Tofino Basin elasmobranch teeth are characterized by a prominent principal cusp and a base (or root) positioned approximately central under the base of the tooth crown. The tooth crown of all the Tofino Basin elasmobranchs has a flanged occlusal crest or cutting edge and is either inclined lingually or compressed labio-lingually (e.g., many of the Superorder Squalomorphii sharks). Examples of tooth forms and terminologies (Figure 3) include Raja sp. (Figure 3.1 and Figure 3.2), a form similar to Isurolamna (Figure 3.3), and Family Squalidae (Figure 3.4 and Figure 3.5).
Determining the type of tooth base (root) and its vascularization are important to complete identification. Figures 3.6 to Figure 3.12 show examples of tooth vascularization. When the tooth base was absent, identification (using binomial systematics) was limited (e.g., only to the Superorder Squalomorphii level) or not possible.
Systematics and list of elasmobranch teeth (following Cappetta 1987):
Class Chondrichthyes
Huxley, 1880
Subclass Elasmobranchii
Bonaparte, 1838
Cohort Euselachii
Hay, 1902
Subcohort Neoselachi
Compagno, 1977
Superorder Squalomorphii
Compagno, 1977
Order Squaliformes
Goodrich, 1909
Family Squalidae Bonaparte, 1838
Family Squalidae indet. Forms A, B, C, D, and E
Family Squalidae indet., Form A
Family Squalidae indet., Form B
Family Squalidae indet., Form C
Family Squalidae indet., Form D
Family Squalidae indet., Form E
Order Hexanchiformes
de Buen, 1926
Suborder Hexanchoidei
Garman, 1913
Family Hexanchidae
Gray, 1851 or Family Heptranchidae
Barnard, 1925
Suborder Hexanchoidei indet., Forms A, B, and C
Suborder Hexanchoidei indet., Form A
Suborder Hexanchoidei indet., Form B
Suborder Hexanchoidei indet., Form C
Superorder Galeomorphii
Compagno, 1973
Order Lamniformes
Berg, 1958
Family Lamnidae
Müller and Henle, 1938
?Isurolamna sp. A
Order Carcharhiniformes
Compagno, 1973
Family Scyliorhinidae Gill, 1862
Family Scyliorhinidae indet., Form A
Superorder Batomorphii
Cappetta, 1980
Order Rajiformes
Berg, 1940
Suborder Rajoidei
Garman, 1913
Family Rajidae
Bonaparte, 1831
Raja sp. A
Unidentified elasmobranch teeth, Forms A, B, C, D, and E
Unidentified elasmobranch tooth, Form A
Unidentified elasmobranch tooth, Form B
Unidentified elasmobranch tooth, Form C
Unidentified elasmobranch tooth, Form D
Unidentified elasmobranch tooth, Form E
Superorder Squalomorphii
Compagno, 1973
Order Squaliformes
Goodrich, 1909
Family Squalidae
Bonaparte, 1834
Family Squalidae indet., Forms A, B, C, D, and E
Remarks: Tofino Basin Squalidae teeth are moderately robust, commonly compressed labio-lingually, slightly convex centrally (not flattened), and asymmetric. Each margin has a well-developed occlusal flange or cutting edge that may be serrated. One margin is longer than the other and convexly curved where it thins to a point basally. The cusp is broad, triangular, commonly angled (some may be more upright), and has apical shadow. The inline is opaque and a similar shape to the outline, is greater than one half to three-quarters the tooth height but does not reach the cusp apex; and has a rounded arc-like apex with approximately vertical striations between the inline and outline apices. The tooth outline is weakly textured with common, short, and irregular striations and may have basal vertical cracks. The base of the tooth is lobed, rarely preserved whole, and basally commonly breaks irregularly. The tooth base (root) is rarely preserved intact.
Greatest variations in the teeth (Forms A to E) include the type of margins, inline striations, cusp curvature and symmetry, and presence (or not) of a distal heel and base with central foramen on the lingual face. Many of the teeth occur together in the same samples suggesting that some may be the same species and that tooth heterodonty may be present.
Distinct characters of the Tofino Basin Squalidae teeth are the apron and lobed tooth base. Elasmobranch teeth from the Family Squalidae, Subfamily Squalinae, have a flanged primary cusp and a tooth apron and base usually with one lobe. Tooth base aprons with more than one lobe are known in Early Miocene to Recent Squalinae genera Etmopterus Rafinesque, 1810 and Squaliolus Smith and Radcliff 1912 (see Cappetta 1987, p. 52-64) but these do not resemble the shorter apron and multi-lobe arrangement of the Tofino Basin specimens.
Two deep-sea subtypes, curved fibrous triangle and fibrous triangle convex margins both Tway et al. 1985 have margins, a fibrous outline texture, and outline shape that is similar to the Tofino Basin Squalidae forms. Also, many of the Tofino Basin specimens similarly break irregularly at the tooth base region. The two deep-sea "fibrous" subtypes are known to range in samples from the middle Eocene to the lower Miocene. The Tofino Basin specimens differ by showing better development of the apron and have a tooth base characteristic of Squalidae.
Squalidae teeth almost identical to Tofino Basin specimens are observed in Cretaceous samples from the Queen Charlotte Islands and the Upper Cretaceous Nanaimo Group, Vancouver Island, and Gulf Islands. Reworked Cretaceous to middle Eocene ichthyoliths in the Tofino Basin are indicated following significant regional tectonic activity and a prominent unconformity that occurred during the interval. The occurrences of west coast Squalidae multi-lobate tooth aprons suggest that this feature was present in Cretaceous forms.
Most Holocene Squalinae are bathyal except for a few species of Squalus that live on the continental shelf (Cappetta 1987, p. 53).
Family Squalidae indet., Form A
Figure 4
a9/b8±12/c(12,13)+19/d14+19/e1/f(4a+b)+9+11+14/g7+8/h3/i3,4/j3,4,10/k5,8,9/m0/n≥1/p0/q1,9,10/r1,2/s1,2/t4/z0
Characters: Tooth asymmetric, compressed labio-lingually, with a primary cusp, distal heel, basal region with a shallow three or greater lobed apron, and one (possibly more) foramen on lingual face of base (root) between basal crown lobes. Tooth primary cusp with both margins convex, one margin basally convex and significantly longer; and a broad flanged lateral occlusal crest where basally on the long margin it thins to a point and curves inwards; apex with lateral and apical shadow; inline apical striations short. Basal labial crown face and base (root) fragmented or not preserved.
Remarks: These teeth are distinctive by having both margins convex and one convex margin that is longer and thins to a point basally. Also, the three or greater lobed basal apron is characteristic as are the one or more foramen on the base between the lobes.
Occurrence: 2 approximately complete specimens, 5 other specimens, 9+ basal fragments; 12+ Form A or B fragments; Hesquiat Peninsula (Oligocene) and one small specimen from offshore Shell Canada well Zeus D-14 (in Miocene sediments that are probably reworked); Squaloid Teeth ichthyolith interval; Cretaceous to middle Eocene, reworked and deposited into Oligocene strata.
Family Squalidae indet., Form B
Figure 5
a9/b8±12/c12+19/d19/e1/f(4a+b)+9+11+14/g7+8/h3/i4/j2,6/k5,8,9/m0/n≥1/p0/q9,10/r1,2/s1,2/t4/z0
Characters: Tooth asymmetric; flanged occlusal crest on one margin convex (especially basally) and may slightly widen centrally and narrow to a point basally, and on second margin straight or slightly concave; first margin longer than second; apical inline striations restricted to lower half.
Remarks: Form B only differs from Family Squalidae indet., Form A by having a straight or slightly concave second margin.
Occurrence: 2 specimens; Hesquiat Peninsula and Flores Island; Squaloid Teeth ichthyolith interval; Cretaceous to middle Eocene, reworked and deposited into Oligocene strata.
Family Squalidae indet., Form C
Figure 6
a9/b8±12/c14+19/d±13+19/e1/f(4a+b)+9+(11,12)+14/g7+8/h3/i2,3,9/j2,3,4,9/k5,8,9/m0/n≥1/p0/q9,10/r1/s1,2/t4/z0
Characters: Tooth asymmetric; with a primary cusp and part of distal heel; angle between distal heel and primary cusp narrow (approximately 20-30°) and V-shaped. Primary cusp first margin straight to slightly convex and basally angled to form the V-shape; other margin straight to slightly convex apically and distinctively basally convex where flanged margin thins to a point. Cusp striations may extend from inline apex into upper half but not to outline.
Occurrence: 4 specimens; Hesquiat Peninsula and offshore well Shell-Anglo Pluto I-87; Squaloid Teeth ichthyolith interval; Cretaceous to middle Eocene, reworked and deposited Oligocene strata.
Family Squalidae indet., Form D
Figure 7
a9/b8+11±12/c14+19/d19/e1/f(4a+b)+9+11+14/g7+8/h3/i2,9/j2,4,9/k5,8,9/m0/n≥0.8/p0/q9,10/r1/s1,2/t4/z0
Characters: Tooth asymmetric; with a primary cusp and part of distal heel; angle between heel and primary cusp moderately broad (approximately 90°) and U-shaped. Primary cusp first margin straight to slightly sigmoid and basally angled to form the U-shape; other margin approximately straight to slightly basally convex where basal flanged margin thins. Cusp striations may extend from inline apex into upper half but not to outline.
Occurrence: 2 specimens; Hesquiat Peninsula; Squaloid Teeth ichthyolith interval; Cretaceous to middle Eocene, reworked and deposited into Oligocene strata.
Family Squalidae indet., Form E
Figure 8
a9/b8±12/c(12,13)+(16, 17)+19/d(1,16,17)+19/e1/f(4a+b)+9+11+14/g7+8/h3/i4/j2,3/k5,8,9/ m0/n≥1/p0/q9,10/r1,2/s1,2/t4/z0
Characters: Tooth asymmetric; with one or both margins serrated/crenulated; flanged occlusal crest on one margin slightly sigmoid to convex (especially basally), widens centrally and narrows to a point basally, and on second margin much shorter and straight or evenly convex; striations between apical inline and outline restricted to lower half.
Remarks: The flanged convex margin that curves inwards to a point basally, makes this Form E similar to Family Squalidae indet., Forms A and Family Squalidae indet., B and differs by having a serrated margin.
Occurrence: 5 specimens, 2 questionable specimens; 3 serrated margin fragments; Hesquiat Peninsula and offshore wells Shell-Anglo Harlequin D-86 and Pluto I-87; Squaloid Teeth ichthyolith interval; Cretaceous to middle Eocene, reworked and deposited into upper Eocene to lower Miocene strata.
Order Hexanchiformes de Buen, 1926
Suborder Hexanchoidei
Garman, 1913
Family Hexanchidae Gray, 1851 or Family Heptranchidae
Barnard, 1925
Suborder Hexanchoidei indet., Forms A, B, and C
Remarks: Tofino Basin Family Squalidae and Hexanchoidei teeth are similarly moderately robust, compressed labio-lingually with a slightly convex central region, and have well- developed occlusal flanged margins or cutting edges. Tofino Basin Hexanchoidei teeth differ by having one crown margin that is convex with a distinct basal angled termination of the flanged occlusal crest that pinches inward and then may slightly curve upwards (slightly hooked). The other flanged margin is concave or straight and basally terminates with a straight across to irregular break. Margins are commonly of a similar length. The tooth crown commonly irregularly breaks basally without preservation of the root/base.
Suborder Hexanchoidei indet., Form A
Figure 9
a9/b8±12/c14+19/d19/e1/f(4a+b)+9+(11,12)+14/g7+8/h1,3,4,5/i3,9/j6,7,8/k5,8,9/m0/n~≥1/p0/q9,10/r1/s1,2/t4/z0
Characters: Tooth moderately robust, asymmetric and curved; with one margin evenly convex to sigmoid and slightly hooked and thinner at base; other margin variably concave and usually longer; both margins with prominent flanged occlusal crest or cutting edge; striations between apical inline and outline may extend into upper half; height to width ≥1; apex moderately acute (not acute, not rounded).
Remarks: A pronounced concave margin is distinct to this form.
Occurrence: 7 specimens; Hesquiat Peninsula and Flores Island; and offshore well Shell-Anglo Pluto I-87; Squaloid Teeth ichthyolith interval; Cretaceous to middle Eocene, reworked and deposited into upper Eocene to Oligocene strata.
Suborder Hexanchoidei indet., Form B
Figure 10
a9/b8±12/c13±14+19/d19/e1/f(4a+b)+9+12+14/g7+8/h1,5,4,3/i2,4/j2,6/k5,8,9/m0/n~≥0.7/p0/q9,10/r1,2/s3/t4/z0
Characters: Tooth asymmetric, robust, centrally and basally moderately inflated; apex blunt to rounded; striations between apical inline and outline may extend from the crown base into upper three quarters with some almost to outline apex; flanged margins of similar lengths and widen basally. One margin straight to convex except at most basal region where it abruptly curves inward (at about 135°) to a point at inline; other margin straight or equally concave and basally breaks straight across at about 90°; height approximately equals width.
Remarks: This robust tooth has margins commonly of the same length with one straight to basally convex and the other straight to slightly concave. Form B differs from Suborder Hexanchoidei indet., Form A by having a rounded apex, being shorter with width approximately equal to height, and by not having one concave flanged margin.
Occurrence: 16 specimens; Hesquiat Peninsula; Squaloid Teeth ichthyolith interval; Cretaceous to middle Eocene, reworked and deposited into Oligocene strata.
Suborder Hexanchoidei indet., Form C
Figure 11
a9/b8±12/c19/d19/e1/f(4a+b)+9+(12,13)+14/g7±8/h1,5/i2,3/j2,3/k5,8/m0/n≥1/p0/q9,10/r1/s1,2/t4/z0
Characters: Tooth approximately symmetric; inflated centrally and basally; flanged occlusal crest not as developed (thin apically and slightly widens basally); both margins slightly evenly convex or straight; about same length striations between apical inline and outline extend into upper half and possibly upper quarter; apex sharp to slightly rounded.
Occurrence: 1 specimen; Hesquiat Peninsula; Squaloid Teeth ichthyolith interval; Cretaceous to middle Eocene, reworked and deposited into Oligocene strata.
Superorder Galeomorphii
Compagno, 1973
Order Lamniformes
Berg, 1958
Family Lamnidae
Müller and Henle, 1838
Isurolamna
Cappetta, 1976
?Isurolamna sp. A
Figure 12
a9/b8/c19/d19/e1/f1/g7/h0/i9/j9/k1/m0/n>1.2/p0/q0/r0,1/s0/t4/z1,2
Characters: Tooth triangular; enameloid smooth; about 2 cm high and 1 cm wide; sigmoidal in mesial view and lingually angled at about 40-50°; with a flattened labial face and an inflated cambered lingual face; occlusal crest or cutting edge closest to labial face and does not quite reach crown base (Figure 5.1.2); basal labial face flares and follows the root slightly beyond cutting edge terminus. Root with prominent lingual protuberance and an elliptical foramen that extends centrally down across lingual root face to root subsurface where opening is largest; lingual face approximately vertical, rounded basally, and evenly convex-curved laterally; labial face vertical and centrally concave.
Remarks: Cappetta (1987, p. 95-96) describes both Isurolamna Cappetta, 1976 and Isurus Rafinesque, 1810. Isurolamna is favoured because the lingual root face is significantly protruding and has an elliptical central foramen that extends to the base surface. Crown lateral cusplets may or may not be present in this genus and could not be observed because the tooth/root is broken at both lateral margins. The identification is questionable because of the fragmentary condition of the specimen.
Isurolamna is known in the upper Paleocene-lower Eocene and Isurus in the upper Paleocene-Recent (Cappetta 1987).
Occurrence: 1 specimen; Rafael Point, Flores Island; upper Eocene to lower Oligocene.
Order Carcharhiniformes
Compagno, 1973
Family Scyliorhinidae
Gill, 1862
Family Scyliorhinidae indet., Form A
Figure 13
a9/b2+8+12/c19/d19/e1/f1/g6+7+8/h0/i6,7/j6,7/k0,1/m0/n0,~1/p0/q0/r0/s3/t4/z0
Characters: Tooth crown lingually inclined; lingually curved and more basally convex. Tooth base missing. Crown with single cusp; margins slightly flare basally and laterally; lingual and labial faces with multiple long lines/ridges that bifurcate basally especially on labial face; lingual face slightly inflated basally; occlusal crest present at both margins; apex rounded.
Remarks: Tooth crown prominent lines/ridges, tall principal cusp that broadens basally, inflated lingual face that arches basally, and a labial face that overhangs the root/base (Figure 13.1.2) characterize many teeth of the Scyliorhinidae. The identification is indeterminate because of incomplete specimens, unknown cusplets, and the missing root/base.
Occurrence: 2 specimens, 1 fragment; Hesquiat Peninsula; Squaloid Teeth ichthyolith interval; Cretaceous to middle Eocene, reworked and deposited into Oligocene strata.
Superorder Batomorphii Cappetta, 1980
Order Rajiformes
Berg, 1940
Suborder Rajoidei
Garman, 1913
Family Rajidae
Bonaparte, 1831
Raja
Linnaeus, 1758
Raja sp. A
Figure 14
a4/b6+8/c2/d2+8/e1/f0/g1,2/h1/i1,2/j2+(11,12)/k1/l3/m0/n1
Characters: Crown with one rounded apex and flattened apical surface; occlusal crest terminates on upper surface of a rounded and inflated labial rim; inline similar shape to outline with abundant long striations that radiate from it. Crown longer than wide (immature specimens may be wider than long) and with prominent rounded basal rim that overhangs the root base (especially labially) and curves under to subcrown collar and crown/root junction. Root base prominently flares basally; base wider than crown mesiodistally; labial face slightly higher than lingual face; and with distinct central, basal, lingual, rounded notch and prominent rounded and arched labial notch. Holaulacorhize root vascularization; root base with two separate kidney-shaped lobes each with flat surface and one or more foramina in central basal concavity.
Remarks: Probable juvenile or immature teeth with a less developed crown cusp or peak are shown in Figure 14.3.1 and Figure 14.4.1.
Occurrence: 3 specimens and 1 questionable specimen; offshore wells Shell-Anglo Zeus I-65 and Apollo J-14; offshore core END-76B-6E; Shadowed Cone ichthyolith Zone; upper Miocene and Pliocene.
Unidentified elasmobranch teeth
Forms A, B, C, D, and E
Some elasmobranch teeth were not completely identified because they were rare and without a tooth base or root. Observing the character of the tooth base/root is required to complete identification to genus or species. Preliminary identifications (including illustration, brief description, and utilitarian code identification) are provided for five elasmobranch teeth (Forms A to E) with distinctive crowns. Future studies hopefully will result in the location of complete specimens so that comparative identifications can be made.
Unidentified elasmobranch tooth, Form A
Figure 15
a9/b2+8+12/c19/d19/e1/f(4a+b)+9+(12,13)+14/g4+7+8/h1,2,4,5/i3,9/j6,7/k5,8,9/m0/n0,≥1/p0/q9,10/r1/s1/t4/z0
Characters: Tooth asymmetric and curved; with one margin concave and longer than slightly convex second margin; striations between apical inline and outline may extend into upper half; both lingual and labial surfaces with ridges that are basally prominent, greater in number, and basally bifurcate; one face has the most ridges that extend from crown base to about three quarters height of tooth but not to occlusal crest, may be discontinuous and irregular.
Remarks: The missing tooth base prevents specimen identification.
Occurrence: 1 specimen; Hesquiat Peninsula; Squaloid Teeth ichthyolith interval; Cretaceous to middle Eocene, reworked and deposited into Oligocene strata.
Unidentified elasmobranch tooth, Form B
Figure 16
a9/b8/c19+20/d19+20/e1/f(4a+b)+9+(11,12)+14/g7/h1,5/i6,7,8,9/j6,7,8,9/k8/m0.05-0.35/n1.2-2.0/p0/q9,10/r1/s3/t4/z0
Characters: Tooth labio-lingual curvature convex/concave; height to width ratio >1.5; one margin concave to sigmoid; other margin concave, slightly sigmoid, or slightly convex; most curvature basal where flanged occlusal crest margins widen and flare outwards; striations between apical inline and outline; asymmetric and acute laterally in cross-section; apex rounded.
Remarks: These unidentified elasmobranch teeth differ from Family Squalidae indet., Form C by being narrower and by having both basal occlusal crest margins that flare outwards (forming concave basal margins), and do not thin to a point basally, and do not curve inwards basally and lingually. These specimens differ from the Family Squalidae forms by not having a textured outline with common, short, and irregular striations, and by having both basal margins that flare outwards.
Occurrence: 4 specimens and 1 fragment; Hesquiat Peninsula; Squaloid Teeth ichthyolith interval; Cretaceous to middle Eocene, reworked and deposited into upper Eocene to Oligocene strata.
Unidentified elasmobranch tooth, Form C
Figure 17
a9/b8±12/c13+19/d13+19/e1/f(4a+b)+9+(12,13)+14/g7+8/h1,5/i9,10/j9,10/k8,9/m0/n>1.5-3/p0/q9/r1/s3/t4/z0
Characters: Tooth labio-lingual curvature sigmoid/reverse sigmoid; height to width ratio >1.5; lingual face more inflated and rounded (especially at base) than flatter labial face; margins sigmoid or reverse sigmoid; flanged occlusal crest on both margins and thin to a point both apically and basally (widest centrally) and basally curve inwards lingually; thick and common long apical striations fill much of area between inline and outline; asymmetric and acute laterally in cross-section; apex rounded; outline with faint irregular ridges/lines.
Remarks: These teeth have a similar outline texture, shape of the inline, and striations between the inline apex and outline as many of the Family Squalidae teeth. They differ by not being compressed labio-lingually, by having basal lingual curvature, and by being more upright.
Occurrence: 5 specimens; Hesquiat Peninsula and Nootka Island; Squaloid Teeth ichthyolith interval; Cretaceous to middle Eocene, reworked and deposited into Oligocene strata.
Unidentified elasmobranch tooth, Form D
Figure 18
a9/b8±12/c19/d19/e1/f±(4a+b)+9+(11,12)+14/g7±8/h1,5,4/i2,6,9/j2,6,9/k8,9/m0/n>2/p0/q9,10/r1/s0,1,2/t4/z0
Characters: Tooth labio-lingual curvature sigmoid/reverse sigmoid; height to width ratio >2; lingual face more inflated and rounded (especially at base) than flatter labial face; margins concave, sigmoid, or straight with thin flanged occlusal crest; common apical striations between inline and outline; asymmetric and acute laterally in cross-section; outline may have faint irregular ridges/lines.
Remarks: These teeth have a similar shape to unidentified elasmobranch tooth, Form C but are smaller, thinner, and not as robust. The tooth apex is commonly damaged but is probably acute because the tooth is tall and narrow.
Occurrence: 4 specimens; Hesquiat Peninsula and Nootka Island; Squaloid Teeth ichthyolith interval; Cretaceous to middle Eocene, reworked and deposited into upper Eocene to Oligocene strata.
Unidentified elasmobranch tooth, Form E
Figure 19
a9/b2+8+12/c19+20/d19+20/e1/f1/g6+7+8/h1,5/i7/j7/k0,1/m0/n~1/p0/q0/r0/s2/t4/z0,1
Characters: Small triangular tooth (<0.4 mm), with base missing. Tooth crown flared and extended laterally; with single cusp and acute apex; labial face with central ridge and occlusal crest at both margins; lingual face with ridges/lines, more common basally and may bifurcate basally; lingual face base with protrusion.
Occurrence: 1 specimen, offshore well Shell-Anglo Prometheus H-68; Bulbous Base ichthyolith Zone; Miocene.