assignment cannot be made safely. Presumably the same conclusions would apply to the missing holotype of H. isaaci. The double-toothed row preservation has prompted a reconsideration of the function of the secondary palatal shelf of the palatine and the parapet of the dentary as providing masticatory surfaces for the lower and upper cheek teeth, repectively, in Diadectes and the closely related Late Pennsylvanian Desmatodon. In addition, pronounced changes in the molarization and occlusion of their cheek teeth from extremely small juveniles to adults may have been accompanied by a shift in diet. The following abbreviations are used throughout the text to refer to repositories of specimens: AMNH, American Museum of Natural History, New York, New York; CM, Carnegie Museum of Natural History, Pittsburgh, Pennsylvania; MCZ, Museum of Comparative Zoology, Harvard University, Cambridge, Massachu- setts. Description of Helodectes paridens, Holotype, AMNH 4346 The holotype and only known specimen of Helodectes paridens, AMNH 4346, consists not only of the articulated left premaxilla and maxilla, as originally de- scribed by Cope (1880), but also portions of the adjoining lacrimal, jugal, palatine, 184 Annals of Carnegie Museum VOL. 67 Fig. 1. — Diadectes sp. (AMNH 4346), holotype of Helodectes paridens, consisting of upper left jaw and portions of adjoining elements. A, lateral (anterior to left); B, occlusal; and C, medial (anterior to right) views. Abbreviations: ec, ectoptyergoid; j, jugal; 1, lacrimal; m, maxilla; o, orbital rim; pal, secondary palatal shelf of the palatine; pm, premaxilla; ue, unidentified element. and ectopterygoid (Fig. 1, 2). The holotype is mediolaterally crushed dorsal to the alveolar shelf, and a large, unidentified fragment of bone adheres to its lateral surface at the premaxillary-maxillary union. The premaxilla lacks the dorsal pro- cess and, as explained below, probably a small portion of its anterior, symphyseal region that included the anteriormost tooth. There is seemingly very little missing 1998 Berman et al. — Helodectes Reassessed as Diadectes 185 Fig. 2. — Diadectes sp. (AMNH 4346), holotype of Helodectes paridens. Outline drawing of Figure IB to identify marginal dentitions of upper and lower jaws. Incomplete teeth shown as cross-sectional outlines drawn at level in which preservation ends. Ankylosed premaxillary-maxillary tooth bases shown as open outlines and isolated dentary tooth crowns as filled-in outlines. Abbreviations as in Figure 1. from the maxilla except for the crowns of its teeth, although poor preservation makes it impossible to define all of its borders. The marginal dentition is undoubtedly the most important and interesting aspect of AMNH 4346. Cope’s (1880) original description of the holotype of Helodectes paridens (AMNH 4346) focused on what he believed to be its unique possession of two rows of marginal teeth. It is now quite apparent that the outer or lateral row, consisting of 14 teeth, is the only series of teeth ankylosed to the premaxilla and maxilla. All that remains of this series of teeth are the bases. Those of the first three teeth are subcircular in horizontal section and decrease serially in size posteriorly, with the third being markedly smaller than the two preceding it. The base of the fourth tooth is equal to the first in basal diameter and has a slightly transverse, oval outline, whereas the fifth tooth base is considerably smaller than the fourth and circular in outline. The bases of the succeeding teeth gradually increase in size, particularly in becoming transversely oval, to the tenth, whereas those of the last four teeth, which are noticeably the smallest of the entire series, gradually decrease serially in size posteriorly. In occlusal view the premaxillary- maxillary tooth row forms a slightly sigmoidal curve, with the anterior half of the series forming a labially convex curve and the posterior half forming a lin~ gually convex curve. The bases of the oval, midseries cheek teeth are not oriented directly transversely, as the labial edges are slightly in advance of the lingual edges. Although the premaxillary-maxillary suture cannot be traced across the entire alveolar surface, it probably passed between the third and fourth preserved tooth bases. This determination is based in part on the fact that in Diadectes jaws examined by us, the last premaxillary tooth is typically much smaller than the preceding teeth of the premaxilla and the first tooth of the maxilla. Because the premaxilla of North American Diadectes possesses four teeth, it is also reasoned that the first tooth and the symphyseal portion of the premaxilla surrounding it have been lost. If this analysis is correct, the total tooth count for the upper jaw is four premaxillary and 1 1 maxillary teeth, which is the standard count for Dia- dectes. Medial to the premaxillary-maxillary tooth row is a row of eight teeth, or more accurately their crowns, that undoubtedly represents a portion of the marginal 186 Annals of Carnegie Museum VOL. 67 series of the opposing dentary of the lower jaw. Although the two series lie closely adjacent to one another for most of their length anteriorly, they diverge posteri- orly. The size relationships of the dentary teeth to each other can only be ap- proximated because of poor preservation and the separation of the crowns from their bases at various levels. The anteriormost two tooth crowns lie adjacent to the third and fourth tooth bases of the upper jaw series, with the first being much smaller than the second in cross-sectional size. The succeeding six tooth crowns of the dentary dentition are closely spaced without gaps and, except for the first crown, are preserved adhering to the ventral surface of the palatine between the levels of the seventh and 14th tooth bases of the premaxillary-maxillary series. The posteriormost four crowns are subequal in cross-sectional size and somewhat expanded transversely, with the labial edge of each lying slightly in advance of the lingual edge. Whereas the two crowns preceding the posteriormost four are substantially larger, their margins are too incomplete to determine outline shape. Very little information about the morphology of the crowns is available, because they are closely attached to the bone of the upper jaw and palate and are too closely spaced to prepare fully. The second and fourth crowns are roughly single, conical cusps and are considerably longer than those of the succeeding dentary teeth. All that can be said of the remaining crowns is that those of the fifth and sixth are low and moderately expanded transversely, without any obvious ap- pearance of medial or lateral cusps. The cross-sectional exposures of the teeth of both rows reveal a labyrinthine structure. By way of comparison with North American Diadectes dentitions and consid- ering the spatial relationships of the dentary crowns to the premaxillary-maxillary teeth, the first and second dentary crowns in AMNH 4346 likely represent the third and fifth tooth positions of the complete series. Continuing, if the gap fol- lowing the second preserved tooth crown is considered equivalent to two teeth, then the remaining six posteriormost crowns represent the eighth through 13th tooth positions of the complete series. If this analysis is correct, then probably somewhere between one and five posteriormost dentary teeth are not represented, as, according to Welles (1941), the number of lower jaw teeth in Diadectes lentus from the Early Permian Wichita Group of Texas varies from 14 to 18. However, because the dentaries of small or juvenile specimens of Diadectes generally con- tain fewer teeth than those of adults, it is suspected that the number in AMNH 4346 was probably 14 or possibly 15. As an example, in the skull of the small, immature specimen Diadectes sanmiguelensis MCZ 2989 from the Lower Perm- ian Cutler Formation of Colorado (Lewis and Vaughn, 1965), the dentary pos- sesses a total of 14 teeth. A small, badly weathered portion of the lacrimal remains articulated with the dorsal lamina of the maxilla, but the union is visible only on the medial surface of AMNH 4346. Only the anterior end of the jugal, where it contacts the maxilla along the jaw margin, is preserved and includes, as Cope (1880) noted, a small portion of its entrance into the orbital rim. The incomplete palatine is represented mainly by a narrow, arcuate shelf of bone that extends ventromedially from its contact with the inner margin of the maxillary alveolar shelf. This portion of the palatine is referred to as the secondary palatal shelf by Olson (1947) and Berman and Sumida (1995), and, although its suture with the maxilla cannot be discerned, it is most easily located in AMNH 4346 by the row of six tooth crowns of the dentary adhering to its ventral surface. Undoubtedly, the complete palatine of AMNH 4346 was identical to those in North American and German Diadectes 1998 Berman et al. — Helodectes Reassessed as Diadectes 187 and Desmatodon (Olson, 1947; Berman and Sumida, 1995; Berman et al., 1998) in which a primary palatal shelf of the palatine (absent here), lying a short distance dorsal to the secondary palatal shelf, extended medially to contact the pterygoid as part of the true palate, A portion of the otherwise small ectopterygoid contacts the posterior margin of the secondary palatal shelf of the palatine, the posterior end of the alveolar shelf of the maxilla, and the anterior ventral margin of the medial surface of the jugal. This portion of the ectopterygoid forms a very short continuation of the secondary palatal shelf of the palatine and the anterior medial border of the subtemporal fenestra, which when complete would have contacted the distal end of the anterior margin of the transverse flange of the pterygoid. Olson’s (1947: fig. 3) reconstruction of the skull of Diadectes in ventral view clearly depicts these structural relationships of the ectopterygoid, although the jugal is mislabeled as the squamosal. A Second Example of Helodectes-vake Dental Preservation A pair of opposing upper and lower, left jaw elements (Fig. 3), belonging to a single Diadectes specimen, provides indisputable evidence for the above inter- pretation of the double row of marginal teeth in the holotype of Helodectes par- idens. Both jaw fragments are part of a large collection of undescribed, isolated, tooth-bearing jaw and mainly disarticulated postcranial elements representing at least three extremely small juveniles of Diadectes from the Lower Permian Wich- ita Group of Texas that were collectively catalogued as MCZ 2780 without spe- cific assignment. For the purpose of description, however, the two jaw fragments which are the focus here have been recatalogued as MCZ 9331. They not only duplicate the double-toothed row condition in AMNH 4346, but most importantly can be rejoined in their original, preserved position to demonstrate that one of the two rows of teeth in each jaw represents the crowns of the opposing jaw dentition. The upper left jaw fragment of MCZ 9331 (Fig. 3) includes essentially the entire maxilla, a small portion of the jugal that extends between its contact with the posterior dorsal margin of the maxilla and the anteroventral rim of the orbit, most of the lacrimal, and at least the greater portion of the secondary palatal shelf of the palatine. Although a series of 12 maxillary teeth is represented (Fig. 3C, 4A), the first and last teeth are represented by empty alveoli, teeth 2 through 4 and 6 through 9 are variably incomplete, teeth 5 and 10 are partially erupted crowns, and only tooth 1 1 is essentially complete. All of the teeth except possibly the last two appear to exhibit some transverse expansion, with those of the mid- series region being the most expanded. The crown of a replacement tooth is visible in the lingual pit of the third tooth. It is possible that the partial alveolus at the anterior end of the maxilla actually held the posteriormost premaxillary tooth, as the maxilla of Diadectes typically possesses 1 1 teeth. Although poor preservation makes it difficult to describe serial size changes in the maxillary teeth, some general remarks are possible. On the basis of their basal diameters, the anterior- most and largest teeth of the series decrease serially in size posteriorly to about the fifth tooth, the succeeding three teeth are subequal, and the last four teeth exhibit a noticeable serial decrease in size posteriorly. Lying medially adjacent to maxillary teeth 7 through 1 1 and adhering to the ventral surface of the secondary palatal shelf of the palatine is a series of four, closely spaced crowns belonging to teeth 7 through 10 of the marginal dentition of the opposing lower jaw dentary. 188 Annals of Carnegie Museum VOL. 67 Fig. 3. — Diadectes sp. (MCZ 9331). A, lateral, and B, medial views of upper and lower jaw fragments joined in original preserved position. C and D, upper and lower jaw fragments, respectively, in occlusal (anterior to left) view. Abbreviations: d, dentary; j, jugal; 1, lacrimal; m, maxilla; o, orbital rim; pal, secondary palatal shelf of the palatine. Serial identifications of the separated dentary tooth crowns, as well as those of the maxilla adhering to the dentary discussed below, are easily determined, as the two jaws can be rejoined so that all the crowns except that of the 12th maxillary tooth contact in an exact union the tooth bases of their origin. The lower jaw fragment of MCZ 9331 (Fig. 3) includes only the dentary, which appears to be missing mainly a small portion of its posterior end. A continuous row of 12 tooth bases extends the length of the alveolar shelf (Fig. 3D, 4B). As the small margin of the shelf posterior to the last tooth does not exhibit any signs of additional teeth, the entire marginal series of the dentary may be represented. Typically, however, the dentary of Diadectes held at least 14 teeth. Using basal 1998 Berman et al. — Helodectes Reassessed as Diadectes 189 A Fig. 4. — Diadectes sp. (MCZ 9331), Outline drawings of Figure 3C and D to indicate marginal den- titions of upper and lower jaws. Ankylosed maxillary and dentary tooth bases of A and B, respectively, shown as open outlines and separated dentary and maxillary tooth crowns of A and B, respectively, shown as filled-in outlines. Abbreviations: a, empty alveolus; c, crown of replacement tooth. diameter as an indication of size, in general the first five teeth, which included the largest of the series, decrease serially in size posteriorly, the succeeding four teeth are noticeably smaller and exhibit a modest serial increase in size posteriorly, and the last three teeth, the smallest of the series, decrease serially in size pos- teriorly. The anterodorsal inclination of the first four tooth bases indicates that these teeth were procumbent as in Diadectes. The crowns of replacement teeth in the lingual pits of the first and third teeth also indicate that the first four teeth were Diadectes-\ik& in being bluntly spatulate and incisiform. Lying immediately lateral to teeth 6 through 9 is a series of four closely spaced crowns belonging to teeth 6 through 9 of the marginal dentition of the opposing upper jaw maxilla. Following a small gap, a fifth crown belonging to the 12th maxillary tooth lies a short distance labial to dentary tooth 1 1 . Discussion Taxonomic Status o/ Helodectes Restudy of the Early Permian holotype of Helodectes paridens AMNH 4346 clearly indicates that it should be regarded as a junior synonym of Diadectes of the same age. The poorly preserved bones of the left side of the skull that com- prise AMNH 4346 deviate in no recognizable way from those in Diadectes. Most noteworthy, the unusual structure of the palatine, specifically the presence of the secondary palatal shelf, is known elsewhere only in Diadectes and the very close- ly related Late Pennsylvanian Desmatodon (Olson, 1947; Berman and Sumida, 1995). The extremely small size of AMNH 4346, as well as the Diadectes spec- imens MCZ 2780 and 9331, represents an early ontogenetic stage of growth sel- dom encountered and rarely described in this genus (Olson, 1947; Lewis and Vaughn, 1965; Berman and Sumida, 1995). Small size combined with concomi- tant differences in skeletal proportions and dental features from those of adult 190 Annals of Carnegie Museum VOL. 67 Fig. 5. — Diadectes sp. (MCZ 2780). A, lateral, and B, occlusal views of partial right juvenile maxilla. specimens undoubtedly misled, as Olson (1947) noted, early investigators to in- terpret juvenile specimens of Diadectes as pertaining to other genera. This con- fusion was partly resolved, however, with the recent description (Berman and Sumida, 1995) of the prominent, ontogenetic changes in the marginal dentition of Diadectes. That study clearly accounts for the differences between the dentition of AMNH 4346 and those of adult specimens of Diadectes as reflecting widely separated, ontogenetic growth stages. The dentition of juvenile specimens of Dia- dectes, including AMNH 4346, can be most easily distinguished from that of adults by the cheek teeth. In contrast to adults, the cheek teeth in juveniles do not exhibit extreme transverse expansion and molarization that results in the cen- tral cusp being flanked lingually and labially by a lower, but prominent, shoulder- like cusp. In order to emphasize this observation, Berman and Sumida (1995) stated that the maxillary and dentary cheek teeth in the very immature specimens of Diadectes MCZ 2780 (including MCZ 9331) and D. sanmiguelensis MCZ 2989 exhibit little transverse widening, and are more accurately described as bulbous, with a weakly developed central cusp and essentially no lingual or labial shoulder- like cusps (Fig. 5). Despite representing an early juvenile stage of development, the dentition of AMNH 4346 is still like that of adult specimens of Diadectes in duplicating in greater or lesser degree numerous features of the teeth, such as their number, arrangement, morphology, and serial changes (Case, 1911; Lewis and Vaughn, 1965; Berman and Sumida, 1995). AMNH 4346 might be suspected of being a juvenile specimen of the rare. Late Pennsylvanian Desmatodon, whose cranial anatomy, including its dentition, is nearly identical to that of the later-occurring Diadectes (Vaughn, 1972; Berman and Sumida, 1995). The potential for this confusion, however, is confidently elim- inated by considering the subtle differences in the juvenile dentitions of these two genera first noted by Vaughn (1972) and later more fully documented by Berman and Sumida (1995). They list four dental features of the juvenile maxilla in Des- matodon which are absent or greatly reduced in mature specimens of this genus and apparently not present in specimens of Diadectes of any age: 1) fewer number of teeth, 2) greater relative spacing of teeth, 3) first two maxillary teeth relatively longer and more incisiform, and 4) absence of wear facets. Features 1 and 2 clearly identify AMNH 4346 as Diadectes, whereas poor preservation prevents the use of features 3 and 4. Specific assignment of AMNH 4346 and MCZ 2780 and 9331 is impossible 1998 Berman et al. — Helodectes Reassessed as Diadectes 191 on morphological grounds = However, if one were to follow the recommendations in Olson’s (1947) study of the systematics of Diadectidae, tentative assignment to Diadectes sideropelicus could be suggested on stratigraphic grounds, as it is the only species that he recommended be recognized from the Wichita beds of Texas until such evidence to the contrary is described. Occlusion of Diadectid Marginal Dentitions On the basis of not only dental anatomy, but also features of the entire skeleton, the diadectids Desmatodon and Diadectes were interpreted by Hotton et al. (1997) as the earliest known examples of vertebrates having a primarily high-fiber diet of terrestrial plants. It is, of course, the unusual features of their dentitions which have provided the most persuasive arguments for considering them as herbivores. The anterior, procumbent, rather spatulate, incisiform teeth are ideally suited for grasping and cropping vegetation, whereas the uniquely molariform cheek teeth are indicative of a highly specialized masticatory structure for dealing with a tough, high-fiber plant diet. Wear patterns confirm a vertical motion of the anterior incisiform teeth and a backward or propalinal motion of the lower jaw cheek teeth against those of the upper jaw during occlusion and mastication (Hotton et al., 1997). The unusual preservation of the dentitions of AMNH 4346 and MCZ 9331 prompts speculation of other aspects of the occlusion of the cheek teeth in Dia- dectes and Desmatodon. In the extremely small marginal dentitions of AMNH 4346 and MCZ 2780 and 9331 most of the crown tips of the cheek teeth are either not exposed or are too poorly preserved to reveal wear facets. In a few instances, however, the crowns in MCZ 2780 are exposed and preserved well enough to note that attrition is either absent or limited to small, subcircular facets on the lingual and labial surfaces of the upper and lower cheek teeth, respectively. The pattern of wear is greatly restricted, despite the cheek teeth being bulbous to moderately expanded transversely, and could have been achieved only if occlusion was restricted to contact between the labial surfaces of the lower and the lingual surfaces of the upper cheek teeth during propalinal movement of the lower jaw. The absence of attrition on the tips of the crowns, however, does not necessarily indicate that they did not participate directly in the mastication of food. To the contrary, there are several features of AMNH 4346 and MCZ 9331 which suggest strongly an additional, occlusal component of the cheek teeth that involves the crown tips. The most persuasive feature is the side-by-side preservation of their upper and lower dentitions, with the crown tips of the dentary cheek teeth con- tacting the ventral surface of the secondary palatal shelf of the palatine and the crown tips of the maxillary cheek teeth contacting the dentary lateral to its den- tition. If considered in combination with the limited attrition pattern of the cheek teeth, then it seems very plausible that the above associations of the crown tips of the dentary and maxillary cheek teeth with the palatine and dentary, respec- tively, indicate an additional, important masticatory component of the cheek teeth. Additionally, in AMNH 4346 and MCZ 9331 the secondary palatal shelf of the palatine curves strongly ventromedially, creating a channel-like space between it and the lingual surfaces of the maxillary cheek teeth. Therefore, it is very likely that the lingual margins, as well as the crown tips, of the cheek teeth of the maxilla contacted the secondary palatal shelf during mastication. If occlusion of the dentary cheek teeth with the palatine is accepted, it forces 192 Annals of Carnegie Museum VOL. 67 a reconsideration of previous interpretations of the functional role of the second- ary palatal shelf of the palatine. Commenting on this unique structure in Diadec- tes, now known to occur also in Desmatodon (Berman and Sumida, 1995), Olson (1947:16) noted that the “ventral surface of the process is rough and there usually are a few small teeth along its posterior margin. This has been the basis for considering its primary function as masticatory” by earlier authors. Olson was convinced, however, that the secondary palatal shelf more probably represents the incipient development of a secondary palate. Crucial to the interpretation of this structure is an excellently preserved, isolated palatine of Desmatodon (CM 47674) described in detail and as identical to that in Diadectes by Berman and Sumida (1995). The ventral surface of the secondary palatal shelf was characterized (Ber- man and Sumida, 1995:322-323) as “distinctly sculptured by short, shallow, ir- regular channels and low ridges, and a few, scattered, minute tubercular promi- nences.” The coarse sculpturing of the secondary palatal shelf suggests the pres- ence of a tough, perhaps keratinized, tissue covering. This would have provided an ideal masticatory surface for the dentary cheek teeth. Also pertinent to this discussion is the description (Berman and Sumida, 1995) of a shagreen of den- ticles distributed over areas of the posterior portion of the channel formed between the primary and secondary palatal shelves of the palatine in Desmatodon. This was interpreted as contradictory evidence of Olson’s (1947) theory that the sec- ondary palatal shelf represents a partially developed secondary palate. A twofold, occlusal pattern of the maxillary cheek teeth during mastication, similar to that suggested for the dentary cheek teeth, can also be hypothesized. In specimens of Diadectes and Desmatodon having complete dentaries there exists an unusual, unique structure that suggests the possible presence of a specialized masticatory surface. Lateral to the bases of the cheek teeth is a shallow groove whose outer wall is formed by a vertical extension of the lateral surface of the dentary into a thin, flange-like ridge or parapet (Welles, 1941; Vaughn, 1972; Berman and Sumida, 1995). Welles (1941:424) remarked that “this flange resem- bles that of the Chelonia and Anomodontia and could very well have served a similar purpose; that is, to support a horny cutting beak.” This interpretation, however, is susceptible to the observation that the outer surface of the parapet exhibits the same coarse surface sculpturing that occurs over the entire lateral surface of the lower jaw. That is, there is no distinct change in the bone texture along the outer jaw margin to support the presence of a horny beak, although this does not preclude a cropping function of the parapet. More important to the discussion here, however, is the channel-like basin formed between the parapet and the cheek teeth, as it approximates the occlusal position of the crown-tips and labial margins of the maxillary cheek teeth. Therefore, the parapet of the dentary not only closely duplicates that of the secondary palatal shelf of the palatine in structure, but probably also functioned as a masticatory surface. The internal surface of the parapet, which also forms the outer margin of the floor of the channel-like basin, is coarsely sculptured in a manner very similar to that of the secondary palatal shelf of the palatine and conceivably could have been cov- ered also by a tough, keratinized tissue. In this connection, it is suspected that both the crown tips and labial margins of the maxillary cheek teeth contacted the inner surface of the parapet and channel during mastication. Although incomplete preservation prevents identification of a parapet and groove lateral to the dentary cheek teeth in the juvenile specimens MCZ 2780 and 9331, these structures are present in D. sanmiguelensis MCZ 2989 (Lewis and Vaughn, 1965) which is 1998 Berman et al, — Helodectes Reassessed as Diadectes 193 Fig. 6. — Maxillary dentitions of adult Desmatodon and Diadectes specimens to illustrate attrition patterns of cheek teeth. A, medial view of left maxilla of Desmatodon hesperis (CM 47654); B, occlusal view of portion of left maxilla (anterior to left) of Desmatodon hoUandi (CM 1938) containing teeth 5-8 and the root of 9; and C, occlusal view of portion of right maxilla (anterior to left) of Diadectes lentus (UC 675) showing teeth 7-9. Wear facets have been whitened. approximately the same size. The cheek teeth of MCZ 2989 also exhibit the same reversed pattern of attrition that does not include the crown tips (Lewis and Vaughn, 1965). Some confirmation on the above hypothesis of occlusion of the cheek teeth of diadectids can be found in the dentition of the nearly complete juvenile right maxilla of Desmatodon hesperis CM 47668 (formally UCLA VP 1748) described by Vaughn (1972). By way of comparison, its estimated length of about 55 mm is perhaps only 5 mm greater than that of the maxilla in AMNH 4346. With regard to this discussion, most notable about the maxillary dentition in CM 47668, consisting of eight complete teeth representing the entire series, is the absence of any signs of attrition. This might indicate that during mastication there was little or no direct occlusion between the upper and lower cheek teeth, but rather they occluded principally with the inner surface of the lower jaw parapet. As was pointed out by Vaughn (1972), rapid tooth replacement offers an alternative but less satisfactory explanation for the absence of wear facets in CM 47668. In the development to full maturity the cheek teeth of Diadectes undergo a number of profound morphological changes that result in a much greater degree of molarization (Fig. 6C): 1) dramatic increase in transverse expansion; 2) strong 194 Annals of Carnegie Museum VOL. 67 reduction in the prominence of the central cusp, which in the maxillary and den= tary dentitions occupy positions labial and lingual to the center of the crowns, respectively; and 3) development of prominent, shoulder-like cusps on the labial and lingual margins of the crown which subsequently acquire true cusps. The attrition pattern of the cheek teeth in adult specimens of Diadectes is also more complicated than in juveniles and progresses typically through four stages before tooth replacement occurs (Fig. 6C): stage 1, in the maxillary cheek teeth a sub- circular wear facet occurs on the lingual shoulder cusp and on the lingual side of the central cusp, whereas in the dentary cheek teeth the reverse pattern occurs, with the wear facets appearing on the labial shoulder cusp and the labial side of the central cusp; stage 2, the wear facets of the shoulder and central cusps of stage 1 enlarge and merge to produce a single principal wear facet that is flat or slightly concave, and an additional small subcircular facet appears on the shoulder cusp on the opposite side of the crown as the principal wear facet; stage 3, the principal and shoulder-cusp wear facets of stage 2 enlarge and partially merge, with an eventual loss of the central and shoulder cusps of the principal facet; and stage 4, extreme attrition produces a single flat or slightly concave wear facet that may extend nearly the entire transverse width of the crown. In mature specimens of Desmatodon the cheek teeth never acquire the advance molarization features seen in mature specimens of Diadectes (Vaughn, 1969, 1972; Berman and Sumida, 1995). In this regard, the most notable differences of the cheek teeth of Desmatodon hesperis include (Fig. 6A): 1) transverse expansion is only moderate; 2) central cusp remains very prominent; 3) lingual and labial shoulders of the crown are only moderately developed and do not possess true cusps; and 4) lingual, shoulder-like cusp is more sharply defined than the labial shoulder-like cusp in the cheek teeth of the maxillary, because the base of the central cusp is much more expanded laterally than it is medially, whereas the reverse condition is true in the cheek teeth of the dentary. Along with a lesser degree of molarization, the degree of attrition of the cheek teeth in adult specimens of D. hesperis is limited to a subcircular facet on the lingual and labial shoulder- like cusps in those of the maxilla and dentary, respectively (Fig. 6A). It is also interesting that in adult specimens of D. hesperis almost all of the maxillary cheek teeth exhibit attrition, whereas only rarely do those of the dentary. As examples, in the complete dentition of the holotypic left maxilla CM 47654 (Fig. 6A; for- mally UCLA VP 1706) and in the five cheek teeth of the partial right maxilla CM 47677 of D, hesperis a distinct wear facet occurs on all the teeth except for two far posterior cheek teeth in CM 47654 (Vaughn, 1969, 1972). This is con- trasted by the complete dentitions of the right lower jaw CM 47670 and left dentary CM 47676 of D. hesperis in which the number of cheek teeth exhibiting wear facets is only three and one, respectively (Berman and Sumida, 1995: fig. 3, 4). The above features of attrition suggest that occlusion between the upper and lower cheek teeth was not only restricted to a lingual-labial or side-to-side con- tact, but, as suggested by the dentary dentitions, in some instances was greatly reduced and, therefore, similar to the condition in very juvenile specimens of Diadectes. In the four cheek teeth of the presumed adult holotypic left maxilla CM 1938 (Fig. 6B) and only-described specimen of D. hollandi the degree of molarization and attrition is basically intermediate between those of D. hesperis and adults of Diadectes. Greater transverse expansion of the cheek teeth in CM 1938 is accompanied by reduction in the prominence of the central cusp and the development of well-defined labial and lingual shoulders that lack true cusps, and 1998 Berman et al. — Helodectes Reassessed as Diadectes 195 attrition can be described as characterizing stage 1 in adult specimens of Diadec- tes. It is concluded from these features that in CM 1938 occlusion between upper and lower cheek teeth had begun to shift to a principally dorsoventral, vertical alignment between the upper crown surfaces. This agrees with the observation (Vaughn, 1969, 1972; Berman and Sumida, 1995) that the cheek-tooth dentition in D. hollandi is more advanced than that in D. hesperis. The preceding discussion suggests that with the increased transverse expansion of the cheek teeth in Diadectes and Desmatodon, either as a result of ontogenetic development or evolution, the occlusion between the crowns of the upper and lower series shifts from an exclusively side-to-side contact to one in which there is an increasingly more vertical alignment and a strictly dorsoventral contact be- tween the upper surfaces of the crowns. It is assumed, although not possible to demonstrate, that mastication in adults of Diadectes and Desmatodon still in- volved contact between the upper cheek teeth and the parapet of the lower jaw and the lower cheek teeth and the ventral surface of the secondary palatal shelf of the palatine. Considering the large number of Diadectes specimens available for study, it seems more than coincidental that the sort of double-toothed row preservation of the cheek teeth seen in the juveniles AMNH 4346 and MCZ 9331 has not been reported in any of the far more plentiful adult specimens. The obvious explanation would seem to be that only in extremely juvenile specimens do the upper and lower cheek-tooth series occupy a strictly side-to-side relationship to one another, whether or not they are actively engaged in mastication. If, therefore, postmortem breakage occurs along the jaw line in an extremely juvenile specimen in which the lower jaws are tightly joined to the skull, it is not likely to result in a complete separation between the upper and lower cheek-tooth series, but rather would pro- duce the double-toothed row condition seen in AMNH 4346 and MCZ 9331. On the other hand, under the same circumstances adult specimens of Diadectes would not be expected to produce the same double-toothed row condition, as their op- posing cheek-tooth series occlude exclusively between the upper crown surfaces. Although only rarely is it possible to examine the cheek teeth in a resting state of occlusion in a mature, undistorted specimen of Diadectes, one unusual example is available. The Diadectes skull CM 24127 from the Early Permian Washington Formation of West Virginia is uniquely preserved as an internal mold of an un- distorted skull and mandible that faithfully reproduces anatomical features (Ber- man, 1971). Most importantly, the lower jaws are in their proper orientation, so that the resting, occlusal state of the upper and lower cheek teeth can be clearly interpreted. Although the lateral margin of the dentition is not represented, latex casts allowed a detailed description of all other aspects of the cheek-tooth series (Berman, 1971). Not only do they exhibit an advanced stage of molarization and a reversed pattern of attrition, with many teeth attaining at least an advanced stage 3 of wear, but also a large degree of dorsoventral occlusion between the upper surfaces of the opposing crowns. Hotton et al. (1997) presented a compelling argument that Diadectes, as well as Desmatodon, was primarily adapted to a diet of high-fiber terrestrial vegetation. Their evidence relied almost exclusively on the complex dental adaptations that allowed for the collection and mechanical processing of a wide variety of plant tissues. It was also argued that the large size and body proportions of diadectids, particularly their bulky, rotund torsos, are suggestive of the possession of a dis- proportionally large gut needed for the endomicrobial fermentation of plant cel- 196 Annals of Carnegie Museum VOL. 67 lulose. The conclusions of Hotton et aL, however, were obviously based on ob- servations of adult specimens, and, as noted above, juveniles of Diadectes exhibit striking differences in their cheek dentitions and the manner in which they oc- cluded. In particular, the cheek teeth of juveniles are far less molariform and the area of occlusion is not only much more limited in area, but is restricted to the lingual and labial, rather than the dorsal, margins of the crowns, and thus must have been far less effective in the comminution of plant tissues. These compari- sons raise the question of whether the dentition of juvenile Diadectes was capable of processing terrestrial vegetation as efficiently and of equivalent toughness or fiber content as that consumed by adults. Changes in the dentition from juveniles to adults are too great not to suspect that they were not accompanied by a change in diet; perhaps the juveniles were primarily insectivorous or were omnivorous but restricted to low-fiber vegetation. Thus, our interpretation of the ontogeny of diadectid dentition and mastication lends strong support to preliminary observa- tions by Vaughn (1972:25) that changes in the maxillary dentition in Desmatodon hesperis from juvenile to adult may have paralleled a shift in dietary habit. Acknowledgments The authors wish to express their thanks to the American Museum of Natural History and the Museum of Comparative Zoology for the loan of specimens. Literature Cited Berman, D. S. 1971. A small skull of the Lower Permian reptile Diadectes from the Washington Formation, Dunkard Group, West Virginia. Annals of Carnegie Museum, 43:33-46. Berman, D. S., and S. S. Sumida. 1995. New cranial material of the rare diadectid Desmatodon hesperis (Diadectomorpha) from the Late Pennsylvanian of central Colorado. Annals of Carnegie Museum, 64:315-336. Berman, D. S., S. S. Sumida, and T. Martens. 1998. Diadectes (Diadectomorpha: Diadectidae) from the Early Permian of central Germany, with description of a new species. Annals of Carnegie Museum, 67:53-93. Case, E. C. 1911. A Revision of the Cotylosauria of North America. Carnegie Institute of Washington, Publication 145. Cope, E. D. 1880. Second contribution to the history of the vertebrates from the Permian Formation of Texas. Proceedings of the American Philosophical Society, 19:38-58. Hotton, N., Ill, E. C. Olson, and R. Beerbower. 1997. Amniote origins and the discovery of herbivory. Chapter 7, pp. 207-264, in Amniote Origins (S. S. Sumida and K. L. M. Martin, eds.). Academic Press, New York, New York. Lewis, G. E., and P. P. Vaughn. 1965. Early Permian vertebrates from the Cutler Formation of the Placerville area, Colorado. U. S. Geological Survey, Professional Paper, 503-C:l-50. Olson, E. C. 1947. The family Diadectidae and its bearing on the classification of reptiles. Fieldiana: Geology, 11:1-53. Romer, a. S. 1958. The Texas Permian redbeds and their vertebrate fauna. Chapter 10, pp. 157-179, in Studies on Fossil Vertebrates. Essays Presented to D. M. S. Watson (T. S. Westoll, ed.), Athlone Press, London, United Kingdom. Vaughn, P. P. 1969. Upper Pennsylvanian vertebrates from the Sangre de Cristo Formation of central Colorado. Contributions in Science, Los Angeles County Museum of Natural History, 164:1-28. . 1972. More vertebrates, including a new microsaur, from the Upper Pennsylvanian of central Colorado. Contributions in Science, Los Angeles County Museum of Natural History, 223:1-30. Welles, S. P. 1941. The mandible of a diadectid cotylosaur. University of California Publications in Geological Sciences, 25:423-432. e ) \ / V } I:-. ' ' •S' - f ■ - - ... 1/ ■ - 'U5' i.. , i ■ ».jc# 't. & I / ■: ' >.' i TV ‘iW ' z.'* • >'■■ ■■> .. ,tT)'i ’*-.' KvAi .4f' -hsiSli gV'. i ni r92M V . 1 ’’ i.^V ‘ -r ■ -.^ r- r-- , ~ V:. , f 4 - II '- I" - t.#.4 fM :J»^ ■ • ^ f ^ n..^ii ' ;> " ^ *lspwyHl '» ,% ^ , -A .^iH^ ;■> ■ INSTRUCTIONS FOR AUTHORS ANNALS OF CARNEGIE MUSEUM consist of contributions to the earth sciences (including paleontol- ogy), life sciences, and anthropology, in 30 by 46 picas format (127 by 195 mm or 5 by inches). Submit all manuscripts to the Office of Scientific Publications. Au- thors should give particular attention to scientific con- tent, format, and general style for the ANNALS. 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The scale or lettering for closely cropped photos can be placed in blank areas close to the figure. Proof. — The author should answer all queried proof marks and check the entire proof copy. Return eorrected page proof with the edited manuscript promptly to the Office of Scientifie Publications. If an author chooses to make extensive alterations to a paper in proof stage, the author will bear the cost. Original manuscripts will not be returned unless request- ed. Figures will be returned to the author only if re- quested prior to publication. ISSN 0097-4463 /I S. 3 f 7^ /\Jff AN N AL S of CARNEGIE MUSEUM THE CARNEGIE MUSEUM OF NATURAL HISTORY 4400 FORBES AVENUE • PITTSBURGH, PENNSYLVANIA 15213 VOLUME 67 13 AUGUST 1998 NUMBER 3 CONTENTS ARTICLES Archaeology of Trants, Montserrat. Part 4. Flaked stone and stone bead industries John G. Crock and Robert N. Bartone 197 On the generic status of Palaeophichthys parvulus Eastman 1908 and Mo- nongahela stenodonta Lund 1970 (Osteichthyes: Dipnoi) Anne Kemp 225 Revision of the Neotropical genus Ischyomius with a discussion on its sys- tematic position (Insecta: Coleoptera: Tenebrionoidea: Pythidae) Darren A. Pollock 245 REVIEW Maples of the World (D. M. van Geldersen et al.) .... Frederick H. Utech 265 Editors, ANNALS, BULLETIN and SPECIAL PUBLICATIONS: J. L. Carter, Ph.D B. C. Livezey, Ph.D D. R. Watters, Ph.D M. A. Schmidt, ELS, Assistant Editor Manuscripts, subscriptions, orders for individual numbers, and changes of address should be sent to: Office of Scientific Publications Carnegie Museum of Natural History 4400 Eorbes Avenue Pittsburgh, PA 15213-4080 Phone (412) 622-3287 Pax (412) 622-8837 E-mail: scipubs@clpgh.org ANNALS OF CARNEGIE MUSEUM is published quarterly by The Carnegie Museum of Natural History, 4400 Forbes Avenue, Pittsburgh, Pennsylvania 15213-4080, by the authority of the Board of Trustees of Carnegie Institute. © 1998 Carnegie Institute. @ This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper). ANNALS OF CARNEGIE MUSEUM ¥ql. 67, Number 3, Pp. 197-224 13 August 1998 ARCHAEOLOGY OF TRANTS, MONTSERRAT PART 4. FLAKED STONE AND STONE BEAD INDUSTRIES John G. Crock* Robert N. Bartone^ Abstract Trants is one of the earliest Ceramic period sites known anywhere in the Caribbean. Attribute analysis of lithic materials from the Trants site (MS-Gl) has determined the reduction processes associated with early Ceramic, or “Saladoid,” period flaked stone and stone bead industries. Both industries feature the systematic, on-site reduction of exotic lithic materials. The flaked stone industry is characterized by the expedient production of flakes for utilitarian purposes, with little investment in tools having fixed morphologies. In contrast, the production of stone bead ornaments is characterized by a more complicated and labor-intensive process. This article describes the reduction sequences and compares the general attributes of the Trants assemblages to lithic industries known elsewhere in the region. Key Words; Trants site, lithic analysis, flaked stone, stone bead, grater board teeth hypothesis Introduction The Trants site (MS-Gl) is located on the windward coast of Montserrat, a small volcanic island in the northern Lesser Antilles (Fig. 1). This article focuses on lithic industries represented at the Trants site, specifically flake production and stone bead manufacture, both of which exhibit flaked-stone reduction techniques. This is the fourth in a series of articles concerned with Trants; other articles deal with field methods (Watters, 1994), vertebrate fauna (Reitz, 1994), and settlement data (Petersen, 1996), respectively. All of the lithic materials discussed here were recovered during archaeological investigations conducted at Trants in 1979 (Wat- ters, 1980) and 1990 (Petersen and Watters, 1993). Specimens studied were re- covered during controlled excavation of 1.0 m X 1.0 m or 2.0 m X 2.0 test units, or during systematic surface collections. Following the 1990 fieldwork, flaked-stone artifacts were analyzed to charac- terize the Saladoid period lithic industries represented at the site. The Saladoid occupation at Trants can be dated ca. 500 b.c.-a.d. 300, and likely later. Two flaked-stone industries were identified during the study, both employing nonlocal materials. The first industry involved the production of flakes and fragments for use. The second industry involved the manufacture of stone bead ornaments. The preliminary results of this analysis were presented at the 14th International Congress for Caribbean Archaeology (Bartone and Crock, 1993). These results are developed further here through a comparison with similar investigations and relevant observations from other Caribbean sites. This article also uses these data to evaluate the hypothesis that reduction of chert was primarily geared toward the production of microliths for use as teeth in wooden grater boards. ' Department of Anthropology, University of Pittsburgh, Pittsburgh, Pennsylvania 15260. 2 Department of Anthropology, State University of New York, Binghamton, New York 13902. Submitted 3 November 1993. 197 198 Annals of Carnegie Museum VOL. 67 ! Fig. 1. — Map of the northern Lesser Antilles showing the location of the prehistoric Trants site, located on the east coast of Montserrat. Materials and Methods A total of 2436 lithic artifacts recovered from the Trants site in 1979 and 1990 were analyzed (Bartone and Crock, 1993). Of this total, approximately one-half {n = 1238) was recovered during systematic surface collections and one-half {n = 1198) during controlled, 1.0 m X 1.0 m (1990) and 2.0 m X 2.0 m (1979) test unit excavations (Table 1). Excavated materials were recovered from both 6.4 mm mesh (41%) and finer 3.2 mm mesh (59%) screens. A portion of the total 1990 lithic inventory remains unanalyzed, including specimens recovered from 0.5 m X 0.5 m test pits, some additional 3.2 mm mesh samples, and all material recov- ered from 1 mm mesh screening of feature sediment. The general format of the analysis followed a processual model developed by Collins (1975) with consideration given to the analytical techniques employed by Crabtree (1982). The approach focused on technological aspects of reduction pro- cesses and expressly avoided the use of functional categories during analysis. As noted by Pantel (1988:18), the identification of gross tool types based on function has inherent limitations for the Caribbean because most existing typologies and tool classification systems have been developed for North American and European lithic assemblages. Lithic analysis began with the separation of materials related to stone flake production from those associated with stone bead manufacture. This distinction was based on artifact shape and raw material. Once separated, all artifacts were placed into process-oriented categories based on the stage of reduction they rep- resented. Following assignment to categories, the dimensions and weights of all artifacts were recorded (Table 2), along with attributes specific to each artifact class. For flake production material (predominantly chert), the primary categories include unmodified raw material, core, core fragment, flake, fragmentary flake. 1998 Crock and Bartone — Trants Archaeology, 4 199 Table 1 . — Vertical distribution of flaked stone artifacts recovered during test unit excavation at the Trants site (MS-Gl), Montserrat. Unit/leve!/ stratum '“C dates Cores/ core frags. Core tools Flake debitage Util./ Mod. Debitage Bead preforms Fully shaped beads Bead debitage Total N396 E571 (1.0 X LO m) 0-10 A 16 2 18 10-20 A 25 1 26 20-30 B 9 1 10 30-40 B 5 2 1 8 40-50 B 1 23 24 50-60 C 480 BC ± 80 28 2 1 1 3 35 60-70 C 480 BC ± 80 1 1 2 70-80 C 4 4 80-90 C 1 1 Unit Total 1 0 112 8 2 2 3 128 N421 E645 (1.0 X 1.0 m) 0-10 A 31 31 10-20 A 86 1 1 1 89 20-30 A 18 1 19 20-30 B 39 39 30-40 B 19 1 2 22 40-50 B 8 8 50-60 B 2 2 60-70 B 1 1 60-70 C AD 210 ± 90 1 60 1 5 67 70-80 C 10 BC ± 90 143 1 144 80-90 C 21 21 80-90 D 3 3 90-100 C 1 1 90-100 D 4 4 Unit Total 1 0 436 2 2 2 8 451 N596 E571 (1.0 X 1.0 m) 0-10 A 1 21 1 1 1 25 10-20 A 25 1 1 27 20-30 A 33 3 36 30-40 B AD 60 ± 70 1 54 5 1 61 40-50 B 27 2 1 30 50-60 B 14 1 1 16 60-70 B 17 3 20 70-80 B 440 BC ± 90 4 1 5 80-90 C 2 2 Unit Total 2 0 197 7 2 11 3 222 1979 T. U. (2.0 X 2.0 m) 0-10 4 63 2 1 70 10-20 1 1 102 9 1 5 119 20-30 2 74 7 4 87 30-40 43 7 1 2 53 40-50 3 1 39 1 44 50-60 1 18 4 23 60-70 1 1 Unit Total 11 2 340 30 2 0 12 397 Total Lithics 15 2 1085 47 8 15 26 1198 200 Annals of Carnegie Museum VOL. 67 •2 § •I 2 s a s s s ^ o K K 60 c 5 "Q K Q K 5^ E X 3 I ■| y f_, ^ ^ E ■^3 ON sn 0 t-H r- m ON Tf 'T- NO 0 NO ON NO ^ r- NO CM CvJ 0) m NO _ 00 — ' IT) 0 CN 00 fN CSJ cn 0 ON 00 in ON 0 (N q q O; r- q d NO q q q CM q q q q q q ON f6 K ^ pr d d d d d NO d d d ON d d d d d —i d d d d d d d 1 00 1 00 1 cn 1 00 1 0 r- 1 in 1 ON NO 1 0 1 CM 1 0 1 i r- 0 1 0 ri 00 *0 eo q q fn CN cn ON q 0 0 r-- r4 d .-1 d d d d d d d .-4 q -4 NO — CM CM NO CM rn cn ON f'' CM NO ON ^ u- CM 0 NO NO 00 00 — ' 0 ON ON -H C" CM 00 d ^ 'It q q q q On ^ d 0 ^ ON C' 00 r-- q q 00 00 NO 9 q q 0 ON q d d d 1 d 1 d d d d d d d 1 CMdd dd d d 1 d d d 1 1 d d d 1 1 d d 1 d d 1 1 0 1 m 1 0 E 1 CO 'f 1 CM 1 «n 1 in 1 u- 1 cn 1 ON 1 cn 1 in 1 cn 1 in 0 00 0 0 CM NO 0 0 0 0 0 in m d d d d d d d d d d d d d d d NO in ON 00 CM 0 ON cn NO 0 in ON NO cn in NO -H in 00 m E- CM ON 00 ON cn m 00 q q CM q q q q q q q q O) q CM q q CM q q — < q q q C50 q q q 'It q d 00 1 d d 1 d d 1 d d 1 d d 1 d d 1 d NO 1 — 1 1 cn 1 0 1—1 0 * 1 »— 1 q * . — 1 * 0 CM in ON y—t 0 s NO 00 q d CM 'It 0 d d q in 0 d m d 0 CM d d * 0 d * d V ON d 'It d d 0 d d d in V V d \/ V e c cd C/5 dJ QJ D e w) S bJ) S w) e TO ^ TO ^ TO — Sp^Sp^Sp^So^iS s bflS &dS *^S W)S boS W)S bag C3 ^ C3 ^ ^ s p^ s SbOgbcgboSbJQCbO OT(-irof-iro(-«TO(-irae-i ^ o [L U „ S *0 -^iiw i™i-i>_ ►-i-'t-i ..2 2 fi ,« S 'O (/J (/3 c 'O e « bO kS E .G O ^ E c« ^ S e £ ^ C« S™, tj .S e « 5 o « g Q. « g .S M a ■So Q D S « a W « Qy c« > fe ^ 6.0 □ Debitage Range in cm H Utilized Debitage] Fig. 9. — Histogram showing frequencies by maximum dimension of utilized and nonutilized debitage from the Trants site (MS-Gl), Montserrat. Walker (1983) addressed some of these concerns by conducting use- wear anal- ysis of flakes from the Sugar Factory Pier site in St. ^tts. Using replicated grater board flakes, the size of which was based on ethnographic samples (after DeBoer, 1975), Walker developed a control for measuring length of use, potential function, and degree of polish. He then compared the replicated sample to flakes recovered archaeologically and made a strong case for the presence of grater teeth manu- facture at the Sugar Factory Pier site. The maximum dimension of grater board teeth in Walker’s replicated and archaelogical samples ranged from 2 mm to 10 mm (Walker, 1980:111). If the main output of flake production at Trants was indeed grater board teeth, then, based on Walker’s study, there should be a clustering of utilized and unuti- lized flakes roughly within the range of 2 mm to 10 mm. The Trants data show no such groupings. Instead, the Trants data show clear preference for the use of flakes with maximum dimensions greater than 1 5 mm and no clear size clusters whatsoever. Only a handful of the utilized flakes identified in the lithic sample from Trants even fit the size criteria as defined by Walker. The Trants data do not support the grater teeth hypothesis based on this line of evidence. The use of grater boards at Trants still cannot be ruled out, however. It may be that the use wear associated with this mode of abrasion is too subtle to be identified given the techniques and equipment employed in this study. It also is possible that the sample analyzed did not include flaked stone material 1998 Crock and Bartone — Trants Archaeology, 4 211 Fig. 10. Wai Wai manioc grater board (specimen in the collection of James B. Petersen). 212 Annals of Carnegie Museum VOL. 67 from specific areas of the site where grater board manufacture and/or use took place. Alternatively, grater boards may have been manufactured at only select sites (other than Trants) and traded widely, as is recorded for the Wai Wai and other groups in South America (Evans and Meggers, 1960:334, quoting Thum, 1883; Yde, 1965:34; Chernela, 1992). As described by Mowat (1989:25): “Grab ers of the Arawakan and Tukanoan tribes were particularly prized as trade objects and would be exchanged for such items as blowpipes, pottery, hammocks and hunting dogs.” Whether or not the inhabitants of Trants and other Saladoid period sites actually produced grater boards, they clearly produced flakes for use as simple, expedient tools. The assumption that chert debitage is directly related to the production of grater boards has obscured the importance of flake tools in Saladoid period lithic assemblages in general. The results of the Trants study indicate that flake tools represent a significant output of the flake production industry and, as such, likely served a critical role in Saladoid lifeways at the site. Beyond testing the grater teeth hypothesis, no attempt was made to attribute flake tools to specific functional categories, although likely uses include a host of everyday tasks such as scraping, cutting, smoothing, and engraving. The use of flake tools for such purposes is documented ethnohistorically among the Taino of the Greater Antilles and ethnographically among similarly adapted groups in low- land South America (e.g., Roth, 1924; Loven, 1935; Rouse, 1948a). Ultimately, it may be possible to determine useful functional categories for flake tools in the Caribbean, but this will require more sophisticated analyses. It can be assumed, however, that artifacts with identical patterns of damage, for example, tools with sharply incurvate utilized edges, were used to perform the same or similar func- tions. Flake tool function also can be inferred from other archaeological evidence such as incised shell specimens. Bead Production Raw Material and Reduction Technique Based on the analysis of 113 bead-related specimens from the 1979 and 1990 investigations, the Trants inhabitants were manufacturing stone beads at the site from raw materials such as white quartz, carnelian, feldspar, and jadeite or neph- rite. None of these materials occur naturally in Montserrat and several, including nephrite, likely originated in mainland South America. Unaltered bead raw ma- terial arrived at Trants in the form of relatively small crystals and stone fragments, and these pieces were subsequently reduced and ultimately manufactured into beads. Carnelian, a red-orange, translucent variety of microcrystalline chalcedony, is the most common material represented in the sample of bead-related specimens. Carnelian comprises the large majority of bead preforms (88%) and the majority of bead material debitage (65%). The source of this material remains unknown, although it likely originated outside of the Lesser Antilles, perhaps in South Amer- ica or in the larger islands of the Greater Antilles (Cody, 1990). Initial stages of bead manufacture employed flaked stone reduction techniques, hence bead manufacture is, in part, a flaked stone industry. First- and second- stage bead preforms exhibit small scars expanding inward from transverse margins indicating that excess material was flaked off in these stages. Some flaking in the first steps was probably done freehand, but given the small size of the flakes and 1998 Crock and Bartone — Trants Archaeology, 4 213 the need for greater control, pressure-flaking techniques likely were employed as well. A total of 12 minimally altered artifacts were assigned to the category of Bead Preform 1 (Fig. 11), and 19 exhibiting further modification were assigned to Bead Preform 2. At the Bead Preform 2 stage, specimens were shaped into “block” - sized pieces suitable for further reduction (Fig. 12). Artifacts grouped in the subsequent stage of bead manufacture {n = 12), des- ignated Bead Preform 3, also exhibit small flake scars, but also have tiny pecked “pits” and abrasion striations, all of which indicate that pecking and/or grinding further transformed individual specimens at this stage of reduction (Fig. 13). In general, reduction at this level includes rounding and end tapering through the removal of the “sharp” edges and corners associated with the earlier preform “blocks.” In the final stage of bead manufacture, the bead raw material was fully shaped (designated Fully Shaped Bead). Specimens at this stage {n = 23) are character- ized by the obliteration of all evidence of flaking and pecking through grinding and polishing (Fig. 14). Although exact techniques of grinding are uncertain, a grooved abrading stone curated at the National Trust Museum on Montserrat sug- gests one possible method. The flat stone has a narrow, rounded “track” engraved in its surface which may have been used to abrade beads during the manufacturing process. The “fully shaped” category includes whole and fragmentary beads which have been partially or completely drilled. The majority of these beads exhibit biconical drill holes, indicating they were drilled from two directions with a pointed in- strument (D. Watters, personal communication, 1992; Bartone and Crock, 1993). Several specimens reveal a remnant “cone” in unfinished drill holes, however, suggesting a hollow instrument may have been employed in some cases. A greater variety of raw materials is represented in the “fully shaped” category, including material such as amethyst, which was not present in earlier stages of manufacture. These beads probably were manufactured elsewhere given that these materials occur only in finished bead form and not as preforms or debitage (Har- rington, 1924; Bartone and Crock, 1993). Although the present study focuses on the reduction process and not on the definition of a finished bead typology, four general bead types were recognized among those grouped in the fully shaped category. These include small, flat “discoid” beads of various thicknesses, convex “barrel-shaped” beads of various lengths, cylindrical beads of various lengths, and beads produced through the minimal alteration of natural crystals (e.g., feld- spar). Within these broad categories, beads vary in transverse section from fully rounded to multifaceted, depending on the material. These types are roughly equivalent to those defined by Watters and Scaglion (1994) in a complementary study of 490 specimens in a private collection from the site. The beads and ornaments were collected from Trants in the early 20th century by planter S. W. Howes, whose original collection is currently curated by the National Museum of the American Indian, Smithsonian Institution (formerly the Museum of the American Indian, Heye Foundation). Based on the attribute analyis conducted by Watters and Scaglion (1994), the S. W. Howes collection contains finished beads manufactured from an even greater variety of materials (e.g., turquoise) and ornament types (e.g., zoomorphic pendants) which are unlike anything recovered during the 1979 and 1990 fieldwork. Watters and Scaglion (1994) grouped specimens by raw material and into preform (“blank”) or finished 214 Annals of Carnegie Museum VOL. 67 Fig. 11. — Unmodified and slightly altered bead raw materials (Bead Preform 1) recovered from the Trants site (MS-Gl), Montserrat. 1998 Crock and Bartone — Trants Archaeology, 4 215 Fig. 12. — Minimally shaped bead materials (Bead Preform 2) recovered from the Trants site (MS-Gl), Montserrat. 216 Annals of Carnegie Museum VOL. 67 Fig. 13. — Well-shaped bead materials (Bead Preform 3) recovered from the Trants site (MS-Gl), Montserrat. 1998 Crock and Bartone— Trants Archaeology, 4 217 Fig. 14. — Fully shaped beads recovered from the Trants site (MS-Gl), Montserrat. 218 Annals of Carnegie Museum VOL. 67 product categories. In part, the results of their analysis indicate that manufacturing debris was not well represented in the Howes collection, which appears biased toward nearly finished and finished ornaments. Saladoid Stone Bead Industries Beads manufactured from exotic raw materials have been documented in Sa- ladoid period contexts at many other sites in the Lesser and Greater Antilles; the closest source for many of the raw materials is mainland South America (Myers, 1981; Boomert, 1987). Stone bead manufacture seems to be less frequently rep- resented in earlier preceramic, or later post-Saladoid contexts dated before and after the Saladoid period, respectively. Along with painted and incised ceramic wares, the stone bead industry and the extensive trade networks that supported it are an integral part of the Cedrosan Saladoid legacy (Rouse, 1992). In addition to Trants, unequivocal evidence of on-site stone bead manufacture has been recorded at the Saladoid sites of Pearls in Grenada (Cody, 1990), Pros- perity in St. Croix (Morse, 1989), and the related “Huecoid” site of Sorce in Vieques (Chanlatte Baik, 1984). The notion of regional manufacturing centers and “ports of trade,” originally proposed by Harrington (1924), has gained support as a result of more recent studies (e.g., Watters and Scaglion, 1994). Locating the sources of different exotic bead raw materials and accurately mapping their dis- tribution remains a problem in the Caribbean, however (Cody, 1993). Manufacture of certain bead materials, such as carnelian, appears to have been restricted to certain sites, like Trants, while the distribution of finished products was more widespread. Examples of finished beads are known from sites such as Hacienda Grande in Puerto Rico (Rouse and Alegria, 1990), Hope Estate in St. Martin (Haviser, 1991), Hichman’s in Nevis (Wilson, 1989), Morel in Guadeloupe (Durand and Petijean Roget, 1991), Prosperity in St. Croix (Morse, 1989), the related “Huecoid” site of Sorce in Vieques (Chanlatte Baik, 1984), and Vive in Martinique (Mattioni, 1979), among many others. Stone beads and other orna- ments provide some of the best evidence for the presence of an extensive maritime trade network during the Saladoid period (Watters, 1997). The exchange of beads and ornaments manufactured from exotic materials and their inherent value is also well documented ethnographically and ethnohistori- cally. Their value was undoubtedly associated with the limited supply of exotic raw materials but also likely reflected the skill and labor involved in bead pro- duction. Flaking, pecking, grinding, and polishing likely were labor intensive tasks, but they probably were expedient relative to drilling. Roth (1924:79) writes of Guiana Amerindians producing quartz beads four to eight inches long, drilled lengthwise, a process purportedly taking up to two lifetimes to complete. To drill the beads, they apparently used pointed shoots of plaintain leaves, fine sand, and water (Roth, 1924:79). At the very least, such accounts provide some measure of the labor and equipment required to produce stone beads by hand. Ethnographic accounts from lowland South America and ethnohistoric accounts of Taino trade record the exchange of exotic stones, particularly “greenstone,” or nephrite (e.g., Loven 1935; Myers 1981; Boomert, 1987). These long-range exchange networks, presumably with mainland South America, were in place throughout the entire Saladoid period based on the vertical distribution of bead manufacture materials at Trants. Manufacturing debris and finished beads were found in deeply stratified midden deposits which have been radiocarbon dated as 1998 Crock and Bartone — Trants Archaeology, 4 219 early as 480 b.c. ± 80 (Beta-44828) at Trants (see Table 1). Bead specimens recovered from even deeper, but as yet undated, levels provide even earlier evi- dence of this Amerindian technology, some of the earliest evidence in the entire Caribbean and northeastern South America. Discussion Flake Production The expedient flake technology of the Saladoid period, although relatively sim- ple and inconspicuous, represents a highly efficient method of lithic tool produc- tion. The inventory of flake tools recovered from the Trants site illustrates that this industry was an important part of, if not the sole purpose of, chert pebble and cobble reduction. Although documenting the production of grater board flakes is a more difficult matter, support for widespread production of grater boards among Saladoid people seems probable. Future studies may even show that man- ufacturing centers existed, with production localized at a few localities. Unlike ceramics and crop cultivation, utilitarian flake production was not in- troduced to the Antilles by Saladoid populations. Earlier Archaic populations uti- lized flakes as makeshift tools long before, in addition to manufacturing blades and, less frequently, stemmed, projectile point-like bifaces. Utilized flakes have been documented in the same early, preceramic contexts as diagnostic blades and/ or projectile point-like bifaces in Antigua (Nicholson, 1976a, Davis, 1993), St. Thomas (Bullen and Sleight, 1963; Lundberg, 1991), Puerto Rico (Ortiz, 1976; Pantel, 1988), Hispanola (Davila, 1978; Ortega and Guerrero, 1985; Veloz Mag- giolo, 1991), and Cuba (Kozlowski, 1974). These sites represent at least two definable cultural groups, the Ortoiroid in the Lesser Antilles and Puerto Rico, and the Casimiroid in the Greater Antilles (Rouse, 1992). Given the presence of minimally modified and utilized flake tools in Archaic sites, the only major element of the Archaic flake tool kit missing from later Ceramic period sites are the well-developed and complex blade industries. While it may appear that this represents a regression in lithic technology, it more likely represents the result of different cultural adaptations and a greater emphasis on other industries after the Archaic period. For example, in characterizing the lithic assemblage among Monserrate period sites in Puerto Rico, Roe et al. (1990) suggest that “while essential continuities remain, the absence of a blade technol- ogy in these sites may be the consequence of a technical devolution,” as ground- stone was increasingly emphasized. The need for well-made flake tools among the preceramic groups was not quite so pertinent for later Saladoid people either. Increased trade and a corresponding increase in the supply and variety of raw materials during the Saladoid period, as documented at Trants, suggest that the time invested in conservatively manufac- turing modified flake blades or knives was unnecessary when the sharp edge of any number of the readily available unmodified flakes would suffice. Raw material trade in the Antilles did not originate with ceramic-producing Saladoid peoples, however. Artifacts manufactured from exotic raw materials have been documented at some preceramic sites (e.g., Crock et al., 1995), and it is rather implausible that Archaic peoples acquired all of their material directly. Many earlier preceramic sites are located at or near chert sources (e.g., Kozlowski, 1974; Ortiz, 1976; Haviser, 1989:8), but Goodwin (1978:12) notes that this was not a prerequisite for Archaic period sites. In some cases Archaic populations 220 Annals of Carnegie Museum VOL. 67 apparently did reduce raw materials at the source areas rather than at their habi- tation sites (Davila, 1978), however, which is a different pattern than that observed at Trants and other Saladoid period sites. The abundant source of lithic raw material at Long Island, Antigua, which dominates the lithic inventory at the Saladoid period Trants site, also was ex- ploited by earlier Archaic populations (Davis, 1974, 1993). Long-term usage of this flint deposit by Amerindians resulted in one expansive quarry site which likely represents countless visits by groups and/or individuals seeking raw ma- terial. As noted by Olson (1973:101), there are: “tools, cores and chips strewn all over the place, by the tens of thousands — several acres literally covered with flint.” Investigations of the Long Island source areas and quarry sites indicate that knappers employed a standardized reduction sequence geared toward the pro- duction of high quality cores for easy transport (Gijn, 1993). The repeated use of this particular locale over time makes it one of the most significant lithic raw material sources in the Lesser Antilles, at least for nonbead specimens. The technologically basic tradition of simply produced utilized flakes continued from the preceramic, throughout the Saladoid period, and carried over into later Ceramic periods. Such tools are documented in collections from numerous post- Saladoid sites such as Blackman’s Midden in Antigua (Nicholson, 1976/?), Villa Taina in Puerto Rico (Goodwin and Walker, 1975), and in collections from several sites in Anguilla recently inventoried (Crock and Petersen, 1997). The continuity of this tradition, from the Archaic period onward throughout Amerindian prehis- tory, testifies to its suitability for West Indian environments. Stone Bead Production Analysis of Saladoid period stone bead manufacture at the local level, such as recently begun for the Trants site, indicates that bead production was a compli- cated process which required a higher level of technical proficiency than any other Saladoid period lithic industry. The skill and labor required for bead production certainly contributed to the significance of these ornaments to Amerindians as did the exotic nature of the raw materials used. Saladoid bead manufacturing “centers” such as at Trants in Montserrat, Pearls in Grenada (Cody, 1993), and Sorce in Vieques (Chanlatte Baik, 1984) provide strong support for specialization by individual villages in the production of beads of certain raw materials during the Saladoid period. It is evident that finished beads of these materials (i.e., carnelian and amethyst) occur at a greater variety of sites than does the evidence of their manufacture. This indicates that lapidaries at these manufacturing “centers” probably worked to satisfy both local and re- gional demands for certain types of beads. Archaeological evidence and ethnographic analogy suggest that, once complete, the beads themselves likely symbolized status and/or rank. Finished beads rep- resenting a variety of raw materials were recovered in burial contexts at the Ha- cienda Grande (Rouse and Alegria, 1990), Morel (Durand and Petijean Roget, 1991), and Vive (Mattioni, 1979) sites. Such ceremonial associations, especially when associated only with certain burials, illustrate the use of stone bead orna- ments to distinguish individuals, perhaps to signal status or rank. Among the ethnohistoric Taino of the Greater Antilles and some lowland South American groups, stone beads were highly valued signs of rank which reportedly were 1998 Crock and Bartone — Trants Archaeology, 4 221 passed down over generations (Loven, 1935:478; Rouse, 1948Z?:553; Boomert, 1987:37). Particular attention to the archaeological context of stone bead ornaments will lead to a better understanding of Saladoid period social organization given the roles these items played in conununicating status, rank, or ideological information. At the regional level, the distribution of bead raw materials and finished beads provides the best available evidence for long-distance trade and exchange during the Salaodoid period. The study of Saladoid period bead production is therefore critical, not only to the investigation of lithic technology during this period, but also to the study of Saladoid period socioeconomic systems. Conclusions As ceramic traditions evolved and changed, Saladoid period lithic technologies endured. Given the associations of flake production and bead manufacture with the earliest occupation of the Trants site, one of the earliest now known for the Ceramic period in the Caribbean, it is evident that both traditions were fully in place at the time of Saladoid colonization. At the same time Saladoid populations were adapting to local conditions by exploiting local resources such as Antigua chert, they continued to maintain a connection with mainland South America through trade in exotic bead materials such as amethyst and nephrite. The integrity of lithic industries throughout the Saladoid period helps document the successful adaptation of these mainland peoples who migrated to the Lesser Antilles and Puerto Rico. Acknowledgments Support for the original research discussed in this article was provided by the University of Maine at Farmington Archaeology Research Center (UMF ARC). Both authors participated in the 1990 fieldwork at Trants which was supported by the UMF ARC and the Carnegie Museum of Natural History (CMNH). Several individuals at the UMF ARC deserve thanks and credit for their help, including former Director James B. Petersen, who provided invaluable support and guidance, and William C. Crandall who helped create and manipulate the computer database. Thanks also go to David R. Watters (CMNH) for his support, patience, and invitation to participate in the Trants sym- posium at the 1993 Society for American Archaeology meeting in St. Louis, Missouri, where an earlier version of this paper was presented by the senior author. Petersen, Watters, and several anonymous reviewers provided useful comments on earlier drafts. The authors are grateful to Rich Scaglion for clarifying certain issues concerning statistical applications. All photographs were taken by James B. Petersen and Thomas R. Buchanan at the UMF ARC. Literature Cited Allaire, L. 1985. Changements lithiques dans I’archeologie de la Martinique. Pp. 299-310, in Pro- ceedings of the Tenth International Congress for the Study of the Pre-Columbian Cultures of the Lesser Antilles (L. Allaire and F. M. Mayer, eds.). Centre de Recherches Caraibes, Universite de Montreal, Montreal, Quebec, Canada. Bartone, R. N., and J. G. Crock. 1993. 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National Museum of Denmark, Copenhagen, Denmark. ANNALS OF CARNEGIE MUSEUM VoL. 67, Number 3, Pp. 225-243 13 August 1998 ON THE GENERIC STATUS OF PALAEOPHICHTHYS PARVULUS EASTMAN 1908 AND MONONGAHELA STENODONTA LUND 1970 (OSTEICHTHYES: DIPNOI) Anne Kemp’ Abstract The genus Monongahela Lund 1970, a species of dipnoan based on small tooth plates from the Duquesne limestone. Upper Pennsylvanian, in Mount Washington, Allegheny County, Pennsylvania, has been designated a junior synonym of Palaeophichthys parvulus Eastman 1908. Alternative inter- pretations of the material are, however, possible. Palaeophichthys parvulus is known from two juvenile specimens from the Francis Creek shale, Carbondale Formation, Middle Pennsylvanian, in the Mazon Creek area of Grundy County, Illinois. These are compressed, with macerated heads. The holotype has poorly preserved upper and lower tooth plates. Characters for comparison with Monongahela can be derived only from the upper tooth plates. None of the tooth plate characters that can be reliably distinguished in P. parvulus are sufficient to separate this species from a range of other lungfish genera, and both taxa are found at localities where other dipnoan genera occur. Because additional material and definition of all of the lungfish in the localities are needed to settle the question of congenerity of Palaeophichthys and Monongahela, separation of the two genera should be retained at present. Key Words: fish, Palaeophichthys, Monongahela, Carboniferous, Dipnoi Introduction Palaeophichthys parvulus is a problematic dipnoan taxon, described by East- man (1908) on a small, poorly preserved specimen. A second specimen, slightly larger, was assigned to this species later (Eastman, 1917), but this material adds little information. Both specimens are compressed and have reasonable preser- vation of the body form and the squamatioe, but the heads are macerated. Mo- nongahela stenodonta, described by Lund (1970), is a genus based on an exten- sive series of small tooth plates and jaw bones from a slightly younger locality. No skull bones or compressed fish are known for this genus. Monongahela has recently been made a junior synonym of Palaeophichthys (Schultze, 1994), an opinion based apparently on the existence in the holotype of P. parvulus of a single, partially preserved upper tooth plate that resembles the tooth plates de- scribed as M. stenodonta. Schultze (1994) has further described P. parvulus as a gnathorhizid, despite characterizing the skull roof as having a series of unpaired median bones. Comparison of the two specimens of P. parvulus with tooth plates and body form in hatchlings of one species of a Recent lungfish, Neoceratodus forsteri, suggests that both specimens of P. parvulus are juvenile, as are the tooth plates belonging to M. stenodonta. Diagnosis of species on juvenile material is possible in a bradytelic group like lungfish, but none of the characters used by Schultze (1994) are reliable at generic or specific level. Consideration of both specimens of P. parvulus and comparison of the upper tooth plates with material from other ' Department of Anatomical Sciences, University of Queensland, St. Lucia, Queensland 4072, Austra- lia. a.kemp@mailbox.uq.edu.au. Submitted 11 October 1995. 225 226 Annals of Carnegie Museum VOL. 67 genera, including specimens of Sagenodus cf. S. periprion, contemporaneous with M. stenodonta, suggests that synonymy is premature. Examination of available characters indicates that although Palaeophichthys and Monongahela are not con- generic, it is impossible at this stage to be sure of the true affinities of P. parvulus. Specimen designations: CM, Carnegie Museum of Natural History, Pittsburgh, Pennsylvania; MCZ, Museum of Comparative Zoology, Harvard University, Cam- bridge, Massachussetts; QM, Queensland Museum, South Brisbane, Queensland, Australia; USNM, National Museum of Natural History, Smithsonian Institution, Washington, D. C. Materials and Methods Comparable Populations of Juvenile Dipnoans A large sample of young Neoceratodus forsteri, collected as eggs from the Brisbane River in southeast Queensland and reared in the laboratory, including juvenile fish of stages 53-57 and isolated tooth plates of stages 54-57, has been used for comparison with the holotype and the hypotype of P. parvulus and with other fossil material. Stage 53 hatchlings are three months old and stage 57 hatch- lings are nine months old. Growth and development of young stages of A. forsteri is described in Kemp (1982), and additional details are provided in the description below. Tooth plates and attached jaws of M. stenodonta came from the type locality of the Duquesne limestone (Lund, 1970). Additional juvenile specimens of Sagenodus cf. S. periprion from the same locality as M. stenodonta were used for an outgroup comparison. Specimens of this taxon provide the only possible outgroup comparison, not because they are necessarily appropriate in cladistic terms, but because this population is the only other large sample of juvenile fossil dipnoans that are less derived than Monongahela and Palaeophichthys. Originals of the holotype and hypotype of P. parvulus have been examined in the National Museum of Natural History, Smithsonian Institution, Washington, D. C., and in the Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts, and compared with latex peels of the same specimens in the per- sonal laboratory of Dr. D. Baird, Pittsburgh, Pennsylvania, and at the Museum of Natural History, University of Kansas at Lawrence. The holotype of M. steno- donta has been examined at Carnegie Museum of Natural History, Pittsburgh, Pennsylvania, and a series of specimens were borrowed for more detailed com- parison and analysis. Scanning Electron Microscopy Isolated tooth plates were mounted on stubs, coated in gold, and examined in a Phillips 505 scanning electron microscope. Biometry Measurements of lengths and breadths of the tooth plates and of the angles between the ridges were done by making a camera lucida drawing of the occlusal surface of each tooth plate, set level, and measuring angles and lengths on the drawing with a ruler and protractor. The enlargements of each specimen were the same in every case. Positions of the measurements of lengths, breadths, and angles taken are shown in Figure 1. Length of the tooth plate from the tip of ridge 1 to the tip of the last ridge is the dimension most suitable for tooth plates that are 1998 Kemp — Upper Carboniferous Lungfish 227 Fig. 1. — Measurements taken of tooth plates based on an upper tooth plate of Neoceratodus forsteri at stage 54, and terms used when describing tooth plates. A, angles between ridges; B, length and breadth. Scale bar = 1 mm. Still cusped. Breadth of the tooth plate is taken from the mediolingual junction to the tip of ridge 2. Angle measurements were done by fitting a straight line as closely as possible to the midpoints of each cusp. The ridges rarely meet in a point if the angles are drawn in this way, and further problems arise if the ridge is curved. However, the angles obtained for M. stenodonta are consistent with those obtained by Lund (1970, 1973), who used a different method. Angles between the ridges of the upper tooth plate of P. parvulus are taken from Schultze (1994) and were also measured on his scanning electron micrograph 228 Annals of Carnegie Museum VOL. 67 Fig. 2. — Drawings of the left pterygopalatine tooth plate of the holotype of Palaeophichthys parvulus. A, actual tooth plate drawn from the scan in Schultze (1994) and from latex peels; B, C, tooth plate reconstructed with five ridges; D, E, tooth plate reconstructed with four ridges; F, G, tooth plate reconstructed with three ridges. B, D, and F have ridges meeting in a point, with the medial portion of the tooth plate reconstructed. C, E, and G have no medial portion, and the tooth plates radiate from a line. Scale bar = 1 mm. of the latex peel of the specimen (Fig, 2), Because measurements made on scan- ning electron micrographs require extensive correction before they can be ac- cepted as accurate, these measurements of angles of P, parvulus are of value for descriptive purposes only. Measurements of the lengths of the body and head of living N. forsteri hatch- 1998 Kemp — Upper Carboniferous Lungfish 229 Table 1 . — Characters used to determine the biological age of the specimens. 1 = squamation; 2 = calvarial bone development; 3 = articulation, calvarial bones; 4 = enamel cover; 5 = severity of wear on cusps; 6 = development of secondary denteons. See text for details. Genus Character 1 2 3 4 5 6 Neoceratodus stage 53 full early none full slight none Neoceratodus stage 55 full partial partial full slight none Monongahela 7 ? 7 full slight none Palaeophichthys full partial partial full slight none Sagenodus ? ? 7 full slight none lings, and the depth of head, body, and tail fin were done using calipers on anesthetized specimens. Measurements of the head and body of P. parvulus are taken from Schultze (1994). Determination of the Biological Age of the Material Biological age of a fossil specimen at death can be deduced from a number of characters by comparison with the juvenile, laboratory-reared material of known age from N. forsteri. Characters used to determine the age of the fossil species are squamation, degree of development and of articulation of the skull bones, enamel cover of the cusps, amount of wear on the cusps of the tooth plate, and the number of secondary denteons developed between the primary ridges. In the juvenile material of N. forsteri, these characters can be defined in full (Table 1). Only some can be determined for the fossil material. Characters for Taxonomic Separation of Juvenile Dipnoans Characters are modified from the lists used for adult lungfish in Kemp (1991 <3, 1992a, 1993a) and the characters used by Lund (1970, 1973). Some characters used on adult tooth plates and jaw bones were of no value for juveniles, as they have not yet developed, and others were too variable in young material. Sagen- odus was used as the outgroup comparison. Characters are listed in the Appendix and explained below. The characters chosen apply for generic separation only, as Palaeophichthys is monotypic. Character 1. — Shape of the upper symphysis is linear in Sagenodus. Other shapes, usually oval or half oval, are regarded as derived. Character 2. — Shape of the lower symphysis is oval in Sagenodus, and other shapes, usually linear, are derived. Character 3. — Sagenodus lacks an ascending pterygopalatine process for artic- ulation with the dermal skull roof, but a process of variable shape is present on the pterygopalatine bone in the other genera examined. Character 4. — Petrodentine, as defined by Lison (1941), is not common in dipnoans (Kemp, 1991^, 1992b; Lund et al., 1992). Absence of blocks of petro- dentine in the tooth plates of juvenile lungfish is primitive, and its presence is derived (Kemp, 1995a). Small quantities of petrodentine may develop in older tooth plates of Sagenodus species, but this hard tissue is absent in juvenile Sa- genodus. Characters 5, 6, 7, and 8.-— Shapes of ridge 1 and of the posterior ridges in each jaw are considered separately because they are not always the same. In the 230 Annals of Carnegie Museum VOL. 67 upper tooth plate, curved ridges are primitive, and straight are derived. In the lower tooth plate, straight ridges are primitive, and curved ridges derived. Characters 9, 10, and 11. — Most cusps in lungfish tooth plates show some compression, and this is often greater in the first ridge of a tooth plate compared to more posterior ridges. Compression is also more obvious in lower tooth plates. Slight lateral compression is regarded as primitive, and strong compression as derived. Characters 12 and 13. — Ridge numbers in both upper and lower tooth plates vary according to growth and also show inherent variation. This character is often the same across a wide range of dipnoan genera, and to be useful, it must be defined with care. Juvenile tooth plates with more than five ridges in the upper jaw and more than four in the lower are primitive, and tooth plates with five ridges or less in the upper and four ridges or less in the lower are derived. This compromise applies only to juveniles in the genera considered here. Adult N. forsteri tooth plates have five or more ridges in both upper and lower jaws. Adult Gnathorhiza tooth plates have four ridges in the upper jaw and three in the lower. Sagenodus periprion has numerous ridges in each adult jaw. Character 14. — Ridge 4 in the upper tooth plate, when present, may show the primitive condition, and begin level with ridge 3, at the mediolingual face, or start half way down ridge 3, independent of the mediolingual face. The latter character state is derived. Characters 15 and 16. — In primitive dipnoans, the prearticular or pterygopa- latine bone below the tooth plate extends beyond the dental tissues. In derived dipnoans, it follows the contours of the attached tooth plate. Characters 17 and 18. — In primitive tooth plates, the ridges originate from a mediolingual line or curve. In derived specimens, the ridges of juvenile dipnoans originate approximately from a point situated medially or mediolingually. Systematic Paleontology Subclass Dipnoi Muller 1845 Family incertae sedis Genus Palaeophichthys Eastman 1908 Synonym: None Palaeophichthys Eastman, 1908:253, fig. 37. Palaeophichthys Eastman, 1917:272, pi. 10, fig. 2. Palaeophichthys Schevill, 1932:85. Palaeophichthys Vorobyeva and Obruchev, 1964:314. Palaeophichthys Romer, 1966:362. Palaeophichthys Jessen, 1973:177. Palaeophichthys Bardack, 1979:511. Palaeophichthys Carroll, 1988:612. Palaeophichthys Maples and Schultze, 1989:257, table 1. Palaeophichthys Schultze, 1992:200. Monongahela Schultze, 1992:201. Palaeophichthys Schultze and Marshall, 1993:212. Palaeophichthys Schultze, 1994:106-107. Monongahela Schultze, 1994:107. Amended Diagnosis. — Dipnoan with gracile and elongate body form; head short in relation to body; scales elongate, parallel sided, longitudinally striated with short, vermiculated free field; pterygopalatine tooth plate having straight radiating ridges with strong lateral compression of the cusps in the first ridge and 1998 Kemp — Upper Carboniferous Lungfish 231 slight compression of the cusps in the second and subsequent ridges; prearticular tooth plate indeterminate. Westphalian D, Upper Carboniferous. Type Species. — Palaeophichthys parvulus Eastman 1908. Palaeophichthys parvulus Eastman 1908 Palaeophichthys parvulus Eastman, 1908:253, fig. 37. Palaeophichthys parvulus Eastman, 1917:272, pi. 10, fig 2. Palaeophichthys parvulus Schevill, 1932:85. Palaeophichthys parvulus Vorobyeva and Obruchev, 1964:314. Palaeophichthys parvulus lessen, 1973:177. Palaeophichthys parvulus Schultze, 1994:107. Diagnosis.— As for genus. Stratigraphic Position and Locality. — Francis Creek shale, Carbondale For- mation, Middle Pennsylvanian (Westphalian D); Mazon Creek area, Grundy County, Illinois, USA. Holotype. — MCZ 5090a, b, compressed fish with macerated head. Hypotype. — ^USNM 4433, compressed fish with severely damaged head region. Description. — Palaeophichthys parx’ulus is a dipnoan of slender and elongate shape. The head appears to be short in relation to the length of the body (14-15%) in both specimens, but because the full extent of the fleshy operculum is not preserved, the head may have been longer in the living fish. The tail is diphycercal, and the squamation well defined. The dentition is poorly preserved, and the only useful dental characters come from the pterygopalatine tooth plates. The right is represented by a lateral view of the first ridge only. The left appears to have three, four, or possibly five straight radiating ridges with strong lateral compression of the cusps in the first ridge and slight compression of the cusps in the second and subsequent ridges (Fig. 2A). Depending on the particular reconstruction used (Fig. 2B-G), ridge 4 may begin halfway down ridge 3, or may originate, like the other ridges, from a medial line or curve (Fig. 2B-G). The lower jaw is undiagnostic in both specimens. Despite the poor preservation, it is possible that there are three ridges in the prearticular tooth plate, but this is found in so many dipnoans, juvenile and adult, that it cannot be used for taxonomic determination. The fragment labelled as a vomer in Schultze (1994) has no characteristics of a vomer in the original specimen. It is present only as an impression, and alternative interpretations of this fragment are more probable, perhaps as one of the unidentifiable pieces of scattered and macerated skull bone. In the holotype, the right pterygopalatine tooth plate is represented by three sharp, laterally flattened cusps of the first ridge, still covered to the tip in shiny brown enamel. No other ridges are visible. This appearance is consistent with any one of the species of small dipnoan contemporaneous with Palaeophichthys or Monongahela. The left pterygopalatine tooth plate is represented by an impression, shown as a positive scan by Schultze (1994). Aside from the poor preservation, the mediolingual junction of the tooth plate is missing, as are the labial extremities of all the ridges. Traces of three ridges are present, as well as a probable fourth and a possible fifth (Fig. 2). Schultze (1994) has provided only one of several potential interpretations, and the characters displayed by the specimen are not entirely consistent with those of other dipnoan genera, even Monongahela. One possibility is that five ridges are present (Fig. 2B, C). Ridge 1 has flattened cusps, ridges 2 and 3 have rounded cusps, ridge 4, with two rounded cusps begins in the middle of ridge 3, and ridge 5, with one cusp, has just begun to grow. It is possible that the ridges are long, straight, and acute, and radiate from a medial or posteromedial position (Fig. 2B). Alternatively, the ridges may radiate from a line or curve and may be short (Fig. 2C). The tooth plate may have only four ridges, the first long and straight with laterally flattened cusps, and the subsequent ridges shorter and also straight but with rounded cusps (Fig. 2D, E). The fourth may have begun to grow midway down ridge 3 (Fig. 2D), or all four ridges may radiate from a line or curve (Fig. 2E). It is also possible that only three long, straight ridges are present, radiating from a medial or posteromedial position, with the first long and straight with laterally flattened cusps, and the subsequent ridges shorter and also straight but with rounded cusps (Fig. 2F), or shorter, and radiating from a line or curve (Fig. 2G). Comments. — Specimens of Neoceratodus forsteri, comparable in size with 232 Annals of Carnegie Museum VOL. 67 Fig. 3. — Living hatchling of Neoceratodus forsteri. Scale bar = 1 cm. those of P. parvulus (Fig. 3), are also slender and elongate. The head is long relative to the length of the body, 26-27%, but in the living juveniles considered for this paper, the head was measured to the back of the operculum, largely a fleshy structure in N. forsteri. The measurement of the head is, therefore, not strictly comparable in the two species. The pterygopalatine tooth plates that are commensurate in size with the tooth plate of P. parvulus have four straight ridges, with cusps in all ridges showing some slight lateral compression. In most upper tooth plates, ridge 4 begins midway down ridge 3 (Fig. 4A). Lower tooth plates are similar but have only three ridges at this age (Fig. 4B). Comparable characters with the ability to discriminate generic status in M. stenodonta are the four radiating ridges in the pterygopalatine tooth plate, the first straight and the second and subsequent curved, all having cusps that are strongly compressed laterally (Fig. 4C, D). Cusps of ridge 4 begin to form midway down ridge 3. Lower tooth plates are similar, but have only three ridges. Characters of the body and head of this genus are not known. Vomerine teeth, however, have numerous small cusps (Lund, 1970). Specimens of Sagenodus cf. S. periprion from the Duquesne limestones are known only from a number of upper and lower tooth plates with attached pter- ygopalatine and prearticular bones, all a little larger than P. parvulus. Pterygo- palatine tooth plates have six or more curved ridges (Fig. 4E), and prearticular tooth plates have five or more straight ridges (Fig. 4F). Cusps of the first ridges show slight compression in both upper and lower tooth plates, and are short, rounded cones in the subsequent ridges, with the newest-formed cusp showing slight lateral compression (Fig. 4E, E). In N. forsteri, M. stenodonta, and P. parvulus, ridges radiate roughly from a mediolingual point, and the angles are wide (Eig. 4, Appendix). In S. cf. S. per- iprion, angles are small, and the ridges radiate from a mediolingual line or curve (Eig. 4, Appendix). Biological Age of the Specimens. — Characters for the assessment of the bio- logical age of the specimens are listed in Table 1 . Measurements of the head, tail, and body of N. forsteri juveniles, stages 53-57 (Kemp, 1982), are illustrated graphically in Figure 5, and mean angles between the ridges of upper and lower tooth plates are listed in Table 2. Corresponding available measurements of the angles between the ridges of M. stenodonta and Sagenodus cf. S. periprion spec- imens, as well as angles between the ridges of P. parvulus, are included in the 1998 Kemp — Upper Carboniferous Lungfish 233 Fig. 4. — Scanning electron micrographs of juvenile lungfish tooth plates. A, B, Neoceratodus forsteri. A, upper tooth plate QM 26023; B, lower tooth plate, QM 26024. C, D, Monongahela stenodonta. C, upper tooth plate, CM 25633; D, lower tooth plate, CM 19415. E, F, Sagenodus cf. S. periprion, CM collection. E, upper tooth plate; F, lower tooth plate. Scale bars = 1 mm. 80 70 60 50 40 30 20 10 0 Annals of Carnegie Museum VOL. 67 * * # * *1W * * * * * -L l_ _JI 1 5 10 15 20 Head length (mm) g body lengths of N. forsteri, with the sizes of P. parvulus material included, data for N. forsteri hatchlings, and stars the data for P. parvulus hatchlings. 1998 Kemp — Upper Carboniferous Lungfish 235 'H, 'S o o o m m r- VO 00 »n +1 +1 +i p »n in M vS FI +1 +1 «n o VO m ^ p p p •n VO ni 00 ^ + 1 +1 +1 VO ^ F- d d m ^ s g s ess m CN CN d +1 FI FI gee see in p VO P d r4 o m ^ ro e S S S i>2 « a a -3 s 2 S ^ o «« ^ ^ Oq P Ov « 2 P. P O S a m 236 Annals of Carnegie Museum VOL. 67 table. Relative dimensions of the body and head of P. parvulus are indicated in Figure 5. Although the head of P. parvulus is apparently shorter in relation to the body compared with juvenile N. forsteri, the body form of both species is elongate. Palaeophichthys parvulus is more gracile than N. forsteri, but both specimens fit within the range of body lengths of N. forsteri juveniles (Fig. 5), with the holotype closest to stage 53 of N. forsteri and the hypotype closest to stage 55 (Fig. 5). The holotype of P. parvulus is 3.6 cm long and the body is 0.45 cm in depth (Schultze, 1994). The hypotype is slightly larger, 5.2 cm long and 0.7 cm in depth (Schultze, 1994). The left upper tooth plate of P. parvulus, although not well preserved, displays the characters of many other small lungfish tooth plates. The few surviving cusps are separate and still distinct, and wear appears to be slight. None of the labial or the most medial cusps can be reliably assessed as the ridge tips and the medial face of the tooth plate are broken. There is no trace of the formation of additional dental tissues, represented by secondary denteons (punctations) that cause thick- ening of dental material between the ridges of the tooth plate. The right upper tooth plate, represented by three whole cusps of the first ridge, reveals that the enamel covering of the cusps reached the tips in this specimen. There is no sign of wear on the cusps. Squamation in both specimens is complete, extending as far as the head, and most of the calvarial, palatal, and mandibular bones appear to be present. The macerated condition of the head in both specimens means that it is not possible to describe with any accuracy the pattern of the dermal skull roof or the cheek region of the holotype or the hypotype. Most of the bones have lost their original relationships, and the few that remain close together suggest one pattern in the holotype and another in the hypotype (Schultze, 1994). Articulation may not have been complete in either specimen. As in both specimens of P. parvulus, a full set of squamation is present in N. forsteri from stage 53 onward (Kemp, 1982). At stage 53 in A. forsteri the larger calvarial bones have begun to form and to ossify but are not yet articulated with each other. The smaller calvarial bones have not appeared. Jaws are represented at stage 53 by well-ossified prearticular, articular, and pterygopalatine bones, and the parasphenoid, ceratohyal, and shoulder girdle are also mineralized. Articula- tions of palatal, mandibular, and shoulder girdle bones are forming. The vomer, positioned on the rostral cartilage of the chondrocranium, is also ossified, but never forms an articulation with any bone. At stage 55, most of the calvarial bones are present and anterior calvarial articulations have begun to form, although these are still incomplete, as is the process of ossification of the bones. The descending process of the anterior mediolateral bone, JLM in the nomenclature of Kemp {\992a), and the ascending process of the pterygopalatine bone are not yet fully articulated. The palate and lower jaw, with associated tooth plates, and the shoulder girdle are well ossified and firmly articulated. The tooth plates of N. forsteri at stages 54-56 parallel exactly the structure of the smaller M. stenodonta and Sagenodus tooth plates, of comparable size, and the upper tooth plates of P. parvulus, as far as can be discerned (Fig. 2, 4). The cusps are clearly demarcated, and increase in size from the oldest, first-formed parts of the tooth plate to the youngest labial fringes. Each ridge has no more than six cusps, and each cusp shows little or no wear, even the medial cusps. In some specimens, a complete coat of enamel extends to the tip. Cusps can be 1998 Kemp — Upper Carboniferous Lungfish 237 Table 3. — Data matrix for juvenile dipnoans, generic characters. Characters are listed in Appendix. Genus Character State Sagenodus 00000 00000 00000 000 Neoceratodus 11100 01000 01111 111 Monongahela 00111 00011 mil 111 Palaeophichthys ????! ?1?0? 0171? ? ? ? distinguished all the way to the mediolingual edge. There is no development of secondary denteons or thickening between the primary ridges. This appears much later, along with removal of cusps through wear, and the appearance of additional ridges and destructive removal of enamel and dentine from the mediolingual face of the tooth plate (Kemp, 1977, 1979). All of the specimens of M. stenodonta fall within a narrow size range (Table 2), The smaller specimens, those comparable in size to P. parvulus, have char- acters consistent with those of other juvenile dipnoans (Fig. 4). Cusps are clearly demarcated, and grade in size from the smallest medial cusps to the largest labial cusps that have fused recently to the underlying bone base. The cusps are few in number, sharp, and show little wear. Where wear is present, it is restricted to the medial cusps. Secondary denteons have not yet appeared, and the ridges are cusped to the mediolingual face. These characters are evident in specimens of S. cf. S. periprion as well. Specimens of this species are slightly larger than those of M. stenodonta, but still have sharp conical cusps on each ridge, and all of the cusps are unworn. Posterior ridges with only one or two cusps indicate that the tooth plate was still growing actively at the time of death (Fig. 4E, F). In larger M. stenodonta tooth plates, but not in Sagenodus, medial cusps show considerable wear, and labial cusps are still sharp. However, in both forms, none of the tooth plates show any development of secondary denteons, or thickening of tooth plate material, between the primary ridges of the tooth plate. The range of specimens available extends to material much larger than P. parvulus. Angles between the Ridges. — The angles listed by Schultze (1994) as specific diagnostic characters of P. parvulus are 40° between ridges 1 and 2 and 28° between ridges 2 and 3. He does not give an angle for ridges 3 and 4. Angles measured on the scan by the methods used for other small lungfish in this work are 35°, 24°, 20°, and 18°, respectively (Table 2). Angles between the ridges of pterygopalatine tooth plates in N. forsteri and M. stenodonta are of comparable size (Table 2), and become progressively smaller in the posterior regions of the tooth plate, as in P. parvulus. Angles between ridges of N. forsteri are 51.6°, 35.7°, and 24.7° for ridges 1-2, 2-3, and 3-4 respectively (Table 2). Correspond- ing angles in M. stenodonta are 40.4°, 40.2°, and 31.2° (Table 2). Angles between the ridges of S. cf. S. periprion are much smaller and closer in size, 10.7°, 16.2°, 14.7°, and 20.2° for ridges 1-2, 2-3, 3-4, and 4-5 (Table 2). Character Determination.-—CharactQY states for the species considered in this paper are given in the data matrix (Table 3). The reconstruction of P. parvulus that is closest to the interpretation of Schultze (1994) has been used for the de- termination of characters. Cladistic analysis of these genera has not been performed because much of the data available for N. forsteri, M. stenodonta, and S. cf. S. periprion cannot be predicted or assumed for P. parvulus. The relevant parts of the specimen are missing, reducing the useful information to six characters (Table 3). An estimation 238 Annals of Carnegie Museum VOL. 67 of phylogenetic relationships among the taxa considered here, based on the six characters and using Sagenodus as the outgroup, suggests that P. parvulus and M. stenodonta are both more derived than N. forsteri, but share few derived characters. Discussion Ontogeny of N. forsteri is not always seen as relevant to the study of fossil dipnoan tooth plates (Campbell and Bar wick, 1995), and it is certainly true that N. forsteri is not as closely related to Monongahela and Gnathorhiza as are the other Recent lungfishes, Lepidosiren paradoxa from South America, and species of Protopterus from Africa (Lund, 1970, 1973). Ontogeny of N. forsteri tooth plates is, however, better known than it is in lepidosirenids (Kerr, 1903, 1910; Kemp, 1977, 1979, 1992^, 1995b), and growth stages of M. stenodonta are com- parable (Lund, 1970). Lepidosirenid lungfish also have tooth plates that are de- rived from the fusion of isolated cusps in radiating ridges, a form of development common in many lungfish, including N. forsteri (Kemp, 1995b). At the present state of our knowledge of lepidosirenid ontogeny, comparisons are limited to N. forsteri, and the juvenile tooth plates of this species are less different to those of M. stenodonta than they first appear to be. The ontogeny of N. forsteri is relevant to the study of fossil dipnoans, particularly the more derived of post-Paleozoic genera. Small size does not necessarily mean that a fossil must have been or have come from a juvenile animal. It is not possible to be certain that a fossil known from limited material of minute size represents a juvenile animal or perhaps an adult, because a representative sample of life stages is rarely present. Criteria derived from comparison with living representatives of the group can be used instead to assess the biological age of a fossil at death. These can be drawn from body and head measurements, squamation, degree of development of skull bones, formation of firm articulations between skull bones, and a number of tooth plate characters, like the form of the cusps, their enamel cover, and the development of secondary denteons. All of the fossil tooth plates examined for this study have the characteristics that define young N. forsteri of known stage of development and known biological age. Based on these criteria, it can be shown that P. parvulus and M. stenodonta specimens are the remains of juvenile fishes, and as such can be compared with the tooth plates of juvenile specimens of the Recent lungfish and with other juvenile fossil dipnoans, like specimens of S. cf. S. periprion. The specimens referred to P. parvulus and M. stenodonta are not as easily comparable with the tooth plates of adult lungfishes, including those of Gnathorhiza. It is possible to discriminate species on juvenile material in a bradytelic group like lungfish, even if the only useful parts are tooth plates, provided that characters are chosen with care. This unfortunately leaves a restricted list. Many of the tooth plate characters used for discrimination among adult lungfish (Kemp, 1992n, 1993a), like the form of the prearticular sulcus, have not yet developed fully in hatchlings and juveniles. Other characters, like the origin of ridges, differ in ju- veniles and adults of the same species. In young N. forsteri, the three or four ridges are medial in origin, but in adults, with six or seven ridgos, they are anterior in origin. (A medial origin for adult N. forsteri, stated in Kemp, 1993a:tabie 1, is a mistake.) Shapes in the jaw bones are not always useful. The form of the 1998 Kemp — Upper Carboniferous Lungfish 239 jaw bone behind the tooth plate is often not preserved, and shows little variation. Most dipnoans have strong articular or quadrate processes on the prearticular and pterygopalatine bones, and the shapes are often similar. The same applies to the relationship between the first ridge and the symphysis. This is almost always parasymphyseal. Ridge length is not a reliable character either. In very young tooth plates, with only three or four cusps in a ridge, the addition of a single new cusp can make a large difference to the total length of the ridge. The only points of comparison between P. parvulus and M. stenodonta are the preserved first ridge of the right pterygopalatine tooth plate and the imperfect impression of the left pterygopalatine tooth plate in the holotype of P. parvulus, and the large number of specimens of M. stenodonta tooth plates known. Generic identity has to be demonstrated on the upper tooth plates, as the only lower tooth plate present is too poorly preserved to distinguish any diagnostic characters. Examination of the few reliable characters indicates that separation of the two genera is appropriate, even using the reconstruction that is closest to the inter- pretation of Schultze (1994). Using any of the other possible reconstructions of the left pterygopalatine tooth plate of P. parvulus only serves to increase the differences between P. parvulus and M. stenodonta. This conclusion is, however, provisional, because reliable data are scarce. Skull bones are not consistent between the holotype and hypotype of P. par- vulus, and, apart from indicating that P. parvulus cannot be a gnathorhizid, have no impact on the question of synonymy with M, stenodonta. Species of Gna- thorhiza and related genera all have a double C bone in the medial series (Berman, 1968, 1976, 1979; Olson and Daly, 1972; Kemp, 19931?). The few discernible postcranial features of P. parvulus are not particularly gnathorhizid in character. Juvenile gnathorhizids have been described as having a rounded eel-like body form, with a short, blunt tail and a head one-fifth of the body length (Dalquest, 1968). Using skull and body-form characters, Palaeophichthys is not a gnathor- hizid. There are at least six possible reconstructions of the upper tooth plate of P. parvulus. Each interpretation suggests different affinities for P. parvulus. It is possible that the tooth plate has five ridges, relating the hatchling to Sagenodus or to one of the undescribed species of dipnoan from either locality. It can also be interpreted as having three ridges. A small number of radiating ridges in a young fossil tooth plate, with cusps entirely covered in enamel and no secondary denteons present, is not uncommon. Equivalent characters are found in young Megapleuron zangleri (Schultze, 1977), a species occurring in Pennsylvanian de- posits in North America, and in many other post-Paleozoic dipnoans, including N. forsteri. A third possibility is that it has four ridges, with the fourth ridge originating midway down the third. Joining of the fourth ridge on to the third lateral to the apex of the tooth plate is found in young Gnathorhiza as well as in M. stenodonta, a large proportion of small N. forsteri, and doubtless other dip- noans. Since the apex is missing, all the ridges, three, four, or five, could meet in a point, as is common in young N. forsteri and in M. stenodonta, or join a line or curve as in young Sagenodus. Schultze (1994) has given a specific diagnosis of P. parvulus based entirely on the angles between ridges, and his measurements fall within the range of angles found in M. stenodonta (Lund, 1970, 1973; Schultze, 1994). Angles between the ridges are only valid as taxonomic determinants if based on a large statistical sample (Vorobyeva, 1967; Kemp, 1977). They should be measured on well-pre- 240 Annals of Carnegie Museum VOL. 67 served and preferably original tooth plates, and care should be taken to distinguish the growth angles from the wear angles. Angles also vary among individuals of one species and between juvenile and adult specimens of the same species (Kemp, 1977). Variability is obvious in all of the species measured on tooth plates in this study. Because a diagnosis is intended to characterize a species or a genus to the exclusion of others, beyond reasonable doubt, angles do not have any validity measured on a cast of a single juvenile tooth plate. Vomerine tooth plates of most post-Paleozoic dipnoans resemble a single ridge of the more complex prearticular and pterygopalatine tooth plates, and have cusps at the actively growing labial end. This is true even in large, old specimens. If the structure identified as a vomerine tooth plate by Schultze (1994) is a vomerine tooth plate, the lack of cusps is, firstly, most unusual and, secondly, a significant point of difference with M. stenodonta. This species has numerous minute cusps on the vomerine tooth plate (Lund, 1970). The deposit from which P. parvulus was obtained contains at least three other species of lungfish (Lund, 1975, 1976; Hook and Baird, 1986, 1993). One of these, Conchopoma, differs fundamentally in skull and tooth-plate characters from most other post-Paleozoic dipnoans (Hook and Baird, 1993), and can be excluded as a genus that is close to Monongahela or Palaeophichthys. The deposit from which Monongahela came also contains other species of lungfish (Lund, 1970, 1973, 1975, 1976). None of these dipnoans is perfectly preserved and none can be characterized completely. Palaeophichthys is best classified as incertae sedis, a monotypic genus of uncertain affinities based on incomplete juvenile material. It is unlikely to be closely related to Gnathorhiza because it has an unpaired median C bone in the dermal skull roof (Schultze, 1994). Monongahela is a genus with close relationships to Gnathorhiza, and is known from an extensive series of dental material and attached jaws (Lund, 1970, 1973). Our knowledge of all of these species is incomplete, and Monongahela should not be regarded as a junior synonym of Palaeophichthys. Additional studies of all of these lungfishes are needed to decide questions of generic status. Acknowledgments I am grateful to Drs. D. Baird, R. Lund, and H.-R Schultze for helpful discussions during the preparation of this paper, and to Dr. Baird for his comments on the manuscript and for diagnostic details of the scales of Palaeophichthys parx’ulus. The Australian Research Council Small Grants Scheme provided funds for a visit to the Carnegie Museum of Natural History and the personal laboratory of Dr. Baird, Pittsburgh, Pennsylvania; the Museum of Comparative Zoology, Harvard University, Cambridge, Massachussetts; and the Museum of Natural History, University of Kansas, Lawrence, Kansas, in 1994. An additional Small Grant from the Australian Research Council permitted a visit to the National Museum of Natural History in Washington, D. C., in 1996. The University of Queensland Foundation has supported further research into the Dunkard fishes. Assistance from all sources is acknowledged with many thanks. Literature Cited Bardack, D. 1979. Fishes of the Mazon Creek fauna. Pp. 501-528, in Mazon Creek Fossils (M. H. Nitecki, ed.). Academic Press, New York, New York. Berman, D. S. 1968. Lungfish from the Lueders Formation (Lower Permian, Texas) and the Gna- f/?<9r/iJza-lepidosirenid ancestry questioned. Journal of Paleontology, 42:827-835. . 1976. Cranial morphology of the Lower Permian lungfish Gnathorhiza (Osteichthyes: Dip- noi). Journal of Paleontology, 50:1020-1033. • . 1979. Gnathorhiza bothrotretata (Osteichthyes: Dipnoi) from the Lower Permian Abo For- mation of New Mexico. Annals of Carnegie Museum, 48:211-230. 1998 Kemp — Upper Carboniferous Lungfish 241 Campbell, K. S. W., and R. E. Barwick. 1995. The primitive Dipnoan dental plate. Journal of Vertebrate Paleontology, 15:13-27. Carroll, R. L. 1988. Vertebrate Paleontology and Evolution. Freeman and Company, New York, New York. Dalquest, W. W. 1968. Lungfishes from the Permian of Texas. Copeia, 1968:194-197. Eastman, C. R. 1908. Devonian fishes of Iowa. Iowa Geological Survey 18, Annual Report, 1907: 29-386. - — — . 1917. Fossil fishes in the collection of the United States National Museum. U. S. National Museum Proceedings, 52:235-304. Hook, R. W., and D. Baird. 1986. The Diamond Coal Mine of Linton, Ohio, and its Pennsylvanian- age vertebrates. Journal of Vertebrate Paleontology, 6:174-190. . 1993. A new fish and tetrapod assemblage in the Allegheny Group (Late Westphalian, Upper Carboniferous) of eastern Ohio, USA. Pp. 143-154, in New Results on Permo-Carboniferous Faunas (U. Heidke, ed.). Pollichia-Buch 29, Pfalzmuseum fiir Naturkunde, Bad Durkheim, Ger- many. Jessen, H. 1973. Weitere Fischreste aus dem Oberen Plattenkalk der Bergisch-Gladbach-Paffrather Mulde (Oberdevon, Rheinisches Schiefergebirge). Palaeontographica (Abteilung A), 143:159- 187. Kemp, A. 1977. The pattern of tooth plate formation in the Australian lungfish, Neoceratodus forsteri (Krefft). Zoological Journal of the Linnean Society, London, 60:223-258. — — •. 1979. The histology of tooth plate formation in the Australian lungfish, Neoceratodus forsteri (Krefft). Zoological Journal of the Linnean Society, London, 66:251-287. . 1982. The embryological development of the Queensland lungfish, Neoceratodus forsteri (Krefft). Memoirs of the Queensland Museum, 20:553-597. . 1991a. Australian Mesozoic and Cainozoic lungfish. Pp. 465-498, in Vertebrate Palaeontol- ogy of Australasia (P. Vickers-Rich, J. M. Monaghan, R. F. Baird, and T Rich, eds.). Pioneer Design Studio, Melbourne, Australia. . \99\b. Palaeopathology and lungfish tooth plates. Pp. 441-464, in Early Vertebrates and Related Problems of Evolutionary Biology (M.-M. Chang, Y.-H Lui, and G.-R. Zhong, eds.). Science Press, Beijing, People’s Republic of China. — . 1992a. New neoceratodont cranial remains from the Late Oligocene-Middle Miocene of northern Australia with comments on generic characters for Cenozoic lungfish. Journal of Ver- tebrate Paleontology, 12:284-293. -. \992b. Ultrastructure of the developing dentition in the Australian lungfish, Neoceratodus forsteri (Krefft). Pp. 1 1-33, in Structure, Function and Evolution of Teeth (P. Smith and E. Tchernov, eds.). Freund Publishing House, Tel Aviv, Israel. . 1993a. Ceratodus diutinus, a new fossil ceratodont from Cretaceous and Tertiary deposits in Australia. Journal of Paleontology, 67:883-886. . 1993b. Problematic fossil dipnoans from Australia. New Mexico Museum of Natural History and Science Bulletin, 3:223-221. . 1995a. Complex dentines in fossil lungfish tooth plates. Pp. 210-215, in Proceedings of the Tenth International Symposium on Dental Morphology ( R. J. Radlanzki, H. Renz, eds.). “M” Marketing Services, Berlin, Germany. — -. 1995b. Marginal tooth-bearing bones in the lower jaw of the Recent Australian lungfish, Neoceratodus forsteri, (Osteichthyes:Dipnoi). Journal of Morphology, 225:1-11. Kerr, J. G. 1903. The development of Lepidosiren paradoxa. III. Development of the skin and its derivatives. Quarterly Journal of Microscopical Science, 46:417-459. -. 1910. On certain features in the development of the alimentary canal in Lepidosiren and Protopterus. Quarterly Journal of Microscopical Science, 54:483-518. Lison, L. 1941. Recherches sur la structure et I’histogenese des dents des poissons dipneustes. Ar- chives de Biologie (Paris), 52:279-320. Lund, R. 1970. Fossil fishes from southwestern Pennsylvania. Part 1. Fishes from the Duquesne limestones (Conemaugh, Pennsylvanian). Annals of Carnegie Museum, 41:231-261. — . 1973. Fossil fishes from southwestern Pennsylvania. Part II. Monongahela dunkardensis, new species, (Dipnoi: Lepidosirenidae) from the Dunkard group. Annals of Carnegie Museum, 44:71-101. — 1975. Vertebrate fossil zonation and correlation of the Dunkard Basin. Pp. 171-182, in The Age of the Dunkard, Proceedings of the First 1. C. White Memorial Symposium (J. A. Barlow, ed.). West Virginia Geological and Economic Survey, Morgantown, West Virginia. — 1976. General geology and vertebrate biostratigraphy of the Dunkard Basin. Pp. 225-239, 242 Annals of Carnegie Museum VOL. 67 in The Continental Permian in Central, West and South Europe (H. Falke, ed.). D. Reidel Pub- lishing Company, Dordrecht, Holland. Lund, R., P. Bartholomew, and A. Kemp. 1992. The composition of the dental hard tissues of fishes. Pp. 35-71, in Structure, Function and Evolution of Teeth (P. Smith and E. Tchernov, eds.). Freund Publishing House, Tel Aviv, Israel. Maples, R. and H.-P. Schultze. 1989. Preliminary comparison of the Pennsylvanian assemblages of Hamilton, Kansas, with marine and non-marine contemporaneous assemblages. Pp. 253-273 in Regional Geology and Paleontology of Upper Palaeozoic Hamilton Quarry Area in Southeastern Kansas (G. Mapes and R. Mapes, eds.). Kansas Geological Survey Guidebook, Series 6. Olson, E. C., and E. Daly. 1972. Notes on Gnathorhiza (Osteichthyes: Dipnoi). Journal of Pale- ontology, 46:371-376. Romer, a. S. 1966. Vertebrate Paleontology. Third Edition. The University of Chicago Press, Chi- cago, Illinois. SCHEVILL, W. E. 1932. Fossil types of fishes, amphibians, reptiles and birds in the Museum of Com- parative Zoology. Bulletin of the Museum of Comparative Zoology, 74:57-105. Schultze, H.-P. 1977. Megapleuron zangleri, a new dipnoan from the Pennsylvanian, Illinois. Field- iana: Geology, 33:375-396. . 1992. Dipnoi. Pp. 1-464 in Fossilium Catalogus, Volume 131 (E Westphal, ed.). Kugler Publications, Amsterdam, The Netherlands. — . 1994. Palaeophichthys parvulus Eastman, 1908, a gnathorhizid dipnoan from the Middle Pennsylvanian of Illinois, USA. Annals of Carnegie Museum, 63:105-113. Schultze, H.-R, and C. R. Marshall. 1993. Contrasting the use of functional complexes and isolated characters in lungfish evolution. Memoirs of the Association of Australasian Palaeontologists, 15: 211-224. Vorobyeva, E. I. 1967. A Triassic ceratod from South Fergana and remarks on the systematics and phylogeny of ceratodontids. Paleontologiskii Zhurnal, 1967:102—111 (translated Paleontological Journal, 1967:80-87). Vorobyeva, E. L, and D. V. Obruchev. 1964. Subclass Sarcopterygii. Pp. 268-321, in Osnovy Paleontologii, Volume Two. Agnatha, Pisces (Y. A. Orlov, ed.). Izdatelstvo “NAUKA” SSSR, Moscow, USSR. (In Russian) Appendix Character Descriptions The characters in the following list were used to assess the Recent and fossil tooth plates from juvenile dipnoans. Characters that could be determined on Pa- laeophichthys parvulus, marked with an asterisk, were too few to permit a da- distic analysis of the groups. Primitive states for each character are coded as “0” and determined from the condition of the outgroup, Sagenodus. The derived state is coded as “1.” 1. Upper symphysis linear (0), upper symphysis not linear (1). 2. Lower symphysis oval (0), lower symphysis not oval (1). 3. Ascending pterygopalatine process absent (0), ascending pterygopalatine process present (1). 4. Petrodentine absent (0), petrodentine present (1). *5. Ridge 1 (upper) curved (0), ridge 1 (upper) straight (1). 6. Ridge 1 (lower) straight (0), ridge 1 (lower) curved (1). *7. Posterior ridges (upper) curved (0), posterior ridges (upper) straight (1). 8. Posterior ridges (lower) straight (0), posterior ridges (lower) curved (1). *9. Cusps of ridge 1 (upper) show slight lateral compression (0), cusps of ridge 1 (upper) show strong lateral compression (1). 10. Cusps of ridge 1 (lower) show slight lateral compression (0), cusps of ridge 1 (lower) show strong lateral compression (1). *11. Cusps of posterior ridges show slight lateral compression (0), cusps of posterior ridges show strong lateral compression (1). *12. Five ridges or more in upper jaw (0), fewer than five ridges in upper jaw (1). * Data available from P. parvulus, using the interpretation of Schultze (1994) and the reconstruction of Figure 2D. Kemp — Upper Carboniferous Lungfish 243 1998 13. Four ridges or more in lower jaw (0), fewer than four ridges in lower jaw (1). *14. Ridge 4 (upper) begins level with ridge 3 (0), ridge 4 (upper) begins midway down ridge 3 (1). 15. Pterygopalatine bone extends beyond dentine ridges (0), pterygopalatine bone follows line of dentine ridges (1). 16. Prearticular bone extends beyond dentine ridges (0), prearticular bone follows line of dentine ridges (1). 17. Upper ridges radiate from a mediolingual line (0), upper ridges radiate from a mediolingual point (1). 18. Lower ridges radiate from a mediolingual line (0), lower ridges radiate from a mediolingual point (1). ( Mf I W i H ' w«\Sa i ■ * A »* , V.1: ( ' 'Ifl - • h. « ’ *ii>it«3 o - jt.rt » iV I ) !l w Wru - . . r' i-» T. y^% .» 0»,-| » r ?i , ... *-i.t < ,• ' '<'V ^ ‘ * ^ * tcw.ai ;..' MWW ey^r '* ’ ' ' tVf'fimifpryi^-^!^" I ' i a-"«) l:rtf^i'}|Ni| ♦- k ., •!».’ '■ 'V •■I •i^.//^- i, iMf- * t^^v'isiiyit i '■ .*■'-^■.-^1 , W'- •* ^ ;/w‘l '. ‘t -»ii. j ','. II- E'-' ^ 4.- V,-- ' ■ 's4iK,‘^,'r<|j|t>»d^t<. .■pjiarjit? ■ ,» * f - ^ ^ f ^ t ■* ■■'■• - ■ ^'gB. ^'T' ' ^’> ■ -^ - •'V ''''4l|!M-ti^(iwjfJ( '<^ ••• “ ''*3uf jflbi ' . ->!.i . ndii^ . •— i :• > 'it _ . " ' ■•I,*’,. . ’- S''*’Tm . . ' T-11 .,,'a ■'iSX'dii ix-«,; . . 4i'.'r,,si- . ^ '.rr ^»e A' .- U » 4 ‘ ) . . u .».'«^v:r * , *<» > .•«.'; V ■ BKi#.''. -ifiKl ■•t awl ■?«|i * ' i-^ ? 3^ t - . • . j* ' • #* H- I . «v N'tti;r L-f:,^piB>i< jw •• .-. ■• •-*^. f> /.)^.. D?. J. A H in t. ^iyj|iji,:r. «1 i' . t X t* .* O ’ ' » <• > -<. .U.- ■‘(.v.> I# ^1* • « *•',4 ^ JJ -a '•^t'#' ■ 'it. ■ .'"k... d t*. . ^ I •■ - ■».. I * , ..« ■• > . ■' • ■■ ► ■«^'*fi- ciy4ii9Tt'-'ct ^.h*kA l -^v,- ’ ■ " « *’ ? .. ... ■JIS ”' »■,%, . L *>j,; -g ■-., ia '' B --^ ' 'r-^ • . Sfel,f^iBi''' »:>!.• .ci/, ^ ' k ■rf^. '-'^ 1 i ■■ #?•: -rs^. ;.;; ^ . . 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ANNALS OF CARNEGIE MUSEUM VoL. 67, Number 4, Pp. 267-279 13 November 1998 CONTRIBUTION TO THE KNOWLEDGE OF THE BRENTIDAE (INSECTA: COLEOPTERA) FROM TROPICAL AFRICA, WITH REDESCRIPTION OF PLESIOBOLBUS SAGAX KOLBE Alessandra SforzP Abstract The author lists distributional data for 42 species of Brentidae from tropical Africa, Reported for the first time are distributional records from Senegal (1 sp.), Gambia (3 spp.), Mali (1 sp.), Republic of Guinea (8 spp.), Sierra Leone (2 spp.), Burkina Faso (1 sp.), Togo (2 spp.), Benin (1 sp.), Cameroon (15 spp.), Equatorial Guinea (2 spp.), Gabon (7 spp.), Angola (1 sp.), Zambia (1 sp.), Rwanda (5 spp.), Kenya (1 sp.), and South Africa (2 spp.). Remarks are given upon the geographic distribution of each species. The redescription of Plesiobolbus sagax Kolbe is also given. Key Words: Brentidae, tropical Africa, new records, Plesiobolbus Introduction Undetermined Brentidae (Insecta: Coleoptera) from the Ethiopian Region were studied in the entomological collections of the Carnegie Museum of Natural His- tory (Pittsburgh, Pennsylvania, USA); additional data were added from other pub- lic and private collections (Museum National d’Histoire Naturelle, Paris, France; Museum of Zoology and Entomology, Lund, Sweden; Zoologisk Museum, Co- penhagen, Denmark; Prof. G. Osella, CAquila and Dr. L. Bartolozzi, Firenze, Italy; Mr. B. J. van Vondel, Hendrik Ido Ambacht, Holland). The following paper lists 42 species and includes many new records for the fauna of various tropical African countries. A specimen of Plesiobolbus sagax Kolbe was discovered, a species whose existence was seriously questioned by Damoiseau in his monograph on African Brentidae (1967 a). The following list follows the classification of tribes from Zimmermann (1994) and the species order of Damoiseau (1967a), awaiting a revised organization of the Brentidae which is urgently needed. Only papers published after Damoiseau’s monograph, or not included in that work, are cited for each species. Institutional Abbreviations CBJV, collection of Mr. B. J. van Vondel, Hendrik Ido Ambacht, Holland; CGO, collection of Prof. Giuseppe Osella, UAquila, Italy; CLB, collection of Dr. Luca Bartolozzi, Firenze, Italy; CMNH, Carnegie Museum of Natural History, Pittsburgh, Pennsylvania, USA; MNHN, Museum National d'Histoire Naturelle, Paris, France; MZL, Museum of Zoology and Entomology, Lund, Sweden; MZUF, Museo Zoologico “La Specola,” Firenze, Italy; ZMB, Museum fur Naturkonde der Humboldt-Universitat, Berlin, Germany; ZMC, Zoologisk Museum, Copen- hagen, Denmark. * Museo di Storia Naturale dell’ Uni versita di Firenze, Sezione di Zoologia “La Specola,” via Romana 17, L50125 Firenze, Italy, alessandra@www.specola.unifi.it Submitted 23 April 1996. 267 268 Annals of Carnegie Museum VOL. 67 Subfamily Brentinae Tribe Brentini Debora forficata (J. Thomson) Debora forficata: Damoiseau, 1967a:342; Damoiseau, 1972:268, Specimens Examined. — EQUATORIAL GUINEA. Mongo, 12 Jun 1947, one male (Palau, ex coll. Frieser; CLB). Locality Records. — This species is known from Ivory Coast, Ghana, Cameroon, Gabon, and People’s Republic of Congo; this is the first record for Equatorial Guinea. Spatherhinus ophthalmicus Kolbe Spatherhinus ophthalmicus: Damoiseau, 1967^:347, Specimens Examined. — CAMEROON. Likomba, evening, at light, 11 Feb 1938, one specimen (Buhr; ZMB); Lokom, 10 Sep 1990, one male (Mourglia; CLB). Locality Records.— Thi^ species is known from Gabon and People’s Republic of Congo; this is the first record for Cameroon. Pericordus freyi Damoiseau Pericordus freyt Damoiseau, 1967«:379. Specimens Examined. — ZAMBIA. Southern Province: Lusaka, Kafue River, Kafue city, 1200 m, Dec 1985, one male and one female (Ferrero; CLB). Locality Records.- — -This species is known from Mozambique and South Africa; this is the first record for Zambia. Symmorphocerus approximatus Damoiseau Symmorphocerus approximatus: Damoiseau, 1967a:382; Damoiseau, 1967c: 130. Specimens Examined. — GABON. Ogove R. (= Ogooue River), one female (Good, ex coll. Holland; CMNH). Locality Records.— species is known from Republic of Guinea, People’s Republic of Congo, and Angola; this is the first record for Gabon, Symmorphocerus alluaudi Senna Symmorphocerus alluaudi: Damoiseau, 1967^:385; Quentin, 1970:219. Specimens Examined. — GAMBIA. About 5 km SSW Gunjur, oil palm- and mangrove veg., close to the beach, at light, (18.45-20.30), UTM 28PCK0554, loc. 8, 13 Nov 1977, one female (Cederholm, Danielsson, Hammarstedt, Hedqvist, Samuelsson; MZL). BURKINA FASO. Quest: Bobo Dioulasso, Oct 1982, one male (Politrap, ex coll. Frieser; CLB). TOGO. Region Centrale: Fazao (580 m), 16 Apr 1985, one female (Mourglia; CLB). KENYA. Machakos: Kibwezi, Nov 1905, two specimens (ScheL fler; ZMB). Locality Records. — ^This species is known from Senegal, Ivory Coast, Nigeria, and People’s Republic of Congo; these are the first records for Gambia, Burkina Faso, Togo, and Kenya. The record from Kenya is also the first one of this taxon for East Africa (Fig. 1). 1998 Sforzi^Brentidae from Tropical Africa 269 Fig. L“—Distributioii of Symmorphocerus alluaudi Senna. Triangles: previous records; circles: new records. Afrocordus vicarius (Kleine) Cordus vicarius: Damoiseau, 1967fl:396; Damoiseau, 19671?: 12; Damoiseau, 1975:60. Afrocordus vicarius: Damoiseau, 1980:31; Bartolozzi and Osella, 1990:241. Specimens Examined.— GABON. Ogove R. (— Ogooue River), one male (Good, ex coll. Holland; CMNH). Locality Records.— Thm species is known from Senegal, Republic of Guinea, Sierra Leone, Ivory Coast, Benin, People’s Republic of Congo, Zaire, Angola, Tanzania, and Malawi; this is the first record for Gabon. 270 Annals of Carnegie Museum VOL. 67 Fig. 2. — Distribution of Hemicordus ivorensis Damoiseau. Triangle: previous records; circles: new records. Hemicordus ivorensis Damoiseao Hemicordus ivorensis Damoiseau, 1980:28. Specimens Examined. — SIERRA LEONE. W. Area: Freetown, S. Michel Lodge near Lakta, 9-24 Jun 1988, one female (Mourglia; TOGO. Quest: Fazao (580 m), 16 Apr 1985, one male (Mour= glia; CLB). CAMEROON. Johann- Albrechts-Hohe, 10 Apr 1898, one specimen (ZMB); same locality, Jul-Aug 1897, one specimen (Conradt; ZMB). Locality Records,— Thh species was known only from Ivory Coast; these are the first records for Sierra Leone, Togo, and Cameroon. These new data greatly extend the range of this brentid to the east and west (Fig. 2). 1998 Sforzi — Brentidae from Tropical Africa 271 Perisymmorphocerus latirostris (Power) Perisymmorphocerus latirostris: Damoiseau, 1967fl:401; Damoiseau, 1968:172; Damoiseau, 1972:268; Bartolozzi, 1991:19. Specimens Examined. — GAMBIA. About 5 km SSW Gunjur, oil palm and mangrove veg., close to the beach, at light (18.45-20.30), UTM 28PCK0554, loc. 8, 13 Nov 1977, one male (Cederholm, Danielsson, Hammarstedt, Hedqvist, Samuelsson; MZL); 3.5 km S Georgetown, hilltop at Sankuli Kunda, about 30 m, at light (18.30-20.15), UTM 28PEK2593, loc. 37, 15 Nov 1977, one male (Ced- erholm, Danielsson, Hammarstedt, Hedqvist, Samuelsson; MZL); Tanji River, 3 km SW Bmfut, at light (19.00-21.00), UTM 28PCK087773, loc. 9, 20 Feb 1977, one female (Cederholm, Danielsson, Hammarstedt, Hedqvist, Samuelsson; MZL). Locality Records. — This species is known from Senegal, Equatorial Guinea, Ivory Coast, Ghana, Togo, Nigeria, Sudan, and Ethiopia; this is the first record for Gambia. Micramorphocephalus consobrinus Kleine Micramorphocephalus consobrinus: Damoiseau, 1967fl:409; Damoiseau, 1972:268. Specimens Examined. — REPUBLIC OF GUINEA. Nimba Mts., Mifergui (700 m), at light, 6 May 1993, one female (Perrin; MNHN). Locality Records.— species is known from Senegal, Sierra Leone, Ivory Coast, Ghana, People’s Republic of Congo, and Central African Republic; this is the first record for the Republic of Guinea. Hadramorphocephalus calvei (Power) Hadramorphocephalus calvei: Damoiseau, 1967«:411. Specimens Examined. — MALL Azaoudd: Timbuctu (= Tombouctou), 17 Aug 1927, one male and one female (Madsen; ZMC). Locality Records. — This species is known from Senegal and Chad; this is the first record for Mali. Tribe Cyphagogini Cormopus guineensis Damoiseau Cormopus guineensis Damoiseau, 1967a:38. Specimens Examined. — CAMEROON. Littoral: 25 km WNW Douala, Modeka, loc. 27, secondary forest and plantation, at light, 18 Jan 1978, one specimen (Gardenfors, Hall, and Samuelsson; MZL). Locality Records. — This species was known only from Guinea; this is the first record for Cameroon. Phobetromimus simulans Kleine Phobetromimus simulans: Damoiseau, 1967a:60; Damoiseau, 1972:260; Bartolozzi and Osella 1990: 234. Specimens Examined. — RWANDA. Cyangugu pref: Nyakabuye, 25 Dec 1982, one specimen (Miihle; CLB). Locality Records. — This species is known from Republic of Guinea, Sierra Leone, Ivory Coast, Ghana, Cameroon, People’s Republic of Congo, and Tanza- nia; this is the first record for Rwanda. 272 Annals of Carnegie Museum VOL, 67 Adidactus cancellatus (Lacordaire) Adidactus cancellatus: Damoiseau, 1967a:67; Quentin, 1970:209; Damoiseau, 1972:261; Sforzi, 1992:3. Specimens Examined. — REPUBLIC OF GUINEA. Seredou, at light, 16 May 1975, one specimen (Zott; ZMB). NAMIBIA. East Caprivi: Mudumu Nat. Park, Nakatwa, (18°10'S-23°26'E), at light, 8- 13 Mar 1992, one specimen (Exp. ZMB 1992, Uhlig; ZMB). Locality Records. — This species is known from Ivory Coast, Ghana, Cameroon, Gabon, People’s Republic of Congo, and Tanzania; these are the first records for the Republic of Guinea and Namibia. Xestocoryphus intermedius Damoiseau Xestocoryphus intermedius Damoiseau, 1967(3:91; Damoiseau, 1972:261; Bartolozzi, 1991:4. Specimens Examined. — CAMEROON. Johann-Albrechts-Hohe, one specimen (Conradt; ZMB). Locality Records. — This species is known from Ghana, Zaire, and Tanzania; this is the first record for Cameroon. Xestocoryphus decellei Damoiseau Xestocoryphus decellei Damoiseau, 1967a:94. Specimens Examined. — CAMEROON. Sud Province: Lolodorf, Dec 1914, one specimen (Good; CMNH). Locality Records. — This species is known from Republic of Guinea and Ivory Coast; this is the first record for Cameroon. Rhytidopterus compressithorax (Senna) Rhytidopterus compressithorax: Damoiseau, 1967(3:103. Specimens Examined. — RWANDA. Cyangugu pref.: Nyakabuye, 24 May 1984, one specimen (Miihle; CLB); same locality, 1-17 Feb 1983, one specimen (Muhle; CLB); Gishoma, 20 Feb 1983, one specimen (Miihle; CLB). Locality Records.— This species is known from Ivory Coast, Cameroon, Equa- torial Guinea, People’s Republic of Congo, and Sudan; this is the first record for Rwanda. Podozemius mustus Kolbe Podozemius mustus: Damoiseau, 1967a: 111. Specimens Examined. — SOUTH AFRICA. Natal: Tugela River, Starger, Jan 1989, one specimen (Werner; CLB). Locality Records. — This species is known from Ivory Coast, Cameroon, Equa- torial Guinea, Gabon, and People’s Republic of Congo; this is the first record for South Africa. Podozemius kolbei (Kleine) Podozemius kolbei: Damoiseau, 1967(3:113; Damoiseau, 1967^:5; Damoiseau, 1967c: 126; Damoiseau, 1968:169; Quentin, 1970:210; Damoiseau, 1972:263; Damoiseau, 1975:56. Specimens Examined. — GAMBIA. Bathurst, Jan 1968, one specimen (Palm; MZL). RWANDA. Cyangugu pref.: Nyakabuye, 15-30 Nov 1982, three specimens (Miihle; CLB). Locality Records.—This species is known from Ivory Coast, Ghana, Togo, 1998 Sforzi — Brentidae from Tropical Africa 273 Nigeria, Cameroon, Equatorial Guinea, People’s Republic of Congo, Angola, Su“ dan, Ethiopia, and Tanzania; these are the first records for Gambia and Rwanda. Anomalopleura babaulti (De Muizon) Anomalopleura babaulti: Damoiseau, 1967a: 117. Specimens Examined. — REPUBLIC OF GUINEA. Seredou, 4 Apr 1975, one specimen (Zott; ZMB). Locality Records, — This species was known only from People’s Republic of Congo; this is the first record for the Republic of Guinea. Zemioses rufostriatus (Kleine) Zemioses rufostriatus: Damoiseau, 1967a: 153. Specimens Examined. — RWANDA. Cyangugu pref: Nyakabuye, 24 May 1984, five specimens (Miihle; CEB). Locality Records. — -This species is known from Ivory Coast, Cameroon, Equa- torial Guinea, and People’s Republic of Congo; this is the first record for Rwanda. Tribe Stereodermini Cerobates {Cerobates) conveniens Kleine Cerobates {Cerobates) conveniens: Damoiseau, 1967a:205; Damoiseau, 1967l?:8; Quentin, 1970:214; Damoiseau, 1972:265; Bartolozzi and Sforzi, 1994:250. Specimens Examined. — SENEGAL. Casamance: 1 km NE Djibelor, about 7.5 km SW Ziguinchor, in forest, at light (19.00-21.00), UTM28PCJ5687, loc. 29, 9 Dec 1977, one specimen (Cederholm, Danielsson, Hammarstedt, Hedqvist, Samuelsson; MZL). CAMEROON. SW Province: Fako distr., Bomana, about 1 1 km N Idenau, 300-400 m, (4°13' N-9°4' E), under bark, 21 Jul 1984, two specimens (Davidson; CMNH). Locality Records. — This species is known from Ivory Coast, Ghana, Equatorial Guinea, People’s Republic of Congo, Zaire, and Kenya; these are the first records for Senegal and Cameroon. Cerobates {Cerobates) corruptus Kleine Cerobates {Cerobates) corruptus: Damoiseau, 1967a: 192; Damoiseau, 1967Z?:8; Bartolozzi and Sforzi, 1994:250. Specimens Examined. — CAMEROON. Sud Province: Lolodorf, Sep 1913, one specimen (Good; CMNH). GABON. Kangve, Ogove R. (= Ogooue River), one specimen (Good; CMNH). Locality Records. — This species is known from People’s Republic of Congo, Zaire, Kenya, Tanzania, and Mozambique; these are the first records for Cameroon and Gabon. Cerobates {Cerobates) cruentatus Senna Cerobates {Cerobates) cruentatus: Damoiseau, 1967a:191; Damoiseau, 1967Z?:8; Quentin, 1970:213; Damoiseau, 1972:264; Bartolozzi and Osella, 1990:236; Bartolozzi, 1991:6; Bartolozzi and Sforzi, 1994:250. Specimens Examined. — GABON. Moyen Ogoove: Lambarene, one specimen (Good; CMNH). Locality Records. — This species is known from Ivory Coast, Ghana, Cameroon, Equatorial Guinea, People’s Republic of Congo, Zaire, Uganda, Rwanda, and Kenya; this is the first record for Gabon. 274 Annals of Carnegie Museum VOL. 67 Cerobates {Jonthocerus) burgeoni De Muizon Cerobates {Jonthocerus) burgeoni: Damoiseau, 1967fl:182; Bartolozzi and Osella, 1990:287. Specimens Examined. — SIERRA LEONE. Northern Province: between Sinikoro and Kondembaia, 3 Dec 1984, one male (Rossi; CLB). Locality Records. — This species is known from Ghana and People’s Republic of Congo; this is the first record for Sierra Leone. Cerobates (Jonthocerus) elegans Damoiseau Cerobates {Jonthocerus) elegans: Damoiseau, 1967a: 184; Damoiseau, 1967Z?:6; Damoiseau, 1967c: 127; Damoiseau, 1972:264; Bartolozzi, 1991:8. Specimens Examined. — REPUBLIC OF GUINEA. Seredou, at light, 4 Apr 1975, one specimen (Zott; ZMB). Locality Records. — This species is known from Ghana, Cameroon, People’s Republic of Congo, Zaire, Angola, and Uganda; this is the first record for the Republic of Guinea. Tribe Trachelizini Microtrachelizus fractus Kleine Microtrachelizus fractus: Damoiseau, 1967a:225. Specimens Examined. — CAMEROON. Sud Province: 15 km E Ebolowa Keeke, loc. 15, secondary forest with cocoa plantation, at light, 5 Jan 1978, two specimens (Gardenfors, Hall, Samuelsson; MZL). Locality Records. — This species is known from People’s Republic of Congo; this is the first record for Cameroon. Pseudoceocephalus formosus Kleine Pseudoceocephalus formosus: Damoiseau, 1967a:264; Damoiseau, 1967Z?:13; Damoiseau, 1972:267. Specimens Examined. — CAMEROON. Johann-Albrechts-Hdhe, 3-28 Aug 1898, three specimens (ZMB); same locality, 11 Apr-27 May 1898, one specimen (Conradt; ZMB). REPUBLIC OF GUIN- EA. Seredou, at light, 16 Apr 1975, one specimen (Zott; ZMB). Locality Records. — This species is known from Ghana, Gabon, and People’s Republic of Congo; these are the first records for the Republic of Guinea and Cameroon. Pseudomygaleicus grandis (Damoiseau) Pseudomygaleicus grandis: Damoiseau, 1967a:275; Damoiseau, 1967c:128. Specimens Examined. — EQUATORIAL GUINEA. Subregion unknown: one specimen (Tessman; ZMB). Locality Records.— This species is known from Ivory Coast, Central African Republic, People’s Republic of Congo, Zaire, and Angola; this is the first record for Equatorial Guinea. Pseudomygaleicus surdus Damoiseau Pseudomygaleicus surdus Damoiseau, 1967a;281; Damoiseau, 1975:59. Specimens Examined. — BENIN. Borgou: Parakou, 5-24 Jul 1989, one specimen (van Vondel; CBJV). 1998 Sforzi — Brentidae from Tropical Africa 275 Locality Records.— This species is known from Ivory Coast, Angola, and Uganda, this is the first record for Benin. Gynandrorhynchus tarsalis (Kleine) Gynandrorhynchus tarsalis: Damoiseau, 1967«:294; Damoiseau, 1972:267; Damoiseau, 1975:58; Bar- tolozzi, 1991:14. Specimens Examined. — REPUBLIC OF GUINEA. Nimba Mts., Bossou (550 m), (on dead Ficus sp.), 5 May 1993, one specimen (Perrin; MNHN). CAMEROON. Efulen, Oct-Nov 1913, 23 specimens (Weber; CMNH). GABON. Ogoove Maritime: Batanga, one specimen (Good; CMNH). RWANDA. Cyangugu pref: Nyakabuye, 20 Jul 1984, one specimen (Miihle; CLB). Locality Records.— This species is known from Ivory Coast, Ghana, Zaire, Angola, and Tanzania; these are the first records for Republic of Guinea, Came- roon, Gabon, and Rwanda. Gynandrorhynchus audax (Kleine) Gynandrorhynchus audax: Quentin, 1966:1641; Damoiseau, 1967a:296. Specimens Examined. — CAMEROON. Johann-Albrechts-Hohe, 14 Sep-6 Oct 1898, one specimen (Conradt; MZB); same locality, 11 Apr-27 May 1898 one specimen (Conradt; MZB); same locality, 30 Jan 1898, one specimen (Conradt; MZB). Locality Records.— This species is known from Ivory Coast, People’s Republic of Congo, Zaire, and Mozambique; this is the first record for Cameroon. Anampyx edentulus Damoiseau Anampyx edentulus: Damoiseau, 1967^:308. Specimens Examined. — CAMEROON. Nasanakang, one specimen (Diehl; ZMB); Sud Province: Lolodorf, 8 Feb-27 Mar 1895, one specimen (Conradt; ZMB); Efulen, 28 Nov-29 Dec 1911/12, eight specimens (Weber; CMNH). Locality Records. — -This species is known from People’s Republic of Congo; this is the first record for Cameroon. Eumecopodus longicornis Calabresi Eumecopodus longicornis: Damoiseau, 1967<3:318. Specimens Examined. — CAMEROON. Efulen, 6 Feb 1912, one female (Weber; CMNH). Locality Records.— This species is known from People’s Republic of Congo and Tanzania; this is the first record for Cameroon. Eumecopodus burgeoni De Muizon Eumecopodus burgeoni: Damoiseau, 1967a:320. Specimens Examined. — GABON. Moyen Ogoove: Lambarene, one female (Good, ex coll. Holland; CMNH). Locality Records. — ^This species is known from Republic of Guinea and Peo- ple’s Republic of Congo; this is the first record for Gabon. Phitoderes gestroi Calabresi Phitoderes gestroi: Damoiseau, 1967<3:323; Bartolozzi and Osella, 1990:239. Specimens Examined. — CAMEROON. Sud Province: Lolodorf, Feb 1914, one male (Good; CMNH); Efulen, Sep 1909 and 5 Dec 1913, two females (Weber; CMNH). 276 Annals of Carnegie Museum VOL. 67 Locality Records. — This species is known from Sierra Leone, Ivory Coast, Equatorial Guinea, Gabon, and People’s Republic of Congo; this is the first record for Cameroon. Thoracobrenthus semistriatus Damoiseau Thoracobrenthus semistriatus: Damoiseau, 1967«:326. Specimens Examined. — CAMEROON. Sud Province: Lolodorf, Nov 1914, one specimen {Good; CMNH); Metet, Apr 1918, one specimen (Good; CMNH); without locality, 5 Apr 1910, one specimen (Weber; CMNH). Locality Records. — This species is known from Ivory Coast and People’s Re- public of Congo; this is the first record for Cameroon. Heterothesis elegans Kleine Heterothesis elegans: Damoiseau, 1967a:332; Damoiseau, 1967c: 129. Specimens Examined. — GABON. Ogove R. (= Ogooue River), 12 males (Good, ex coll. Holland; CMNH). Locality Records. — This species is known from Cameroon, People’s Republic of Congo, and Angola; this is the first record for Gabon. Paryphobrenthus latirostris (Gerstacker) Paryphobrenthus latirostris: Damoiseau, 1967a:334; Damoiseau, 1968:171; Sforzi, 1992:5. Specimens Examined. — ANGOLA. Huila distr.: Mulondo (3600 ft), 23 Nov 1930, one male (CMNH). SOUTH AFRICA. NE Transvaal: Ofcolaco, Makhutswe River, Jan 1989, one female (Wer- ner; CLB). Locality Records. — This species is known from Zaire, Sudan, Tanzania, and Mozambique; these are the first records for Angola and South Africa. Tribe Atopobrentini (sensu Damoiseau, 1961 a) Neoceocephalus freyi De Muizon Neoceocephalus freyi: Damoiseau, 1967<3:241; Bartolozzi, 1991:9; Bartolozzi and Sforzi, 1994:251. Specimens Examined. — ZAMBIA. Nmkande, Nov 1973, three specimens (Piacenza; CGO, CLB). Locality Records. — This species is known from Zaire, Kenya, Tanzania, Mo- zambique, and South Africa; this is the first record for Zambia. Neoceocephalus punctatus Damoiseau Neoceocephalus punctatus: Damoiseau, 1967fl:247; Damoiseau, 1972:266. Specimens Examined. — REPUBLIC OF GUINEA. Seredou, at light, 4 Apr 1975, one specimen (Zott; ZMB). Locality Records. — This species is known from Ghana, Cameroon, Gabon, and People’s Republic of Congo; this is the first record for the Republic of Guinea. Neoceocephalus fausti (Senna) Neoceocephalus fausti: Damoiseau, 1967«:251; Damoiseau, 1975:58. Specimens Examined. — REPUBLIC OF GUINEA. Seredou, at light, 4 Apr 1975, two specimens (Zott; ZMB). 1998 Sforzi-=-Brentidae from Tropical Africa 277 Fig. 3.—Plesiobolbus sagax Kolbe: habitus (total length 16.1 mm). 278 Annals of Carnegie Museum VOL. 67 Locality Records. — This species is known from Ivory Coast, Cameroon, Peo- ple’s Republic of Congo, Angola, and South Africa (Transvaal); this is the first record for the Republic of Guinea. Tribe Taphroderini (sensu Damoiseau, 1967fl) Plesiobolbus sagax Kolbe Plesiobolbus sagax: Damoiseau, 1967a:463; Bartolozzi, 1991:26. Specimens Examined. — CAMEROON. Efulen, Nov 1913, one male (Weber; CMNH). Taxonomy and Distribution. — ^This species was briefly described by Kolbe in 1916, but Damoiseau {1961a) questioned the validity of this taxon and even the genus Plesiobolbus Kolbe. His statement was based on the fact that the types of the two species of Plesiobolbus {P. sagax and P. cribriceps Kolbe) could not be found, and Damoiseau never saw any African Taphroderinae with the characters of the genus. Two specimens in the Museum of Tervuren identified as Plesiob- olbus by the well-known German specialist Kleine, were in fact females of the widespread species Anisognathus czikii Bolkay (Damoiseau, 1967a:463). Barto- lozzi (1991) confirmed the validity of the genus Plesiobolbus and described the new species P. martini from Tanzania. In the material of CMNH there is a male specimen of P. sagax, the first to be found after Kolbe’s short description of the species. The specimen agrees well with the short diagnosis of Kolbe, although its length (16.1 mm) is much greater than the type specimen of P. sagax (9.5 mm). It is useful to redescribe and illustrate this very rare and interesting insect. Redescription. — Male: total length 16.1 mm, maximum width (at pronotum) 2.3 mm, antennae 3.9 mm. Body reddish brown with pronotum almost black (Fig. 3). Head longer than broad, with slightly rounded sides, well separated from neck. Occiput slightly projecting posteriorly on neck and bilobed. Temples longer than meta-, meso-, and prorostmm together. Upper surface covered with strong, shining granulation. Rostrum (length 2.4 mm). Metarostrum slightly shorter than distance between eyes, nar- rowing anteriorly; a fovea present between meta- and mesorostrum. Mesorostrum almost as long as metarostrum. Prorostrum as long as distance between eyes, enlarged anteriorly. Upper surface punc- tuated. Anterior border of prorostrum almost straight with very small projection in middle. Mandibles as long as prorostrum, acuminated, left one with tooth in middle. Pronotum as long as head and rostrum together, narrowed anteriorly. Surface smooth, dull, with sparse short golden pubescence; granulation on distal half of pronotal disc. Elytra shorter than pronotum and head together, narrowed posteriorly and narrower than pronotum. Upper surface convex; striae slightly punctuated, interstriae flattened with sparse short golden pubescence; interstriae 1, 2, and 3 elevated on declivity. Base of elytra slightly concave; each elytron apically rounded. Femora laterally compressed at base, unarmed. Underside reddish, shining, with golden pubescence only under rostrum. Two small longitudinal de- pressions laterally on mesorostrum. Small rounded foveola near base of head. Female: unknown. Acknowledgments I thank Dr. Luca Bartolozzi (MZUF) for his precious help in the identification of the material and for his useful suggestions in the preparation of this paper. I am also grateful to Mr. Robert Davidson (CMNH), Dr. Helene Perrin (MNHN), Dr. Roy Danielsson (MZL), Dr. Fritz Hieke (ZMB), Dr. Ole Martin (ZMC), Mr. Bernhard J. van Vondel, Prof. Giuseppe Osella, Prof. Walter Rossi, Mr. Hans Miihle, and Mr. Karl Werner for having allowed me to study the material, and to Dr. Ada Grandinetti (MZUF) for her drawing. I also thank anonymous referees for the revision of the text. Literature Cited Bartolozzi, L. 1991. Secondo contribute alia conoscenza dei Brentidae (Insecta, Coleoptera) deir Africa orientale. Bollettino del Museo regionale di Scienze natural! (Torino), 9:1-31. 1998 Sforzi — Brentidae from Tropical Africa 279 Bartolozzi, L., and G. Osella. 1990. Contribution to the knowledge of Brentidae (Insecta, Cole- optera) of Sierra Leone and Ghana. Accademia Nazionale dei Lincei. Problemi attuali di Scienza e Cultura. Sezione: Missioni ed Esplorazioni. XII. Ricerche Biologiche in Sierra Leone (Parte III) (Roma), 265:233-244. Bartolozzi, L., and A. Sforzi. 1994. Contribution to the knowledge of Brentidae from Kenya (Coleoptera: Brentidae). Koleopterologische Rundschau (Wien), 64:249-256. Damoiseau, R. 1967fl. Monographic des Coleopteres Brentidae du Continent africain. Annales du Musee royal de F Afrique centrale, Serie 8vo (Sciences Zoologiques) (Tervuren), 160:1-507. — . 1961 b. Resultats scientifiques de F Expedition pedo-zoologique hongroise au Congo-Braz- zaville. 15. Brentidae (Coleoptera-Curculionoidea). Bulletin de ITnstitut Royal des Sciences Na- turelles de Belgique (Bruxelles), 43:1-14. . 1967c. Brentidae d’ Angola (Coleoptera-Curculionoidea). Publicagoes culturais Companhia de Diamantes de Angola, Museu do Dundo (Luanda), 77:123-132. — — ^ — . 1968. Coleoptera d’ Afrique Nord-Orientale. Brentidae. Notuiae Entomologicae (Helsingfors), 48:169-174. . 1972. Entomological explorations in Ghana by Dr. S. Endrodi-Younga. 6. Brentidae (Cole- optera). Annales Historico-naturales Musei nationalis Hungarici (Budapest), 64:259-269. — — . 1975. Brentidae d’ Angola (Deuxieme note). Publicagoes Culturais Companhia de Diamantes de Angola, Museu do Dundo (Luanda), 89:53-60. — — “. 1980. Les Amorphocephalini (Coleoptera-Brentidae). 2. Cordus Schoenherr et les genres voisins. Bulletin de ITnstitut Royal des Sciences Naturelles de Belgique (Bruxelles), 52:1-32. Kolbe, H. J. 1916. Beitrage zur Morphologic und Systematik der Taphroderinen (Familie der Bren- tiden) Afrikas. Deutsche Entomologische Zeitschrift (Berlin), 1:50-67. Quentin, R. M. 1966. Contribution a la faune du Congo (Brazzaville). Mission A. Villiers et A. Descarpentries. XXXVL Coleopteres Brenthidae. Bulletin de ITnstitut Fran9ais de FAfrique Noire (serie A) (Abidjan), 28:1631-1670. — — . 1970. Contribution a Fetude des Coleopteres Brentidae. III. Premieres recoltes a la Station d’Ecologie tropicale de Lamto, et aper9u general sur la faune de la Cote d’Ivoire. Annales de FUniversite d’ Abidjan (Serie E, Ecologie), 3:205-222. Sforzi, A. 1992. Contribution to the knowledge of East African Brentidae (Coleoptera). Opuscula Zooiogica Fluminensia (Flumserberg-Grossberg), 93:1-8. Zimmermann, E. C. 1994. Australian Weevils (Coleoptera: Curculionoidea). Volume II. Brentidae, Eurhynchidae, Apionidae and a Chapter on Immature Stages by Brenda May. Commonwealth Scientific and Industrial Research Organization, Melbourne, Australia. itif) A d i> J * ' H‘. '■ . •■-»Ui.»< • . '-r . > sv-L- . < •l •|).,11• k «>k4n ■ ;z* , ’ ■ •' * '-U ‘-t- ■ ♦ i r«%*f .;■' I»> i -u .l#/'<'*»T/ ,t\J •- t. '?*4'3illlWI^ ’i»r "1 w<' /( ' '“ •'V' ¥ ^ jf j*! ,A^yj i*" It » .ir ** I ^ .H..-/'.4,t, > ..'• v,r . . - ' 'l*c. M»'. 3 - ‘i - ., ^ --'‘u ■(»'!'"• < ''^ '*^'d:, ^ ■ .. ,»’Vj.<. t . :iii.:i ■,r pt‘' . ... 1* ‘i'j ^12^ -'v \ V- / •<^A'. V ' -..liv-.'''. .shj^.'A'’*r IP ■.. .'i ■■^/. *’ Vv .'t^it* *'Vi' .Y ■ ,'T»> ' / *iiV^ ♦-••■. *:i5t .•■•‘H i;', 'tVi’ '''..’■<<' .■? iff* .'' '' ^'‘ Vv .fV .'■ •■ ^ »‘' ,i jroPikiS.fc*,^4' ■ ■■^'>* rt'o '^i SP ^ <* '•' ')-''^s’A‘<\ ’^'•'' ' 4TvV# vTr-r ,7-' g "i ‘ f .. 7,. ' TTriiBi' ' ' ‘ '. f .'t- ''..W-f* '-' .'S®. /^1 ■ w A''a.7y^^-»'’s 0 . . , ^'»f*.* . *' ;,L'' V. '?' v . ;V :iiiiQ-'f'l I 'if.-.-r. .V, ^^'■*’7' ' ■.-«r- *. ” 'iti "'-JS- . ' ’>. my v' -t ,, ' ' ■ i ■ - ^ , ■% -ft f ' ' • i -f*’ ' -. .5 ■ -. “>■ I’, .*>'.■ T. t,''' V '* ...» 40 u «- . tf- -• '-^t. -a *C ‘'^41.'*®. . ■' i3t> .« -'■ . - • -^ „ ... ■7' -. I. Hi. ; ■’ •*''• '*'''. *•’..■•* *«r ■ ■’. '■•'■> * fl' li' j,. Pf- ,.Hri ’ -’ ■'’, ''>•■ ' '■^r *. tV v/ ^ j rS. ... , ! :- \g I .J’.v‘.^.rt,\ i^-r- .UlvW,-. Mrf Cll[r,f0 sn ov o\ m IT) r- 00 cn ov VO r- ^ cn o w VO r- VO ^ m VO VO VD N'r^»nA'r^r-' 000000000000000 « ^ a d* o o Q o ffi O *7 D & 308 Annals of Carnegie Museum VOL. 67 Description and Discussion. — The isolated specimen referred here to Petauristodon varies from those of other species of the genus only in its larger size and less prominent crenulations in the talonid basin of the tooth (James, 1963; Lindsay, 1972; Engesser, 1979). If better represented, the Pratt Quarry specimen might well represent a new species. Sciurion Skwara, 1986 cf. Sciurion sp. (Fig. 2J) Referred Specimen. — UNSM 101769, isolated left M, or M2. Measurements. — a-p, 1.42 mm; tra, 1.50 mm; trp, 1.56 mm. Description. — The single lower molar is smaller than all of the other sciurids from Pratt Quarry. It is rhomboid, with no evidence of the anteroposterior compression present in the molars of Ammos- permophilus and Spermophilus. There are faint enamel wrinkles in the talonid and trigonid basins. The metalophulid I runs from the apex of the metaconid along the anterior border of the tooth, ending in a distinct cuspule (= anterostylid) anterior and lingual to the protoconid. The trigonid basin is small and blocked posteriorly by a complete but low metalophulid II. The ectolophid is low and obliquely oriented with just a trace of a mesoconid. The posterolophid is continuous from the hypoconid to the entoconid. The entoconid is distinct and round, not compressed into the lingual end of the postero- lophid as in the spermophiles. A small mesostylid is present posterior to the metaconid. Discussion. — Sciurion campestre was named from the Hemingfordian of Sas- katchewan (Skwara, 1986). The Pratt Quarry specimen differs from the type ma- terial in having a complete metalophulid II and less well-defined mesoconid. UNSM 101769 is also larger than the referred lower molar of S. campestre (Skwa- ra, 1986:table 1). In size, UNSM 101769 is close to the European Blackia. Blackia was originally named for a small “flying squirrel” from the Miocene of Europe (Mein, 1970). UNSM 101769 differs from the described species of Blackia from Europe in having less pronounced wrinkling of the enamel on the cheek teeth, a complete metalophulid II, and a more distinct entoconid, mesostylid, and anter- oconid. The only occurrence of Blackia in North America is based on several isolated teeth from the Hemingfordian of California (Hutchison and Lindsay, 1974) and the medial Barstovian of Nebraska (Voorhies, \990b). The Pratt Quarry specimen is slightly smaller than the Barstovian lower molar from Nebraska (UNSM 85558) and has a complete metalophulid II (trigonid completely open in the Barstovian specimen) and a better developed entoconid, mesoconid, and mesostylid. The Pratt Quarry specimen differs from the Hemingfordian specimens in being larger and the features discussed by Skwara (1986) for S. campestre. Family Castoridae Gray, 1821 Subfamily Castoroidinae Trouessart, 1880 Dipoides Jager, 1835 Dipoides tanneri, new species (Fig. 3, 4, 5A-C, Table 4) Type Specimen. — UNSM 101612, little worn P4. Referred UNSM 101613-101620, P4; UNSM 101621-101631, P^; UNSM 101632- 101653, 101795, 101798, FAM 64483-64487, isolated molars; UNSM 101654-101657, isolated in- cisors; UNSM 101730, distal humerus; UNSM 101731, proximal femur; FAM 64488, 64517, man- dibles with P4-M2; FAM 64482, complete skull with associated mandible and postcranial elements. Diagnosis. — Similar in size to D. stirtoni; differs from all other species of the genus in the lower crown height of the cheek teeth (premolars develop roots in 1998 Korth — Late Clarendon Rodents and Lagomorphs 309 very late stages of wear), upper premolars not attaining the S-pattern of the oc- clusal surface, and the lower premolar having a parastriid shorter than the me- sostriid (parastriid not reaching the base of the tooth) as in D. stirtoni, and a small metafossettid retained until very late stages of wear (absent in other species). Etymology. — Patronym for Lloyd Tanner, in recognition of his many years of work for the UNSM. Description. — The nearly complete skull with heavily worn dentition (FAM 64482) lacks only parts of the zygomatic arches. Due to the advanced age of the individual, some of the cranial sutures are not distinguishable. Also, damage to the pterygoid and basicranial area has obscured the features and foramina of this area. In general shape, it is not as deep dorsoventrally as the skull of Recent Castor, and the rostrum is relatively longer. Wahlert (1972) fully described the skull of a Hemphillian Dipoides and noted only four differences in the cranial foramina between Dipoides and earlier Eucastor and Monosaulax: 1) posterior palatine foramina medial to M’ in Dipoides, medial to the boundary of M’ and in the other genera; 2) lateral margin of the infraorbital foramen forms part of the masseteric tubercle in Dipoides but continues into the tubercle in the other genera; 3) the sphenopalatine foramen is dorsal to the premolar in Dipoides and more posterior in the other genera; and 4) the ethmoid foramen is entirely within the frontal bone in Dipoides and passes through the frontal-orbitosphenoid suture in the other genera. Nearly complete skulls of E. tortus from the Valentine Formation of Nebraska (UNSM 85600) and M. pansus from New Mexico (FAM 64945) were compared with FAM 64482. All of the differences cited by Wahlert (1972) hold true for the skull of D. tanneri except for the morphology of the infraorbital foramen. In the skull of D. tanneri the foramen continues into the masseteric tubercle as in Eucastor and Monosaulax. Other than these differences, there is little dis- tinction between the skulls of these three beavers. One other feature of the cranial foramina of the Hemphillian Dipoides figured by Wahlert (1972: fig. 16) is the presence of three interorbital foramina posterior and ventral to the optic foramen. In the specimens of Eucastor and Monosaulax at hand, there are only two interorbital foramina. The skull of D. tanneri has three foramina as in the Hemphillian Dipoides. In size, the skull of D. tanneri is at least 50% larger than the skulls of the other two beavers. The molars of D. tanneri are hypsodont and rootless. The occlusal patterns of both the upper and lower molars quickly wear to the characteristic S -pattern of Dipoides. The flexi remain open and do not close to form fossettes (-ids) until the latest stages of wear. M^ is distinct from the anterior molars. It does not taper towards the base of the crown. On the buccal side of the tooth, there is only one persistent stria, the mesostria. The parastria is much shorter, and a parafossette is formed after only moderate wear. The hypostria is continuous to the base of the crown and the hypoflexus is as in the anterior molars. In late stages of wear the parafossette is lost and the hypoflexus extends to the buccal margin of the tooth. On little-wom specimens there is a short metastria. The metaflexus and mesoflexus isolate the metacone. After the metastria has been worn away, the metacone remains isolated. The mesoflexus extends to the lingual margin of the tooth and is strongly concave posteriorly. The premolars of D. tanneri develop roots only in the very late stages of wear. On the lower premolars, the buccal hypostriid and lingual mesostriid are continuous to the base of the crown, thus the hypoflexid and mesoflexid remain open throughout the life of the individual. The parastriid is slightly shorter than the mesostriid, ending only a few millimeters from the base of the crown. Because of the length of the parastriid, in very late stages of wear, the paraflexid closes lingually to form a parafossettid. In occlusal view, the paraflexid, mesoflexid, and hypoflexid are all concave anteriorly and cross nearly the entire width of the tooth. The only variation is the presence of a small metafos- settid at the lingual end of the hypoflexid. This metafossettid is present on all specimens except those with extreme wear. The upper premolars are square in occlusal outline and the crown is strongly curved (concave buccally). On the lingual side of the tooth, the hypostria is continuous to the base of the crown. On the buccal side, the parastria and mesostria extend to within 2-A mm of the base of the crown, but the metastria extends less than half of the height of the crown toward the base of the crown. On the occlusal surface of the tooth, the hypoflexus and paraflexus meet on the buccal half of the tooth, being separated only by a thin enamel wall. The mesoflexus extends to the lingual border of the tooth. The metaflexus is only about half the tooth in length and closes off buccally after only moderate wear to form a small fossette. All of the flexi on are concave posteriorly. Discussion. — Dipoides tanneri is referable to this genus based on Stirton’s (1935:440) diagnosis — high-crowned cheek teeth with striae (-ids) persistent to the base of the crowns. The skull of D. tanneri also has the distinctive features 310 Annals of Carnegie Museum VOL. 67 1998 Korth — Late Clarendon Rodents and Lagomorphs 311 of Dipoides, not present in Eucastor (posterior palatine foramina more anterior, ethmoid foramen entirely within the frontal bone, three interorbital foramina pres- ent posterior and ventral to the optic foramen, and the sphenopalatine foramen more anteriorly positioned). Dipoides tanneri is clearly distinguishable from contemporaneous species of Eucastor because of these differences. The Clarendonian Eucastor malheurensis is similar to D. tanneri in crown height of the cheek teeth, but is much smaller than D, tanneri (Shotwell and Russell, 1963:table 8) and the striae (-ids) of the cheek teeth do not extend as far toward the bases of the crowns, thus forming fossettes (-ids) earlier in wear. Two species of Eucastor have been identified from the Clarendonian of the Great Plains, E. dividerus from Nebraska (Webb, 1969a) and E. philisi from Kansas (Wilson, 1968). Both of these species are similar in size to D. tanneri but do not have the persistent striae (-ids) of the cheek teeth of D. tanneri (Stirton, 1935:fig. 107-110; Wilson, 1968:texLfig. 15). Dipoides tanneri differs from all other species of Dipoides in having lower- crowned cheek teeth, maintaining a metafossettid on P4 until late stages of wear, having roots develop on the premolars in late stages of wear, having and not attain the S -pattern of the other molars, and having the parastriid shorter than the mesostriid on P4. All of these features are viewed as primitive and are shared with species of Eucastor. Among species of Dipoides, D. tanneri most closely resembles D. stirtoni from the early Hemphillian of Oregon (Wilson, 1934). Dipoides stirtoni, like D. tanneri has a P4 with a shorter parastriid than mesostriid. In all other species both of these striids extend to the base of the crown. Similarly, P"^ and of D. stirtoni do not wear to the occlusal S-pattem characteristic of Dipoides. Dipoides tanneri differs from D. stirtoni in having lower-crowned cheek teeth and developing roots on the premolars. The postcranial material assigned to D. tanneri differs from that of D. stirtoni only in being slightly smaller and less robust (Wilson, 1934: fig. 1). Dipoides tanneri is the earliest occurrence of the genus. All other species of Dipoides are known only from the Hemphillian and Blancan (Korth, 1994:148). Tliis earlier occurrence is compatible with the more primitive features of the dentition of D. tanneri. Eucastor Leidy, 1858 Eucastor planus Stirton, 1935 (Fig. 5D-F, Table 5) Referred Specimens. — UNSM 101594, partial mandible with incisor and P4-M1; UNSM 101610, b; UNSM 101611, P4; UNSM 101596, 101600, 101602, 101605, 101607-101609, 101611, 101658, Mj or M2; UNSM 101603, M3; UNSM 101597, 101601, 101604, Pf UNSM 101595, 101606, 101794, Ml or Mb UNSM 101598, 101599, Mb Discussion.—The specimens referred here to Eucastor planus do not differ from the holotype from the Clarendonian of Oklahoma (Stirton, 1935) or other <— Fig. 3. — Skull of Dipoides tanneri, FAM 64482. A, lateral view (zygoma removed). B, ventral view. C, dorsal view. Abbreviations of foramina: eth, ethmoid; iof, infraorbital; in, incisive; ito, interorbital; op, optic; pgl, postglenoid; ppl, posterior palatine; spl, sphenopalatine. Bar scale = 1 cm. Dashed lines indicate probable orientation of sutures. 312 Annals of Carnegie Museum VOL. 67 Fig. 4. — Unworn premolars of Dipoides tanneri. A, UNSM 101612, holotype, right P4; occlusal view (above), buccal view (left), and lingual view (right). B, UNSM 101628, right P-^; occlusal view (above), buccal view (left), and lingual view (right). Bar scale = 5 mm. 1998 Korth — Late Clarendon Rodents and Lagomorphs 313 Table 4. — Dental measurements o/’Dipoides tanneri. Abbreviations as in Table 1. Crown height mea- sured only on unworn or little worn lower premolars. Measurements in mm. Isolated premolars n M or 5 cv P4 a-p 9 7.03 6.48-7.86 0.50 7.1 tra 8 3.92 3.49-4.55 0.40 10.2 trp 9 4.69 3.98-5.38 0.41 8.7 crown height 5 14.20 13.40-15.60 0.85 6.0 p4 a-p 11 4.47 3.92-5.01 0.29 6.5 tr 11 5.22 4.70-5.59 0.29 5.6 Teeth in jaws FAM FAM No. No. 64482 64488 64517 64482 (Left) (Right) P4 a-p 6.28 6.10 6.39 P”^ a-p 6.10 6.31 tra 4.41 4.10 4.40 tr 5.78 5.52 trp 5.02 5.35 5.21 M, a-p 4.59 4.14 4.30 M* a-p 3.90 3.81 tr 4.80 5.09 4.88 tr 4.58 5.15 M2 a-p 3.40 3.82 4.25 M^ a-p 3.87 3.80 tr 4.39 5.01 4.64 tr 4.99 5.20 M3 a-p 4.76 M^ a-p 4.49 4.25 tr 4.00 tr 3.93 4.29 P4-M3 19.79 20.10 19.88 p4_M3 19.22 18.37 referred specimens of this species. This species is clearly separable from the other castorids from Pratt Quarry by its markedly smaller size, having higher-crowned cheek teeth than Hystricops, and having less well-developed striae (-ids) on the cheek teeth than in Dipoides. Eucastor planus has previously been identified in the early Clarendonian Burge and medial Clarendonian Minnechaduza faunas of northcentral Nebraska as well (Webb, 1969a; Voorhies, 1990a). Subfamily uncertain Hystricops Leidy, 1858 Hystricops venustus Leidy, 1858 (Fig. 6, Table 6) Hystrix {Hystricops) venustus Leidy, 1858. Hystrix venustus Leidy, 1869. Erethizon venustus (Leidy) Hay, 1901. Hystricops venustus Leidy: Stirton, 1935. Referred Specimens.— 101660, P4; UNSM 101661, 101662, 101775, M, or M2; UNSM 101663, 101664, 101667, M3; UNSM 101655, 101666, P^; UNSM 101668, 101669, upper molars; UNSM 101747, incisor fragment. Description. — The cheek teeth are mesodont and strongly rooted, much lower crowned than the other castorids from Pratt Quarry, and clearly much larger in size (Table 6). One incisor fragment is preserved, UNSM 101747. It has a broad, gently convex, and smooth anterior surface. The lower premolar is the largest of the cheek teeth. It has two roots. The only striids that are continuous to near the base of the crown are the hypostriid and the mesostriid. All others are very short, disappearing after only moderate wear. The mesoflexid is the longest of the flexids, extending nearly to the buccal edge of the tooth. The mesoflexid is gently curved anteriorly. The hypoflexid is much shorter, extending only about one-fourth the width of the tooth, posterior to the mesoflexid, in 314 Annals of Carnegie Museum VOL. 67 Fig. 5. — Cheek teeth and mandibles of castorids from Pratt Quarry. A-C, Dipoides tanneri FAM 64482. A, left B, right P4-M3. C, lateral view of mandible. D-F, Eucastor planus. D, UNSM 101594, left P4-M,. E, UNSM 101609, left P''. F, UNSM 101594, lateral view of mandible. Bar scale for teeth (above) = 5 mm. Bar scale for mandibles (below) = 1 cm. 1998 Korth — Late Clarendon Rodents and Lagomorphs 315 Table 5. — Dental measurements o/Eucastor planus from Pratt Quarry. Abbreviations as in Table 1. Measurements in mm. n M OR S cv P4 a-p 2 4.40 4.27-4.52 tra 2 2.84 2.70-2.98 trp 2 3.47 3.34-3.60 Ml or M2 a-p 8 3.10 2.72-3.51 0.32 10.2 tr 8 3.74 3.02-4.09 0.42 11.6 M3 a-p 1 2.50 tr 1 2.89 p4 a-p 3 3.64 3.51-3.80 0.15 4.0 tr 3 4.01 3.46-4.36 0.48 12.0 M> or M2 a-p 2 3.09 3.00-3.18 tr 2 3.60 3.50-3.60 M3 a-p 2 2.74 2.67-2.80 tr 2 2.63 2.41-2.85 a posterolingual direction. On the anterior half of the tooth is a large, anteriorly concave parafossettid. Irregularities of the enamel are preserved along its borders. Anterior to the parafossettid is a much smaller fossettid that is straight and oriented transversely. Along the lingual border of the tooth between the mesoflexid and the parafossettid is a short flexid that is oriented posterobuccally. The associated striid on the lingual side of the tooth is very short (approximately one-eighth the remaining crown height) and would soon disappear after only a little more wear. The posterior half of the tooth is dominated by a transversely elongated hypofossettid. There are two bends in the hypofossettid, giving it a zig-zag shape. Posterior and lingual to the hypofossettid is a minute fossettid that is oriented obliquely. The anterior lower molars are three-rooted, with one posterior and two smaller anterior roots. They are nearly square in occlusal outline. Only one flexid, the hypoflexid, remains open to near the base of the crown. On all other specimens the only lingual flexid is the mesoflexid on one specimen (UNSM 101775). There is no evidence of the parafossettid or metafossettid communication with the lingual margin of the tooth on any of the available specimens. The parafossettid is transversely elongated and the only fossettid on the anterior half of the tooth. It has several subtle bends in it, giving it an irregular shape. In one specimen, UNSM 101661, the buccal end of the parafossettid is separated from the rest of the fossettid, and forms a small, obliquely oriented fossettid. The mesofossettid is straight and extends a little over half of the width of the tooth. It is oriented slightly anterobuccally. The hypoflexid extends about half the width of the tooth and is posterolingually oriented, paralleling the mesofossettid. A short metafossettid is the only feature of the posterior half of the tooth. It is mostly parallel with the mesofossettid, but has a bend near its buccal end, similar to the metafossettid in P4. The last lower molar is essentially identical to the anterior molars except it is narrower and longer. P^ is the largest of the upper cheek teeth. The only persistent flexus is the hypoflexus, the hypostria extending nearly to the base of the crown. On UNSM 101666 the mesoflexus has remained open, but the mesostria extends less than one-fourth of the remainder of the crown height, indicating that the flexus would close after only a little more wear. There is no indication of a parastria or metastria on either of the available specimens. The hypoflexus is oriented in an anterobuccal direction and extends about one-third the width of the tooth. Its buccal end abuts the lingual end of the parafossette. The parafossette follows the direction of the hypofossette, is transversely elongated, and retains some minor irregularities along its enamel outline. On UNSM 101666 there is a minute enamel fossette anterior to the parafossette. It appears that this accessory fossette would erode after only a little more wear. The mesofossette (or mesoflexus) extends nearly the entire width of the tooth and curves posteriorly, ending along the posterior margin of the tooth. The metafossette is short and obliquely oriented. The upper molars are similar to P"^ in occlusal pattern but smaller in size, being greatly shortened anteroposteriorly. There is no evidence of buccal striae on any of the referred upper molars. The mesofossette is long, posteriorly curved, and meets the posterior wall of the tooth. The parafossette and metafossette are small and placed as in P"^. The hypoflexus, again, is oriented as in P'^. Discussion. — The type specimen of Hystricops venustus is an isolated lower premolar and lower molar, USNM 1180, from the ‘'Niobrara River fauna” of northcentral Nebraska (Leidy, 1858; Stirton, 1935). The single P4 from Pratt Quar- 316 Annals of Carnegie Museum VOL. 67 Fig. 6. — Cheek teeth of Hystricops venustus. A, UNSM 101660, right P4. B, holotype, USNM 1180, left P4. C, UNSM 101775, right M, or M2. D, UNSM 101661, right M, or M2. E, UNSM 101667, left M3. F, UNSM 101666, left Ph G, UNSM 101665, right Ph H, UNSM 101668, right M* or M^. Bar scale = 5 mm. 1998 Korth — Late Clarendon Rodents and Lagomorphs 317 Table 6. — Dental measurements of Hystricops venustus from Pratt Quarry. Abbreviations as in Table 1. Measurements in mm. Isolated Cheek Teeth UNSM P4 M| or M 2 M3 p M' No. a-p tra trp a-p tra trp a-p tra trp a-p tra trp a-p tr 101660 101661 1L90 7.65 9.11 7.59 8.14 8.33 101662 8.00 8.00 7.33 101775 101663 7.45 8.01 7.14 7.35 6.51 6.16 101664 — 5.80 5.84 101667 101665 5.86 5.80 5.68 8.50 9.39 8.82 101666 101668 8.42 9.42 8.65 6.32 6.51 101669 5.74 7.00 ry (UNSM 101660) is similar in size to the holotype premolar and differs only in the presence of a small reentrant valley on the lingual side of the tooth anterior to the mesoflexid and the minute accessory fossettid lingual to the metafossettid. It appears that this accessory valley and the accessory fossettid would be worn away after only slightly more wear. This is evident on the short flexid because there is no striid on the lingual side of the tooth continuous with it. The remainder of the occlusal morphology is nearly identical with the holotype. The lower molars referred here to H. venustus have all of the lingual flexids closed early in wear, and are nearly identical to the isolated molar included in the holotype of H. venustus (Stirton, 1935:fig. 60). The reference of the Pratt Quarry material to H. venustus is almost certain. There are no known castorids of this size and morphology reported from the lower horizons (medial to late Barstovian) in the Niobrara River valley of northcentral Nebraska, even though they have been extensively collected and described (Voorhies, 1990^). The morphology of the upper premolars is also nearly identical to the unnamed species of Hystricops reported from the Clarendonian of Oregon (Shotwell and Russell, 1963). The Hemphillian H. browni from Oregon (Shotwell, 1963) is also similar in occlusal morphology to the upper premolars referred to H. venustus from Pratt Quarry, but differs in the depth of the striae. This similarity verifies the assignment of these later Tertiary beavers to Hystricops. There is also a great similarity in the morphology of the cheek teeth of H. venustus to those of an unnamed castorid from the Hemingfordian of Colorado referred by Wilson (1960) to lAnchitheriomys sp. (KU 10173). It appears that the Colorado specimen is also referable to Hystricops rather than Anchitheriomys. Some of the skull characters of the Colorado skull also differ from those of Anchitheriomys (Korth and Emry, 1997). Dorsally, the nasals extend farther pos- teriorly than the premaxillary bones, a character only otherwise present in Ag- notocastor and Neatocastor among castorids (Korth, 1996a). Likewise, the ros- trum is more elongated than in typical castorids, another feature of Agnotocastor and Neatocastor. Besides being referable to Hystricops, the Hemingfordian skull from Colorado appears to relate this genus to the Agnotocastorinae (Korth, 1996a; Korth and 318 Annals of Carnegie Museum VOL. 67 Emry, 1997). However, a detailed description and study of the Colorado specimen is necessary before such an allocation can be made. Family Eomyidae Deperet and Douxami, 1902 Pseudotheridomys Schlosser, 1926 cf. Pseudotheridomys sp. (Fig. 8A) Referred Specimen. — UNSM 101748, right mandible with fragment of lower incisor and alveoli for all cheek teeth. Description. — The mandible is small (alveolar length of cheek teeth = 3.5 mm) and nearly complete, lacking only the base and the posterior processes (coronoid, condyle, and angle). There are alveoli for four cheek teeth: the premolar was two-rooted and the molars three-rooted (one posterior, two anterior). The masseteric scar ends anteriorly below the anterior root of the premolar in a V-shape. On the ascending ramus, the base of the incisor is a lateral bulge that tapers posteriorly to a small ridge. The diastema is shorter than the tooth row and concave. The mental foramen is near the dorso ventral center of the mandible, below the center of the diastema. Discussion. — Only three genera of eomyids persist in North America into the Barstovian and later times — Leptodontomys, Kansasimys, and Pseudotheridomys (Korth, 1994:161). Of these only Leptodontomys and Pseudotheridomys are sim- ilar in size to UNSM 101748 from Pratt Quarry. UNSM 101748 most closely resembles the mandible of Pseudotheridomys because of the more anterior extent of the masseteric scar, shorter diastema, morphology of the base of the incisor on the ascending ramus (low, shelf-like structure on Leptodontomys), and the align- ment of the alveolar margin of the cheek teeth with the alveolus of the incisor. Engesser (1979:fig. 7a) described the mandible of Leptodontomys in detail and noted that the alveolar margin of the cheek teeth generated a line that, if extended anteriorly, would pass the incisor well above its alveolus. In UNSM 101748 and other species of Pseudotheridomys this line would be level with the incisor al- veolus. In size, UNSM 101748 is nearly identical to P. pagei from the Barstovian of Oregon (Shotwell, 1967^). However, the lack of cheek teeth does not allow a for a definite specific identification of the Pratt Quarry specimen. Previously, the last reported occurrence of Pseudotheridomys in North America was the Barstovian (Engesser, 1979; Fahlbusch, 1979). The recognition of this genus in the late Clarendonian greatly extends its record in North America. In Europe, Pseudotheridomys disappears even earlier in the fossil record (Fahlbusch, 1979). Because UNSM 191748 was recovered from the base of the channel at Pratt Quarry, it is possible that it is reworked from the Barstovian layers of the Valentine Formation below the channel. However, there is no previous record of Pseudotheridomys from the Valentine Formation (Klingener, 1968; Korth, 1979; Voorhies, \990b). Family Heteromyidae Gray, 1868 Subfamily Mioheteromyinae Korth, 1997 Mioheteromys Korth, 1997 Mioheteromys sp., cf. M. agrarius (Wood, 1935) (Fig. 7A, 8B) Referred Specimens. — UNSM 101750, isolated P'*; UNSM 101573, partial mandible with lower incisor. Measurements. — UNSM 101750, a-p, 1.67 mm; tra 0.96 mm; trp, 1.56 mm. 1998 Korth — Late Clarendon Rodents and Lagomorphs 319 Fig. 7. — Scanning electron micrographs of cheek teeth of heteromyid rodents from Pratt Quarry. A, Mioheteromys sp., cf. M. agrarius, right UNSM 101750. B, C, Lignimus sp. B, right M* or M^, UNSM 101576. C, left M, or M2, UNSM 101531. D-G, Cupidinimus prattensis. D, left P^-M’, UNSM 101502. E, Holotype, left P4 (occlusal view above, lingual view below), UNSM 101501. F, left Mj or M2 (occlusal view above, buccal view below), UNSM 101512. G, right M‘ or M^ (occlusal view above, lingual view below), UNSM 101510. Bar scale = 1 mm. Discussion. — A single isolated heteromyid is distinct from the Cupidinimus specimens from Pratt Quarry because of its larger size and lower crown height. The union of the protocone to the metaloph is also lingual rather than central as in Cupidinimus. All of these distinctive characters are typical of Mioheteromys. The size of the specimen is comparable with P'^s of M. agrarius from the early and middle Clarendonian of Nebraska (Korth, 1997). 320 Annals of Carnegie Museum VOL. 67 Fig. 8. — Mandibles of geomyoids from Pratt Quarry. All lateral views. A, cf. Pseudotheridomys sp., UNSM 101748. B, Mioheteromys sp., cf. M. agrahus. C, Cupidinimus prattensis, holotype, UNSM 101501. D, Lignimus sp., UNSM 101532. Bar scale = 5 mm. 1998 Korth — Late Clarendon Rodents and Lagomorphs 321 Table 7. — Dental measurements of Cupidinimus prattensis. Abbreviations as in Table L Additional abbreviations (crown height variables from Barnosky, 1986a): CHEV, height of enamel chevron on lingual side of upper molars; LEH, buccal enamel height of lower molars; LEHP, buccal enamel height of lower premolar; T, maximum transverse width; TOTH, crown height of lingual side of upper molars; TP, transverse width of P 4. n M OR s cv P4 a-p 1 0.91 tra 1 0.84 trp 1 0.91 LEHP 1 0.48 LEHP/TP 1 0.53 M, or M2 a=p 4 1.15 0.98-1.26 0.12 10.4 tra 4 1.44 1.33-1.59 0.11 7.7 trp 4 1.34 1.24-1.49 0.11 8.1 LEH 3 0.65 0.54-0.73 0.10 16.1 LEU/T 3 0.44 0.38-0.47 0.05 11.3 p4 a-p 11 1.41 1.28-1.55 0.10 7.0 tra 11 0.75 0.67-0.83 0.04 6.0 trp 11 1.33 1.20-1.43 0.08 6.2 M' or M2 a-p 7 1.06 0.95-1.24 0.10 9.8 tra 7 1.32 1.16-1.43 0.10 7.7 trp 7 1.26 1.15-1.34 0.07 5.5 CHEV 7 0.19 0.13-0.28 0.06 30.4 TOTH 6 0.88 0.73-0.97 0.09 10.7 CHEV/T 7 0.14 0.09-0.23 0.05 33.9 TOTH/T 6 0.64 0.53-0.70 0.06 9.7 The partial mandible preserves the alveoli for P4 and Mj only. The rest of the mandible posterior to Mj is missing. It does not differ from the mandible of M. agrarius described previously (Wood, 1935; Korth, 1997). Subfamily Dipodomyinae Gervais, 1853 Cupidinimus Wood, 1935 Cupidinimus prattensis, new species (Fig. 7D~G, 8C; Table 7) Type Specimen. — -UNSM 101501, right mandible with Ii and P4. Referred Specimens. — UNSM 101502, maxilla with UNSM 101528, edentulous maxilla; UNSM 101503-101509, 101513, 101519, 101521, isolated R; UNSM 101510, 101511, 101514, 101515, 101520, isolated M^ or M^; UNSM 101512, 101517, 101529, 101530, isolated Mi or M2; UNSM 101522-101527, mandibles lacking cheek teeth. Diagnosis. — Intermediate sized, smaller than C. kleinfelderi, C. avawatzensis, C. quartus, and C. bidahocMensis, larger than C. eurekensis and C iindsayi; lower crowned than C. nebraskensis, C. whitlocki, and C. halli, higher crowned than C kleinfelderi, C. boronensis and C smaragdinus with better-developed enamel chevrons on upper molars; also differs from C. whitlocki in having a relatively longer P^; differs from C saskatchewanensis in lacking the central anteroposterior loph on P4; differs from C. tertius and C. cuyamensis in lacking accessory cus- pules on protoloph of P"^; differs from C. nebraskensis and C. madisonensis in having as wide as P^ rather than having P^ wider than Mb Etymology. — Latin, -erisis, suffix meaning from; allusion to Pratt Quarry. Description. — The only known P4 of C. prattensis is in the holotype. It is simple in occlusal mor- 322 Annals of Carnegie Museum VOL. 67 phology, consisting of only four cusps with no accessory cuspules. The crown-height index of the specimen (LEHP/TP) is 0.53, intermediate among species of Cupidinimus (Bamosky, 1986i2:fig. 12). The cusps of the metalophid (protostylid, metaconid) are nearly equal in size and essentially round in outline. The hypolophid cusps (hypoconid, entoconid) are also equal in size but are oval in outline, the long axis being transversely oriented. The two lophs fuse at the center of the tooth. The upper premolar on the only maxillary specimen (UNSM 101502) has a transverse width equal to that of M‘. P"^ is longer (anteroposteriorly) than wide in nearly all specimens. The protoloph is made of a single round to oval protocone. None of the specimens of P'* have any accessory cuspules on the protocone. The metaloph of P'^ is made of three aligned cusps as is typical for the genus. The fusion of the lophs is central. The occlusal morphology of the molars, again, is typical of the genus, consisting of two rows of three cusps. On the lower molars, the anterior cingulum originates at the protoconid and continues buccally to the protostylid. The anterior cingulum on the upper molars originates at the paracone, passes anterior to the protocone, and ends lingually at the protostyle. The crown height index for the lower molars (LEH/T) averages 0.44, higher than in C. madisonensis and C. lindsayi, but lower than all the other species measured by Barnosky (1986fl:fig. 10). The upper molars have a crown height index (TOTH/T) that averages 0.64, lower crowned than all the species measured by Bamosky (1986fl: fig. 13). The index of the height of the enamel chevron (CHEV/T) on the upper molars of C. prattensis is also lower than other species previously measured (Barnosky, 1986fl;fig. 11). Discussion. — Cupidinimus prattensis has relatively low-crowned cheek teeth, exceeding the crown height of only a few species either not measured or unknown to the last reviewer of the genus Bamosky (1986a): C boronensis, C. kleinfelderi, and C. smaragdinus (Whistler, 1984; Korth, 1996Z?). The lack of accessory cus- pules on either the upper or lower premolars of C. prattensis is elsewhere con- sistent only with specimens of C lindsayi, which is smaller and higher crowned than C. prattensis (Bamosky, 1986a). Among its contemporary species of Cupidinimus (C. quartus, C. tertius, C. cuyamensis, and C. avawatzensis), C. prattensis has lower-crowned cheek teeth. It also lacks the accessory cuspules on or P4 of these species. Cupidinimus prattensis is also smaller than C. quartus and C avawatzensis. The unnamed species of Cupidinimus from the Clarendonian Mission fauna of South Dakota is smaller than C. prattensis and the figured P4 appears to have an anterostylid (Green, 1971:text-fig. 2D). As noted by Bamosky (1986a), the species of Cupidinimus from the Great Plains typically have lower-crowned cheek teeth than those from the Rocky Mountains and farther west. Cupidinimus prattensis is clearly part of this more eastern radiation of Cupidinimus. The remainder of the species in this eastern radiation (C. nebraskensis, C. kleinfelderi, and C. smaragdinus) are also charac- terized by conamonly having accessory cuspules on the premolars, especially P4 (Storer, 1975; Korth, 1979, I996b\ Bamosky, 1986a). Cupidinimus prattensis dif- fers from these species in this regard, lacking accessory cuspules on either upper or lower premolars. Cupidinimus prattensis is similar in size and occurrence of accessory cuspules on the premolars to the Barstovian C. whitlocki but differs from the latter in having slightly lower-crowned cheek teeth and having P'^ longer than it is wide. Cupidinimus whitlocki is diagnosed as having a shortened upper premolar (Barnosky, 1986a). The recognition of Cupidinimus prattensis increases the number of Clarendon- ian species of this genus to five (see Korth, 1994:186). This is a decrease from the nine Barstovian species, and part of the trend toward the diminishing numbers of species of this genus into the Hemphillian, where only a single species is recognized (Baskin, 1979). 1998 Korth — Late Clarendon Rodents and Lagomorphs 323 ?Subfamily Harrymyinae Wahlert, 1991 Lignimus Storer, 1970 Lignimus sp. (Fig. 7B, C, 8D) Referred Specimens.— V^SM 101531, isolated M, or Mj; UNSM 101516, 101533, isolated M' or M^; and UNSM 101532, partial edentulous mandible. Measurements. — Lower molar (UNSM 101531): a-p, 1.22 mm; tra, 1.32 mm; tip, 1.35 mm. Upper molars: UNSM 101516, a-p, 1.19 mm; tra, 1.43 mm; tip, 1.39 mm; UNSM 101533, a-p, 1.26 mm; tra, 1.75 mm; trp, 1.71 mm. Description. — The cheek teeth are nearly identical in morphology to those of Lignimus montis and L. austridakotensis except in size. The mandible referred to Lignimus is robust for a heteromyid. The diastema is shallow and long. The masseteric scar has a strong ventral ridge that extends to a point just anterior to the roots of P4, near the center of the mandible. At its anterior end it curves posteriorly. There is a very faint ridge marking the dorsal margin of the masseteric fossa that disappears below the alveolus for Mj. The mental foramen is small and directly anterior to the end of the masseteric ridge. Discussion.— specimens referred here to Lignimus are intermediate in size between L. montis and L. austridakotensis from the Great Plains (Storer, 1975; Korth, 1996b). The cheek teeth are not transversely elongated as in L. transversus from Wyoming (Bamosky, 1986^). The mandible of Lignimus has never before been described. The allocation of the Pratt Quarry specimen to Lignimus, even though it lacks cheek teeth, is based on its comparable size with the cheek teeth identified as Lignimus and a lack of any other geomyoid in the fauna to which it could be allocated. The most intrigu- ing character of the mandible is the masseteric scar. It is not typical of heteromyids with a small shelf that extends anterior to the tooth row, but is nearly identical to the masseteric scar on the mandible of the harrymyine Harrymys irvini (Wah- lert, 1991 :fig. 1). This similarity of the mandible reinforces the suggestion that Lignimus belongs in the Harrymyinae rather than any other group of geomyoid. Lignimus is one of several species present at Pratt Quarry that is known else- where only from much earlier horizons (early to middle Barstovian) and thus these specimens might simply be reworked from the lower horizons present at Pratt Quarry. Once again, this is not likely because Lignimus specimens are known from the fossiliferous horizon 10 m above the basal channel and are not conspe- cific with any of the named species of the genus from the Barstovian. Family ?Geomyidae Bonaparte, 1845 Phelosaccomys Korth and Reynolds, 1994 Phelosaccomys hibbardi (Storer, 1973) (Fig. 9, Table 8) Lignimus hibbardi Storer, 1973. Parapliosaccomys hibbardi (Storer) Korth, 1987. Phelosaccomys hibbardi (Storer) Korth and Reynolds, 1994. Referred Specimens.— WSM 101561, 101773, P4; UNSM 101577, isolated dP4; UNSM 101562, 101570, isolated Fs; UNSM 101563-101569, 101571, 101574-101576, isolated molars; UNSM 101572, isolated upper incisor. Description. — The dP4 referred to P. hibbardi is low crowned and the roots are splayed. The tooth is much longer than any P4 referred to the species. The hypolophid is made of four cusps that fuse together to form a wide loph. The largest cusps recognizable on the hypolophid are the hypoconid and entoconid, the former being the largest. A small hypoconulid is present posterior to the junction of the two main cusps. Buccal to the hypoconid is a small hypostylid. Anterior to the entoconid is a deep valley that separates the entoconid from the large, round metaconid. The remainder of the tooth 324 Annals of Carnegie Museum VOL. 67 Fig. 9. — Scanning electron micrographs of cheek teeth of Phelosaccomys hibbardi from Pratt Quarry. A, left dP4, UNSM 101577. B, left P4 (occlusal view above, lingual view below), UNSM 101773. C, 1998 Korth — Late Clarendon Rodents and Lagomorphs 325 is surrounded by a series of seven small cuspules that run along the buccal and anterior sides of the tooth and on the lingual side anterior to the metaconid. A small triangular cusp is located just anterior to the hypoconid in the center of the tooth. Of the two P4S represented from Pratt Quarry, one is unerupted (UNSM 101561) and the other is relatively well worn (UNSM 10173). On the unempted tooth all cusps are visible. The metaconid and protostylid are nearly equal in size. There are two small anterostylids present, the lingual one is attached to the metaconid. Together these four cusps form an anteriorly convex lophid, isolating a central basin. The hypolophid is straight and made of two major cusps (entoconid, hypoconid) and a smaller, central hypoconulid. On the worn tooth the cusps have been lost on the occlusal surface and only a small enamel lake is present near the center of the tooth. On the sides of the unworn specimen there is a distinct dentine tract. The ratio of the dentine tract height (measured on the buccal side) to the posterior width of the tooth is 1.41. There are no roots evident on the worn P4. As with the lower premolars, of the two P'^s referred here to P. hibbardi, one is essentially unworn (UNSM 101570) and the other is well worn (UNSM 101562). On the unworn specimen there are three cusps of the metaloph that form a posteriorly convex arc (metacone, hypocone, hypostyle). The protoloph is narrower (buccolingually) than the metaloph and straight. It is made of two cusps, a larger protocone and a smaller, lingual protostyle. The lophs unite lingually. On the worn tooth the cusps are no longer distinguishable. The metaloph is still a wider curved loph and the protoloph a straight, shorter loph. The lophs, as in the unworn specimen, unite lingually. On the worn P"*, the dentine tracts extend well up the sides of the tooth. The dentine tract height/transverse width ratio of this tooth is 75%. Below the base of the enamel there are the beginnings of roots forming on the worn P^. The molars are hypsodont and the occlusal pattern of cusps is worn away an all specimens available, indicating that the cusps are lost at an early stage of wear On the sides of the teeth there are dentine tracts. The ratio of the height of these dentine tracts to the transverse width of the teeth is 67-89% (mean = 79%). None of the available permanent teeth show any signs of having roots at any time, regardless of the amount of wear. Discussion. — The small geomyid specimens from Pratt Quarry are clearly re- ferable to Phelosaccomys hibbardi originally known from the Clarendonian of Kansas as diagnosed by Storer (1973). The only difference in the morphology of P4 from Pratt Quarry is the presence of two anterostylids rather than three. The dentine tract on the sides of P4 are also slightly higher in the Nebraska specimens than on those from Kansas. However, these differences are quite minor and are well within any range of variation of a single species. The upper premolar of P. hibbardi has never before been identified. It is quite similar in occlusal morphology to that of P. shotwelli from the Clarendonian of California (Korth and Reynolds, 1994). Both P. hibbardi and P. shotwelli have anteroposteriorly compressed protolophs that form a straight line unlike that of older species where the protocone is more oval in shape (Korth, 1979). Phelo- saccomys hibbardi and the Pratt Quarry specimens differ from P. shotwelli in having less well-developed dentine tracts on the cheek teeth, being larger, and having a P4 that is longer than wide. The dP4 of Phelosaccomys has never been described previously. This tooth differs from those of heteromyids in having a reduced protoconid and numerous cuspules that line the buccal and anterior sides of the tooth. This type of deciduous lower premolar is more typical of geomyids (Akersten, 1973). Lindsay (1972:fig. 34b) figured the dP4 of the primitive geomyid Mojavemys. This tooth is very similar to dP4 of P. hibbardi. The similarity of the deciduous premolars of these right P4 (unworn), UNSM 101561. D, left P"* (occlusal view above, lingual view below), UNSM 101562. E, right P"^ (unworn: occlusal view above, buccal view below), UNSM 101570. Bar scale = 1 mm. 326 Annals of Carnegie Museum VOL. 67 3 - o ^ S .Q -§: S t ^•1 Z? K ■d ^ d 2 IS •$ c/. .2? >> ^ s J ^.2 1/) o 3 ^ K c/^ o ZZ D . 0^ , ON ' o ' o ^ m d- in 00 00 c4 ^ ^ ^ ON ON m in — < n (N — ' — i o NO NO O 00 O > 1 O CQ X 3.S .1 S V X Q o 3 0i)§ ° ^ ^ 2 ^ ^ „ CN d . ^ Os .5 -a os os u 2G o-fi ^ 3 X fc 2 3 -§ (U _C H -tJ •- J ^ g J Pd X « « -o M X o d 4_i ^ 3 O n c« 3 ^ m (U ^ ^ Gu II II ^ xc C<2 1^ S CN -s: a ^ Os 00 as X > <1> ^ 3 2 Z 3 ^ X „ t, (D 50 2 O X 3 -S Ph H O ^ r S ^ 2 VOL. 67 1998 Davidson and Ball — Rawlinsius papillatus, new genus, new species 375 American pattern; and Eucheila cordova Ball and Shpeley, Galerita sulcipennis species group, Eripus Dejean, Cyrtolaus Bates, and Dyschromus Chaudoir, the MesO“American Montane pattern. The details of the Meso-American Montane pattern are of no further interest in the present context. The relict status of the northern-based taxa suggests marked changes of some type that led to their isolation in the Mexican highlands, possibly drying of the intervening areas, with associated changes in vegetation. The fact that at least two different ages of dispersal are indicated suggests that either the climatic changes were cyclic in nature, or that the later-arriving N. pygmaeus had different adap- tations from the earlier arrivals, and was thus able to spread southward. Ball and Negre (1972:528) and more recent authors (e.g., Jameson 1990:412) hypothesized cyclic climatic changes within northern Mexico, resulting in conditions favorable for dispersal alternating with unfavorable conditions that resulted in more or less extensive range disruption with consequent differentiation of isolated populations that managed to survive in favorable areas or refugia. We believe, then, that the ancestral Broscina were widespread in North Amer- ica, and were possibly riparian in habitat preference. Unfavorable climatic con- ditions led to extinction of the Broscina over large parts of North America, with a single lineage persisting in southern Mexico. To the north, in western North America, one or more lineages survived and differentiated, but the descendants did not evolve the adaptations required to recover the whole of the ancestral range. Similarly, the Mexican lineage became highly specialized to live in association with fast-flowing water, and is thus unable to re-occupy the ancestral range in lowlands. Concluding Remarks The discovery of Rawlinsius papillatus was an exciting event in the develop- ment of knowledge of the Middle American carabid fauna. Additional populations of this species are not likely to be encountered casually and therefore the species may have a much wider range than known currently. Specimens probably will be found only at night, in very patchy microhabitats, mostly difficult of access. They are likely to be found only by the most energetic and persistent collectors, and then only with a good deal of luck. We hope that the discovery of this species will encourage adventurous collectors to search vigorously for further localities or additional species of Rawlinsius throughout southern Mexico, Central America, and perhaps even the northern mountains of South America. A taxon with such secretive and elusive members could turn up in any of the mountain ranges that add so much wonder and beauty to the tropics of the New World. This remarkable relictual species forces attention on the area in which it is known to live — the Sierra de Atoyac de Alvarez. This mountain range, as well as the other parts of the Sierra Madre del Sur, gives evidence of a substantial amount of precinction, especially at higher altitudes. With extensive tracts of forest still intact, this mountain range would seem to be prime ground for estab- lishment of an ecological reserve, as well as the focal point for extensive explo- ration of its fauna and flora. Undertaking such activities is a challenge for the government of Mexico, and for the Mexican biologists who are interested in biodiversity, the country’s “rich biological patrimony” (Toledo and Ordonez, 1993:775). 376 Annals of Carnegie Museum VOL. 67 Acknowledgments A preliminary draft was reviewed by B. S. Heming (Department of Biological Sciences, University of Alberta), D. A. Pollock (Carnegie Museum of Natural History), and S. Roig Jufient (Instituto Ar- gentino de Investigaciones de las Zonas Aridas, Mendoza, Argentina), all of whom made significant improvements to the manuscript. In particular, thanks to his detailed and extensive knowledge of the broscine genera. Dr. Roig Junent’s substantial advice enabled us to extend our ideas about the rela- tionships of Rawlinsius, and to correct some important errors of omission and commission. We are also grateful to editors B. C. Livezey and M. A. Schmidt for many useful comments on and improve- ments to the manuscript. We received substantial assistance with preparation of illustrations from members of our respective departments: D. Shpeley and G. B. Braybrook (Department of Biological Sciences and Department of Earth and Atmospheric Sciences, respectively. University of Alberta) did the required preparatory work and photography with the scanning electron microscope; J. S. Scott (Edmonton, Alberta) prepared the SEM plates for publication; M. Klingler (Carnegie Museum of Natural History) made the final line drawings and the plates on which they were mounted for publication; and J. E. Rawlins took and prepared the black-and-white habitus photographs and the color photographs that are reproduced as Figures 1-3, respectively, S. A. Thompson of Carnegie Museum of Natural History identified plants and sent others out for determination to B. Allen (Missouri Botanical Garden), W. Buck (New York Botanical Garden), and R, C. Moran (Institute of Biological Sciences, Aarhus University); P. D. Perkins (Museum of Comparative Zoology) identified the hydrophilid. We thank all of these individuals for their efforts on our behalf. Finally, we record a special note of thanks to John Rawlins for exercising his insatiable curiosity when he saw in the light of his headlamp the first known specimen of R. papillatus, wading in water at the edge of a small stream on a mountain road in Guerrero. Literature Cited Acorn, J. H., and G. E. Ball. 1991. The mandibles of some adult ground beetles: Structure, function, and the evolution of herbivory (Coleoptera: Carabidae). 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Das Abdomenende weibiicher, terrestrich lebender Adephaga (Coleoptera) uod seine Bedeutung fiir die Phylogeeie. Zoomorphologie, 84:113-193. Bousquet, Y., and A. Larochelle. 1993. Catalogue of the Geadephaga (Coleoptera: Trachypachidae, Rhysodidae, Carabidae, including Cicindeliei) of America north of Mexico. Memoirs of the En- tomological Society of Canada, 167:1-395. 1998 Davidson and Ball — Rawlinsws papillatus, new genus, new species 377 Brooks, D. R., and D. A. McLennan. 1991. Phylogeny, Ecology, and Behavior — A Research Pro- gram in Comparative Biology. University of Chicago Press, Chicago, Illinois. Cooper, K. W. 1990. Linear, longitudinal markings on the outer eiytral surface of beetles: Intemeurs or striae? Quaestiones Entomologicae, 26:695-699. Crowson, R. a. 1981. The Biology of the Coleoptera. Academic Press, London, United Kingdom. Curtis, J. 1839. Descriptions &c. of the insects collected by Captain P. P. King, R. N. F. R. S. & L. S. in the survey of the Straits of Magellan. Transactions of the Linnean Society of London, 18: 181-205. Dejean, P. F. M. A. 1828. Species general des Coleopteres, de la collection de M. le Comte Dejean. Tome troisieme. Mequignon-Marvis, Paris, France. Deuve, T. 1993. L’ abdomen et les genitalia des femelles de Coleopteres Adephaga. Memoires du Museum National d’Histoire Naturelle, Tome 155, Zoologie. Paris, France. Emden, F. I. van. 1942. A key to the genera of larval Carabidae. Transactions of the Royal Ento- mological Society of London, 92:1-99. Erwin, T. L. 1974. Studies on the subtribe Tachyina (Coleoptera: Carabidae: Bembidiini), Part II: A revision of the New World-Australian genus Pericompsus LeConte. Smithsonian Contributions to Zoology, 162:1-96. . 1985. The taxon pulse. Pp. 437-493, in Taxonomy, Phylogeny and Zoogeography of Beetles and Ants: A Volume Dedicated to the Memory of Philip Jackson Darlington, Jr. (1904-1983) (G. E. Ball, ed.). Dr. W. Junk, Publishers, Dordrecht, The Netherlands. Eschscholtz, j. F. 1830. Nova genera coleopterorum faunae Europaeae. Bulletin de la Societe Im- periale des Naturalistes de Moscou, 2:63-66. Goulet, H. 1983. The genera of Holarctic Elaphrini and species of Elaphrus Fabricius (Coleoptera: Carabidae): Classification, phylogeny, and zoogeography. Quaestiones Entomologicae, 19: 219-482. Guerin-Meneville, M. F. E. 1838. Insectes du voyage de la Favorite. Magasin du Zoologie, 8: 225-238. Halffter, G. 1987. Biogeography of the montane entomofauna of Mexico and Central America. Annual Review of Entomology, 32:95-114. Hennig, W. 1966. Phylogenetic Systematics. University of Illinois Press, Urbana, Illinois. Horn, G. H. 1881. On the genera of Carabidae with special reference to the fauna of boreal America. Transactions of the American Entomological Society, 9:91-196. Jameson, M. L. 1990. Revision, phylogeny and biogeography of the genera Parabyrsopolis Ohaus and Viridimicus, new genus (Coleoptera: Scarabaeidae: Rutelinae). Coleopterists Bulletin, 44: 377^22. Jeannel, R. 1928. Monographic des Trechinae: Morphologic comparee et distribution geographique d’un groupe de Coleopteres. (Troisieme livraison). Les Trechini cavemicoles. UAbeille, Journal d’Entomologie, 35:1-808. — . 1941. Coleopteres carabiques. Premiere partie. Faune de France 39. Librairie de la Faculte des Sciences, Paris, France. Kavanaugh, D. H. 1978. The Nearctic species of Nebria Latreille (Coleoptera; Carabidae: Nebriini): Classification, phylogeny, zoogeography, and natural history. Unpublished Ph.D. Thesis, Univer- sity of Alberta, Edmonton, Alberta, Canada. Kohlmann, B., and G. Halffter. 1990. Reconstruction of a specific example of insect invasion waves: The cladistic analysis of Canthon (Coleoptera: Scarabaeidae) and related genera in North America. Quaestiones Entomologicae, 26:1-20. LeConte, J. L. 1869. List of Coleoptera collected in Vancouver’s Island by Henry and Joseph Mat- thews, with descriptions of some new species. Annals and Magazine of Natural History (Series 4), 4:369-385. Liebherr, j. K. 1986. Barylaus, new genus (Coleoptera: Carabidae) endemic to the West Indies with Old World affinities. Journal of the New York Entomological Society, 94:83-97. . 1994. Biogeographic patterns of montane Mexican and Central American Carabidae (Cole- optera). Canadian Entomologist, 126:841-860. Liebherr, J. K., and K. W. Will. In press. Inferring phylogenetic relationships within the Carabidae (Insecta: Coleoptera) from characters of the female reproductive tract. Bollettino del Museo Re- gionale di Scienze Natural! (Torino). Lindroth, C. H. 1961a. On Miscodera Esch., and related genera (Col. Carabidae). Opuscula Ento- mologica, 26:145-152. — I96lb. The ground-beetles (Carabidae, excL Cicindelinae) of Canada and Alaska. Part 2. Opuscula Entomologica Supplementum, 20:1-200. Llorente-Bousquets, j., and a. Luis-Martinez. 1993. Conservation-oriented analysis of Mexican 378 Annals of Carnegie Museum VOL. 67 butterflies: Papilionidae (Lepidoptera, Papilionoidea). Pp. 147-178, in Biological Diversity of Mexico: Origins and Distribution (T. P. Ramamoorthy, R. Bye, A. Lat, and J. Fa, eds.). Oxford University Press, New York, New York. Moore, B. P. 1964. Australian larval Carabidae of the subfamilies Broscinae, Psydrinae and Pseu- domorphinae. Pacific Insects, 6:242-246. Noonan, G. R. 1973. The anisodactylines (Insecta: Coleoptera: Carabidae: Harpalini). Quaestiones Entomologicae, 9:266^80. Panzer, G. W. F. 1813. Index entomologicus, sistens omnes insectoram species in G. W. F. Panzeri Fauna Insectorum Germaeica descriptas atque delineatas secundum methodum Fabricianam: Ad- jectis emendationibus, observationibus. Pars 1. Eleutherata. Felssecker, Norimbergae, Norway. Roig Junent, S. 1995. Revision sistematica de los Creobina de America del Sur (Coleoptera: Carab- idae: Brosciei). Acta Entomologica Chilena, 19:51-74. Roig Junent, S., and G. E. Ball. 1995. Nothobroscus chilensis, new genus and new species from southern South America (Coleoptera: Carabidae; Broscini). Coleopterists Bulletin, 49:301-312. Shpeley, D., and G. E. Ball. 1993. Classification, reconstructed phylogeny and geographical history of the New World species of Coptodera Dejean. Proceedings of the Entomological Society of Ontario, 124:3-182, Simpson, G. G. 1944. Tempo and Mode in Evolution. Columbia University Press, New York, New York. Straneo, S. L., and G. E. Ball. 1989. Synopsis of the genera and subgenera of the tribe Peleciini, and revision of the Neotropical and Oriental species. Insecta Mundi, 3:73-179. Tanner, V. M. 1927. A preliminary study of the genitalia of female Coleoptera. Transactions of the American Entomological Society, 53:5-50. Thiele, H.-U. 1977. Carabid beetles in their environments: A study on habitat selection by adaptations in physiology and behaviour. Springer- Verlag, Berlin, Germany. Thompson, R. G. 1979. Larvae of North American Carabidae with a key to tribes. Pp. 209-291, in Carabid Beetles: Their Evolution, Natural History, and Classification (T. L. Erwin, G. E. Ball, D. R. Whitehead, and A. L. Halpem, eds,). Dr. W. Junk bv, Publishers, The Hague, The Nether- lands. Toledo, V. M., and M. de Jesus Ordonez. 1993. The biodiversity scenario of Mexico: A review of terrestrial habitats. Pp. 757-777, in Biological Diversity of Mexico: Origins and Distribution (T. P. Ramamoorthy, R. Bye, A. Lat, and J. Fa, eds.). Oxford University Press, New York, New York. Vargas-Fernandez, L, J. Llorente-Bousquets, and a. Luis-Martinez. 1992. Listado Lepidopter- ofaunistico de la Sierra de Atoyac de Alvarez en el estado de Guerrero: Notas acerca de su distribucion local y nacional (Rhopalocera: Papilionoidea). Folia Entomologica Mexicana, 86: 41-178. INDEX TO VOLUME 67 CONTENTS Significance of nearshore trace-fossil assemblages of the Cambro-Ordovician Deadwood For- mation and Aladdin Sandstone, South Dakota . Thomas M. Stanley and Rodney M. Feldmann Diadectes (Diadectomorpha: Diadectidae) from the early Permian of central Germany, with description of a new species . . . David S Berman, Stuart S. Sumida, and Thomas Martens New species of Cernotina Ross (Insecta: Trichoptera: Polycentropodidae) from the Amazon Basin in northeastern Peru and northern Brazil Jan L. Sykora Atokan (Late Bashkirian or Early Moscovian) brachiopods from the Hare Fiord Formation of Ellesmere Island, Canadian Arctic Archipelago . . John L. Carter and Vladislav 1. Poletaev Taxonomic status of the Early Permian Helodectes paridens Cope (Diadectidae) with discussion of occlusion of diadectid marginal dentitions David S Berman, Amy C. Henrici, and Stuart S. Sumida Archaeology of Trants, Montserrat. Part 4. Flaked stone and stone bead industries ....... . John G. Crock and Robert N. Baitone On the generic status of Palaeophichthys parvulus Eastman 1908 and Monongahela stenodonta Lund 1970 (Osteichthyes: Dipnoi) Anne Kemp Revision of the Neotropical genus Ischyomius with a discussion on its systematic position (Insecta: Coleoptera: Tenebrionoidea: Pythidae) Darren A. Pollock [Review of] Maples of the World (D. M. van Geldersen et al.) Frederick H. Utech Contribution to the knowledge of the Brentidae (Insecta: Coleoptera) from Tropical Africa, with redescription of Plesiobolbus sagax Kolbe Alessandra Sforzi The butterflies of the Isle of Pines, Cuba: Eighty years on David Spencer Smith, Luis Roberto Hernandez, and Neil Davies Rodents and lagomorphs (Mammalia) from the Late Clarendonian (Miocene) Ash Hollow For- mation, Brown County, Nebraska William W. Korth The tribe Broscini in Mexico: Rawlinsius papillatus, new genus and new species (Insecta: Coleoptera: Carabidae), with notes on natural history and evolution Robert L. Davidson and George E. Ball 1 53 95 105 181 197 225 245 265 267 281 299 349 379 380 Annals of Carnegie Museum VOL. 67 new taxa new genera and species ■\Antecalomys, new genus 332 ■\Antecalomys phthanus, new species 333 "fAntronaria annosa, new species 146 '\Camarium nuperum, new species 149 ■fCareoseptum, new genus 139 '\Careoseptum septentrionalis, new species 139 Cernotina aestheticella, new species 99 Cernotina ecotura, new species 99 Cernotina harrisi, new species 96 Cernotina nigridentata, new species 96 "fCranaena nassichuki, new species 174 ’tCupidinimus prattensis, new species 321 tDiadectes absitus, new species 56 fDipoides tanneri, new species 308 'fElassonia sverdrupensis, new species 144 'tElinoria ellesmerensis, new species 162 "fExlaminella, new genus 142 Exlaminella insolita, new species 142 fFimbrinia borealis, new species 123 '\Heteraria canadiensis, new species 158 Ischyomius championi, new species 260 Ischyomius nevermanni, new species 260 ^Lazarevia, new genus 125 fLazarevia stepanowensis, new species 126 ^Liraria paucispina, new species 133 ^Maemia gelida, new species 127 ■\Meristorygma, new species A 171 tNucleospira aquilonaris, new species 151 '\Orthotichia dorsistrigis, new species 116 f Parachoristites tellevakensis, new species 163 '\Plicatiferina kalashnikovi, new species 117 fPronotolagus whitei, new species 341 RawUnsius, new genus 361 Rawlinsius papillatus, new species 361 fRugivestis pristina, new species 122 '\Tiramnia grunti, new species 155 ^Tiramnia walteri, new species 154 Fossil taxa 1998 Index to Volume 67 381 AUTHOR INDEX Ball, George E. ...................................................... 349 Bartone, Robert N. .................................................... 197 Berman, David S ................................................... . 53, 181 Carter, John L. ...................................................... . 105 Crock, John G. ...................................................... . 197 Davidson, Robert L. ................................................... 349 Davies, Neil ......................................................... 281 Feldmann, Rodney M. ................................................. 1 Henrici, Amy C. ..................................................... . 181 Hernandez, Luis Roberto ................................................ 281 Kemp, Anne ........................................................ 225 Korth, William W. .................................................... 299 Martens, Thomas ..................................................... 53 Poletaev, Vladislav I. .................................................. 105 Pollock, Darren A. .................................................... 245 Sforzi, Allesandra ..................................................... 267 Smith, David Spencer .................................................. 281 Stanley, Thomas M. ................................................... 1 Sumida, Stuart S. .................................................. . 53, 181 Sykora, Jan L. ....................................................... 95 Utech, Frederick H. ................................................... 265 . /i'.'Oi’ It ‘:-^ '■ U ^ ' '\i ^ * ■■ ■' V- ^ ' ^,' •■'; **> J*, '>»>.!■>;> ^\i . f«y^ X :’* . . ' * 'iSf . J'-.' ■4 IT’S-"* ■}:.C'.^^» ,Vi ' * ' ■ '■‘’'* ' "■ **' * "V ■.'*!> • *■ ■•• » *S; • *r I ^ ) *, •# 1 ‘*Jai 'f iVicf? ;■' * .irjj'liiA^f .;^.v ,|^ ''%■* « ■ '^«>, * ^ - ^ /:V3>. :; ' '" ■'’if-'t/S'l .•*■'«» V-'W» -(.^i^apft, • .."!«. ■ - . '■ . ' ' ' »-* -f.S-..-|y^]^ ; : ■ t *■ * # r rjy '4 * » « , 'I iX ‘tL-^A*. ,tt* '*.. (-v’ «■ •,' ■ .'■' -'4-4if ,. » . -'KMdP,:. 'X Xi 5^.*- ■ "A ■ • ’ « > '»- < ft ^ m i~ , v; >4r*.w^ ;V-) • ' « i 1 ' . ■: ■«(. 4^“’ •?E i ^ ^ J' Miai %;4«4i 'I 4h ■ ' r •■ . _r./f ■-’'> . ■ ■¥>r-0--t>,. r INSTRUCTIONS FOR AUTHORS ANNALS OF CARNEGIE MUSEUM consist of contributions to the earth sciences (including paleontol- ogy), life sciences, and anthropology, in 30 by 46 picas format (127 by 195 mm or 5 by 7% inches). Submit all manuscripts to the Office of Scientific Publications. Au- thors should give particular attention to scientific con- tent, format, and general style for the ANNALS. 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