Cretalamna Glikman, 1958

Siversson, Mikael, Lindgren, Johan, Newbrey, Michael G., Cederström, Peter & Cook, Todd D., 2015, Cenomanian-Campanian (Late Cretaceous) mid-palaeolatitude sharks of Cretalamna appendiculata type, Acta Palaeontologica Polonica 60 (2), pp. 339-384 : 342-346

Genus Cretalamna Glikman, 1958

Type species: Otodus appendiculatus Agassiz, 1843 , by original designation; early Cenomanian to early Coniacian interval, Lewes, England.

Emended diagnosis.—Quadrate process more than twice as high as dorsoventrally narrowest part of palatoquadrate, anterior to quadrate process but posterior to section harbouring anterior tooth files. Most distal upper anterior tooth and most mesial upper lateroposterior tooth not markedly reduced in size relative to adjacent teeth. Intermediate teeth absent. Root typically without median groove. Median indentation of basal edge of root shallow in labial/lingual views in lateroposterior teeth. Lobes of root more-or-less quadrangular in lateroposterior tooth files. Radial lamellae of vertebral centra moderately robust with spacing equivalent to thickness. Neural arch foramina subovate, extending full length of centrum between corpus calcarea.

Remarks.—There is no visible evidence of an orbital process sensu Compagno 2001) on the palatoquadrates of LACM 128126 ( Fig. 13A, B ). However, the dorsal edge of the more complete, right palatoquadrate is not well enough preserved to reveal if a cartilaginous orbital process was originally present.

Agassiz’s former syntypes of Otodus appendiculatus Agassiz 1843 : pl. 32: 1–25) are a heterogeneous mix, comprising several genera ( Cretodus ? [Sokolov, 1965] [Agassiz 1843: fig. 9], Cretalamna [Agassiz 1843: e.g., fig. 10], Dwardius Siverson, 1999 [Agassiz 1843: fig. 15], and Cretoxyrhina Glikman, 1958 [Agassiz 1843: fig. 25]) and probably about half a dozen species if not more. Siverson (1999) designated the upper? lateroposterior tooth in Agassiz’s (1843: fig. 10) as lectotype for C. appendiculata ( Fig. 1 ). This species concept conformed to that of most current authors of that time (e.g., Müller and Diedrich 1991; Manning and Dockery 1992; Welton and Farish 1993; Williamson et al. 1993). However, the evolution of C. appendiculata sensu lato turns out to be more complex than perhaps anyone could have imagined. The lectotype of C. appendiculata was collected from “White Chalk” near Lewes, East Sussex, southern England (Agassiz 1843: 270). About a dozen more-or-less overgrown quarries are spread around Lewes and expose or exposed strata of early Cenomanian to mid-Santonian age ( Dineley and Metcalf 1999). The Turonian stage is particularly well represented with current or former exposures in a majority of the local quarries ( Dineley and Metcalf 1999: fig. 13.18). Mining activity in the mid 1800’s exposed strata of Cenomanian to early Coniacian age (Andrew Gale, personal communication 2010), effectively bracketing the possible age range of the lectotype.

The lectotype of C. appendiculata is from the Gideon A. Mantell collection but may not have been transferred to the Natural History Museum in London as it was not listed as a British Museum ( NHMUK) specimen by Woodward (1889). Several of the paralectotypes of “ Otodus ” appendiculatus , derived from Mantell’s collection, did however end up in the NHMUK collections (Agassiz 1843: pl. 32: 1, 5, 6, 11, and 14). One could argue that C. appendiculata is a nomen dubium because the whereabouts of the lectotype are currently unknown, the type stratum is likewise unknown and possible source rocks around Lewes exposed in the mid 1800s span approximately 10 Ma (early Cenomanian to early Coniacian). However, as C. appendiculata is the type species of a nominal genus that has become well known to many palaeontologists outside the arena of fossil shark studies, there is arguably a strong case for preserving taxonomic stability. Doing so would also be in the spirit of the latest edition of the International Code of Zoological Nomenclature ( Ride et al. 2000).Complete teeth of Cretalamna with an uncorroded root are seemingly extremely rare in the Cenomanian–early Coniacian of the Lewes area, as we are only aware of one well preserved specimen (although the tip of the mesial lobe is missing), a lower lateroposterior of C. aff. C. gertericorum sp. nov. from the middle Cenomanian Acanthoceras rhotomagense Zone, Southerham Grey Pit (David Ward collection). Lateroposterior teeth in species of the C. borealis group (like C. gertericorum sp. nov.) are readily separable from the lectotype of C. appendiculata by their strongly distally curved cusp in teeth with a comparable profile view.

The thin, conglomeratic marl of early Turonian age, constituting the Tourtia de Bettrechies in the SECAB quarry, Bettrechies, northern France, is right in the middle of the possible age range of the lectotype (Cenomanian–early Coniacian) and produces abundant teeth of Cretalamna . Examination of several collections of Cretalamna teeth from the Tourtia de Bettrechies in the SECAB quarry, derived from bulk sampling and surface in situ collecting, resulted in the recognition of three species of Cretalamna , one of which includes teeth with a morphology very close to that of the lectotype of C. appendiculata (straight thick cusp, rounded root-lobes and a profile view indicating an upper jaw position; Fig. 2E ). The latter was illustrated by Agassiz (1843: pl. 32: 10) in profile and labial views. The profile view ( Fig. 1B ) is, however, of somewhat limited use in this case (other than for indicating an upper jaw position) as the tooth does not appear to have been illustrated at right angles to the vertical height axis, hence the position of the apex of the cusplet in the same horizontal plane as the summit of the lingual protuberance of the root. A good match was achieved by tilting the cusp of the tooth in Fig. 2E 2 about 45° towards the viewer. The relatively low protuberance of the lectotype is probably a result of abrasion. Several of the Cretalamna teeth from Bettrechies are very well preserved except for a “polished-appearing” lingual protuberance (see high magnification view of e.g., Fig. 2B 4 , the area between the main lingual foramen and the root/neck boundary). The ten teeth from the Tourtia de Bettrechies in the SECAB quarry, assigned to C. appendiculata ( Figs. 2 , 3 ), enabled the formulation of a meaningful diagnosis for the species and a much improved understanding of its relationship to other Cretaceous Cretalamna .

The spelling of the name of the genus is controversial. While strictly adhering to the Code of Zoological Nomenclature in force at the time ( Ride et al. 1985), Siverson (1999) reinstated the original spelling Cretalamna , replacing the unjustified but universally adopted subsequent spelling Cretolamna . Within a few years, usage of the original spelling was embraced by a growing number of workers (e.g., Cook et al. 2008; Everhart 2005; Kriwet and Benton 2004; Kriwet et al. 2008; Shimada 2007; Shimada et al. 2006). Over the last few years a majority of researchers discussing or mentioning the genus in publications have used the original spelling Cretalamna (e.g., Adolfssen and Ward 2015; Andreev and Motchurova-Dekova 2010; Becker et al. 2010; Bogan and Gallina 2011; Bourdon et al. 2011; Bourdon and Everhart 2011; Cook et al. 2011, 2013; Cumbaa et al. 2010; Cuny et al. 2012; Einarsson et al. 2010; Eriksson et al. 2011; Hamm and Cicimurri 2011; Retzler et al. 2013; Sato et al. 2012; Shimada et al. 2010; Siverson et al. 2013; Underwood and Cumbaa 2010; Underwood et al. 2011; Ward 2009). The replacement of Cretolamna with Cretalamna was strongly opposed by Cappetta (2000, 2012). He presented his case to a representative for the International Commission for Zoological Nomenclature ( Cappetta 2000) who (in litt.), while citing Article 33.3.1 of the 2000 Edition of the Code, support- ed the retention of the spelling Cretolamna . However, it is our view that the 2000 edition cannot be used retrospectively, without a formal ruling by the Commission, to alter justified nomenclatural acts carried out before the 2000 edition came into effect. Moreover, article 33.3.1 states that “when an unjustified emendation is in prevailing usage and is attributed to the original author and date it is deemed to be a justified emendation”. Given that a majority of current students have adopted the original spelling Cretalamna , Article 33.3.1 can no longer be invoked in support of the spelling Cretolamna . Cappetta (2012) indicated that the spelling Cretolamna is used by an overwhelming number of workers and cit- ed a number of publications, all of which were published before Siverson’s (1999) work. Cappetta would have had a stronger case if the original spelling had been reinstated in 2000 or later as this would have represented a violation of Article 33.3.1. As it stands, however, the reinstatement of the original spelling strictly adhered to the Code in force at the time and Cretalamna is in prevailing usage and, if anything, is supported rather than threatened by Article 33.3.1 of the 2000 edition.

Geographic and stratigraphic range.— Cretalamna sensu stricto ranges from the Late Albian (Early Cretaceous) to the Ypresian or possibly Lutetian (Eocene). It has been reported from all continents except Antarctica.

Cretalamna appendiculata (Agassiz, 1843)

Figs. 2 , 3 .

1843 Otodus appendiculatus [partim]; Agassiz 1843: 270, pl. 32: 10. Lectotype: lateroposterior tooth, presumably from the upper jaw (repository currently unknown).

Type horizon: Cenomanian–early Coniacian, Late Cretaceous.

Type localit y: Lewes area, East Sussex, southern England.

Material.—Ten teeth ( WAM 13.5.1–7, UMT-BET 1–3) from the Tourtia de Bettrechies (early Turonian, Late Cretaceous); SECAB quarry, Bettrechies, France.

Emended diagnosis.—Cusplets relatively small in upper lateroposterior teeth and moderately divergent. Cusp labially curved in profile view in upper lateroposterior teeth. Lower anterior teeth with high root but moderately tall, triangular cusp in lingual view. Root massive in lower anteriors of large individuals, displaying protruding lingual protuberance in profile view and triangular outline in basal view. Upper anterior and anteriorly situated upper lateroposterior teeth with deeply U-shaped median indentation of basal edge of root and round- ed lobes. Distal lobe of root labiolingually compressed relative to mesial lobe in anteriorly situated upper lateroposterior teeth. Lingual protuberance of root high but apico-basally narrow in profile view in upper and lower lateroposterior teeth.

Description.— First upper anterior tooth: A 15.5 mm high tooth is referred to the first upper right file ( Fig. 2A ). One cusplet is missing, presumably the distal one. The cusp is straight in labial/lingual views and lingually curved in profile view. The preserved cusplet is symmetrical and set at a 45° angle to the cusp. The lingual protuberance of the root is well demarcated in basal view. The root is almost symmetrical in labial/lingual views although the mesial? lobe is somewhat wider and labiolingually less compressed.

Second upper anterior tooth file:A well-preserved, 17 mm high tooth is assigned to the right A2 position ( Fig. 2B ). The robust cusp is moderately tall and very slightly curved in labial/lingual views, presumably towards the commissure. In profile view, the cusp is slightly labially curved. The cusplets are sub-triangular and moderately divergent. The root is relatively gracile and slightly asymmetrical in labial view, with the mesial lobe somewhat shorter and broader than is the distal lobe. In basal view, the root is asymmetrical with the distal lobe more compressed than the mesial lobe.

Upper lateroposterior tooth files: Of the four well-preserved upper lateroposterior teeth at hand range, three of them Fig. 2C–E ), 14–15 mm high, are from anteriorly situated tooth files (probably within the LP 1– LP 4 range), whereas one Fig. 2F ), 9 mm high, is a posteriorly situated tooth (estimated position LP 7 or LP 8). All teeth have cutting edges somewhat labially curved in profile view. The cusp is slightly- to moderately distally inclined in the anteriorly situated teeth but strongly distally curved in the posteriorly situated tooth. The root is markedly asymmetrical in the two most anteriorly situated teeth ( Fig. 2C, D ) which both also have a deep, U-shaped median indentation of the basal edge of the root. The lateral cusplets are sub-triangular, moderately large and relatively upright on all four teeth. In basal view, the root is symmetrical in the posteriorly situated tooth ( Fig. 2F 3 ) but asymmetrical in the anteriorly situated teeth with the distal lobe labiolingually compressed relative to the mesial lobe ( Fig. 2C 1 , D 2 , E 3 ).

Second lower anterior tooth file: A complete, 21.5 mm high tooth from a large individual is referred to the second lower anterior file ( Fig. 3A ). The triangular cusp is very slightly distally inclined in labial/lingual views and moderately thick. The cutting edges of the cusp are straight in profile view. The root is massive and asymmetrical in labial view, as the mesial lobe is longer and narrower than the distal lobe. A triangular outline characterises the basal view. In profile view, the neck and lingual protuberance rise sharply from the lingual demarcation of the cusp.

Lower lateroposterior tooth file: Three teeth are referred to the lower lateroposterior files. The largest of the three (probably an lp1), measuring 18 mm in height, is well preserved with the exception of a small missing portion of the mesial lobe of the root ( Fig. 3B ). The cusp is slightly distally inclined in labial/lingual views and very slightly lingually curved in profile view. The root is markedly asymmetrical with its angular and broad distal lobe and more acute and narrower mesial lobe. The lingual protuberance is relatively narrow and well demarcated in basal view ( Fig. 3B 4 ). The median indentation of the basal edge of the root is deeply U-shaped. The medium-sized tooth is missing the tip of the cusp ( Fig. 3C 3 ). As preserved it measures 10 mm in height and would have been located in the middle part of the inferred lower lateroposterior hollow. The cusp is moderately distally curved in labial/lingual views and slightly lingually curved in profile view. The mesial cusplet is noticeably larger than the distal cusplet. The median indentation of the basal edge of the root forms a wide-angled V. The smallest of the well-preserved lower lateroposterior teeth is 6.5 mm high ( Fig. 3D ). The apex of the distally inclined cusp is missing. The rounded median indentation of the basal edge of the root is both wide and deep. Both lobes of the root are sub-rectangular in basal view. A narrow median groove divides the lingual protuberance of the root.

Remarks.—Teeth of C. appendiculata most closely resemble the teeth of the early Campanian C. sarcoportheta sp. nov., and both species are included in the C. appendiculata group. Presumed derived characters uniting them include a marked asymmetry of the root in basal view in anteriorly (but noticeably not in posteriorly) situated upper lateroposterior teeth, with the distal lobe being labiolingually compressed relative to the mesial lobe ( Figs. 2C 1 , D 2 , E 3 , 5 B 2 , E 1, F 1 ), a strongly compressed distal lobe in presumed second upper anteri- or teeth ( Figs. 2B 2 , 4D 1 , E 3, F 3 ; note that the cusp appears to be somewhat distally curved in C. appendiculata but very slightly recurved in C. sarcoportheta sp. nov.) and a very prominent lingual protuberance of the root in lower anterior teeth ( Figs. 3A 2 , 7A 2, B 2, C 2 ). Apart from the morphology of the putative A2, comparable teeth of the Campanian species differ from those of C. appendiculata by their more protruding lingual protuberance of the root (see e.g., Figs. 2C 4 , 5F 3 ), less concave labial face in profile view in upper lateroposterior teeth from comparable files (compare Figs. 2D 4 and E 2 with 5F 3 and H 2), somewhat broader cusp (on average), more divergent cusplets, and less tightly curved median indentation of the root in anteriorly situated lateroposterior teeth.

Anterior teeth of the co-existing C. gertericorum sp. nov. are much taller relative to their width than are those of C. appendiculata . The cusp is straight in profile view in the lower anterior tooth at hand of C. appendiculata ( Fig. 3A 2 ), but lingually curved in the probable a1 of C. gertericorum sp. nov., as it is in other species of the C. borealis group ( Figs. 10A 4 , 11F 2 ). Upper lateroposterior teeth of C. gertericorum sp. nov. have generally a more strongly distally curved cusp than have teeth of C. appendiculata . The median indentation of the basal edge of the root is smaller and more tightly curved in lateroposterior teeth of C. gertericorum sp. nov. than it is in comparable teeth of C. appendiculata (compare Figs. 2F 1 , 3D 2 with Fig. 11E 1, G 1 ). The lingual protuberance of the root is very small but well demarcated in basal view of most lateroposterior teeth of C. gertericorum sp. nov. ( Fig. 11E 3, G 3 ) but much wider and less well demarcated in C. appendiculata ( Figs. 2F 3, 3D 4 ).

Lower anterior teeth of C. deschutteri sp. nov. have a lingually curved cusp in profile view (straight in the probable a2

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of C. appendiculata ), a lingual protuberance of the root well defined in basal view (poorly defined in C. appendiculata ) but poorly demarcated from the main body of the root in profile view (protruding in C. appendiculata ). Upper lateroposterior teeth of C. deschutteri sp. nov. have a symmetrical root in basal view (as in C. hattini sp. nov., whereas it is strongly asymmet-

4

1

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rical in at least more anteriorly situated teeth of C. appendiculata ) and a labiolingually compressed cusp. In contrast, the cusp is thick and robust in comparable upper lateroposteriors of C. appendiculata (compare Fig. 2F 2 with Fig. 18E 2 ).

Herman (1977: pl. 9: 2) illustrated 12 teeth (five from the late Cenomanian Praeactinocamax plenus Zone [uppermost Tourtia de Mons”] and seven from the middle Turonian Terebratulina rigida Zone [= T. lata Zone , see Appendix 2]) from the SECAB quarry, Bettrechies, as Cretolamna appendiculata var. appendiculata . One of the teeth from the T. lata Zone ( Herman 1977: pl. 9: 2c) is an upper anterior (probably A2) of a C. borealis -group species (possibly C. gertericorum

sp. nov.). We are uncertain about the identity of the other 11 teeth as we have not had the opportunity to examine them first hand. Even moderate abrasion of the root may render a Cretalamna tooth indeterminable at species level.

Geographic and stratigraphic range.— At this stage the species can only be positively identified from the type area in England and the early Turonian of northern France.

Figures

Fig. 13. Otodontid shark Cretalamna hattini sp. nov., holotype, LACM 128126, latest Santonian or earliest Campanian (Late Cretaceous); upper Smoky Hill Chalk, Niobrara Formation, close to MU 16, Logan County, Kansas, USA. A. Right palatoquadrate in lingual view. B. Anterior section of right palatoquadrate showing embedded replacement teeth, re-interpreted as (from left to right) A2, LP1 and LP2 (two cusps) C. Disarticulated tooth assigned to the upper “intermediate” file by Shimada (2007) and to the LP5 file herein. D. In situ tooth assigned to the the upper “intermediate” file by Shimada (2007) and to the LP1 file in this work. E. Disarticulated tooth assigned to the LP1 file in this work.

Fig. 1. Otodontid shark Cretalamna appendiculata (Agassiz, 1843), Lewes area, East Sussex, England, Cenomanian–early Coniacian interval, lectotype in labial (A) and tilted profile (B) views (reproduced from Agassiz 1843: pl. 32, fig. 10).

Fig. 2. Otodontid shark Cretalamna appendiculata (Agassiz, 1843), upper jaw teeth, Tourtia de Bettrechies, early Turonian (Late Cretaceous); SECAB quarry, Bettrechies, northern France. A. WAM 13.5.2, right A1?; labial (A), basal (A), profile (A), and lingual (A) views. B. UM-BET 1, right A2?; 1 2 3 4 labial (B 1), basal (B 2), profile (B 3), and lingual (B 4) views. C. UM-BET 2, right LP1?; basal (C 1), lingual (C 2), labial (C 3), and profile (C 4) views. D. UM- BET 3, left LP2?; labial (D 1), basal (D 2), lingual (D 3), and profile (D 4) views. E. WAM 13.5.7, left LP3?; labial (E 1), profile (E 2), basal (E 3), and lingual (E ) views. F. WAM 13.5.3, right LP8?; labial (F ), profile (F ), basal (F ), and lingual (F ) views.

Fig. 4. Otodontid shark Cretalamna sarcoportheta sp. nov., upper anterior teeth, informal Belemnellocamax mammillatus zone, latest early Campanian (Late Cretaceous); Åsen locality, Kristianstad Basin, southern Sweden. A. Paratype, LO 10889 t, left A1?; labial (A1), profile (A2), basal (A3), and lingual (A4) views. B. Paratype, LO 10890 t, right A1?; labial (B1), basal (B2), profile (B3), and lingual (B4) views. C. Paratype, LO 10891 t, right A1?; basal (C1), labial (C2, stereopair), profile (C3), and lingual (C4) views. D. Holotype, LO 10888T, left A2?; basal (D1), lingual (D2), labial (D3, stereopair), and profile (D4) views. E. Paratype, LO 10892 t, left A2?; labial (E1), profile (E2), basal (E3), and lingual (E4) views. F. Paratype, LO 10893 t, right A2?; profile (F1), labial (F2), basal (F3), and lingual (F4), views.

Fig. 3. Otodontid shark Cretalamna appendiculata (Agassiz, 1843), lower jaw teeth, Tourtia de Bettrechies, early Turonian (Late Cretaceous); SECAB quarry, Bettrechies, northern France. A. WAM 13.5.6, right a2? from a large individual; labial (A), profile (A), lingual (A), and basal (A) views. 1 2 3 4 B. WAM 13.5.5, left lp1?; labial (B 1), profile (B 2), lingual (B 3), and basal (B 4) views. C. WAM 13.5.4, right lp6?; profile (C 1), labial (C 2), lingual (C 3), and basal (C ) views. D. WAM 13.5.1, right lp8?; profile (D ), labial (D ), lingual (D ), and basal (D ) views.

Fig. 5. Otodontid shark Cretalamna sarcoportheta sp. nov., anteriorly situated upper lateroposterior teeth, informal Belemnellocamax mammillatus zone, latest early Campanian (Late Cretaceous); Åsen locality, Kristianstad Basin, southern Sweden. A. Paratype, LO 10894t, left LP1?; labial (A), basal (A), 1 2 lingual (A), and profile (A) views. B. Paratype, LO 10895t, left LP2?; lingual (B), basal (B), labial (B), and profile (B) views. C. Paratype, LO 10896t, 3 4 1 2 3 4 right LP2?; lingual (C ), profile (C ), and labial (C ) views. D. Paratype, LO 10897t, right LP3?; labial (D ), lingual (D ), and profile (D ) views. E. Paratype,

Fig. 7. Otodontid shark Cretalamna sarcoportheta sp. nov., lower anterior teeth, informal Belemnellocamax mammillatus zone, latest early Campanian Late Cretaceous); Åsen locality, Kristianstad Basin, southern Sweden. A. Paratype, LO 10907t, left a1?; labial (A 1, stereopair), profile (A 2), basal (A 3), and lingual (A 4) views. B. Paratype, LO 10908t, right a1?; lingual (B 1), profile (B 2), and labial (B 3) views. C. Paratype, LO 10909t, right a2?; lingual (C 1), profile (C ), and labial (C ) views.

Fig. 10. Otodontid shark Cretalamna borealis (Priem, 1897), lower jaw teeth, informal Belemnellocamax mammillatus zone, latest early Campanian (Late Cretaceous); Ignaberga (A) and Åsen (B) quarries, Kristianstad Basin, southern Sweden. A. LO 6350t, right a1?; lingual (A 1), basal (A 2), labial (A 3), and profile (A 4) views. B. LO 11137t, left lateroposterior tooth; labial (B ), lingual (B ), and profile (B ) views.

Fig. 11. Otodontid shark Cretalamna gertericorum sp. nov., Tourtia de Bettrechies, early Turonian (Late Cretaceous); SECAB quarry, Bettrechies, northern France. A. Paratype, WAM 13.5.12, left A2?; labial (A 1), profile (A 2), basal (A 3), and lingual (A 4) views. B. Paratype, UM-BET 4, right upper lateroposterior tooth; labial (B 1), profile (B 2), basal (B 3), and lingual (B 4) views. C. Paratype, WAM 13.5.9, right upper lateroposterior tooth, distal half of distal lobe of root broken off; basal (C 1), labial (C 2), and lingual (C 3) views. D. Paratype, WAM 13.5.8, left upper lateroposterior tooth; labial (D 1), basal (D 2), lingual (D 3), and profile (D 4) views. E. Paratype, WAM 13.5.11, left upper lateroposterior tooth; labial (E 1), profile (E 2), basal (E 3), and lingual (E 4) views. F. Paratype, WAM 13.5.10, a1?; labial (F 1), profile (F 2), lingual (F 3), and basal (F 4) views. G. Holotype, UM-BET 5, right lower lateroposterior tooth; labial (G ), lingual (G ), and basal (G ) views.

Fig. 18. Otodontid shark Cretalamna deschutteri sp. nov., Tourtia de Bettrechies, early Turonian (Late Cretaceous); SECAB quarry, Bettrechies, northern France. A. Paratype, WAM 13.5.18, left A1?; labial (A 1), basal (A 2), profile (A 3), and lingual (A 4) views. B. Paratype, WAM 13.5.14, left upper lateroposterior tooth; labial (B 1), basal (B 2), profile (B 3), and lingual (B 4) views. C. Paratype, WAM 13.5.15, left upper lateroposterior tooth; labial (C 1), profile (C 2), basal (C 3), and lingual (C 4) views. D. Paratype, WAM 13.5.19, a1?; labial (D 1), basal (D 2), profile (D 3), and lingual (D 4) views. E. Paratype, WAM 13.5.13, right upper lateroposterior tooth; labial (E 1), profile (E 2), basal (E 3), and lingual (E 4) views. F. Holotype, WAM 13.5.17, left a2?; labial (F 1), profile (F 2), lingual (F 3), and basal (F 4) views. G. Paratype, UM-BET 6, right upper lateroposterior tooth; basal (G 1), lingual (G 2), labial (G 3), and profile (G 4) views. H. Paratype, WAM 13.5.16, right lower lateroposterior tooth; labial (H ), profile (H ), lingual (H ), and basal (H ) views.