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Otoliths of Fishes from the Tertiary Formations of New Zealand. By G. Allan Frost, F.L.S., F.G.S. [Read by J. Marwick before the Wellington Philosophical Society, 14th October, 1931; received by the Editor, 27th November, 1931; issued separately, 28th February, 1933.] Through the courtesy of Dr. J. Marwick, of the Geological Survey Office, Wellington, New Zealand, I have received for description a remarkable series of fossil otoliths from various localities in New Zealand which, although their stratigraphical value may be of slight extent, are palaeontologically of great interest. With the exception of three, the 25 species figured are all new forms, distinct from any yet described. The type examples are clear cut and in excellent preservation, and in some cases show a striking likeness to the otoliths of present-day New Zealand fishes. It is desirable when describing fossil otoliths to compare them, if possible, with those of recent fishes, and not to rely on the determinations of previous authors, which are in some instances inaccurate. This has been done with the present examples, and all have been carefully compared with recent forms in the author's collection. Among the deep-sea forms described there are 13 species of scopelid otoliths—four belong to the Macruridae and one is referred to the family Gadidae. Of the remainder, three belong to the family Congridae, one to the Pleuronectidae, two are from percoids, and there is one example from a species of the order Scleroparei. The majority are from the Tutamoe Survey District; of these, 11 are from the Tutamoe Beds, Island Creek, and 11 from the Ihungia Beds, four species occurring in both places. Two are from Waingake, Patutahi Survey District, three from Waingaromia Survey District. Of the latter, one occurs also in the Tutamoe Beds and one in the Ihungia Beds. One comes from the Otapiri grey marls, one from the marl of Terakohe, Takaka, and three from the Mimi Survey District, one of these also appearing in the Tutamoe Beds. Order Iniomi, fam. Myctophidae. Otolithus (Scopelus) marwicki sp. n. (Figs. 1–3, inner side; Figs. 2–4, outer side). Description: Dimensions 4 mm. × 3 mm. Shape ovate, outer side convex, with vertical groove from the dorsal rim ending above the ventral rim; inner side flat, rostrum and antirostrum present, dorsal rim domed, with posterior part lower than the anterior, ventral rim slightly serrated. Sulcus bold, divided, cauda rectangular with groove below. Material: Three examples, including Holotype.

Distribution: Loc. 1358, Ihungia Mudstone, Haupatua Stream; and Loc. 1349, Ihungia Beds, N.W. tributary Konganaru Stream, Tutamoe Survey District (type). Age: Hutchinsonian, Upper Oligocene. Remarks: This species is remarkable for the vertical groove of the outer side, resembling in this feature a scopeloid otolith described by Bassoli (1906 XII, p. 50, Pl. II, Fig. 24) O. (Berycidarum) sulcatus Bass, from the Miocene of Monte Gibio, Italy. Otolithus (Scopelus) mammatus sp. n. (Fig. 5, inner side; Fig. 6, outer side). Description: Dimensions 3 ½ mm. × 2 ¾ mm. Shape ovate, outer side convex, umbonated, groove above rostrum; inner side flat, dorsal rim domed anteriorly, ventral rim slightly serrated, rostrum stout, no antirostrum. Sulcus with upper and lower angles, cauda circular. Material: Two examples, including Holotype. Distribution: Loc. 1262, Island Creek (Tutamoe Beds); Loc. 1349, Ihungia Beds, tributary Konganaru Stream, Tutamoe Survey District (type). Age: Awamoan, Lower Miocene; Hutchinsonian, Upper Oligocene. Remarks: The general shape resembles that of O. (Scopelus) tuberculatus Bassoli (1906 XII, Pl. 11, Fig. 25–26) from the Miocene and Pliocene of Italy. It differs, however, in the circular cauda. Otolithus (Scopelus) excisus sp. n. (Fig. 7, inner side; Fig. 8, outer side). Description: Dimensions 4 ½ mm. × 3 ½ mm. Shape ovate, outer side slightly convex, furrowed on dorsal and ventral rims; inner side very flat. Dorsal rim low with posterior angle; rostrum, antirostrum, and concave excisura present. Sulcus horizontal, long, cauda oblong. Material: Four examples, including Holotype. Distribution: Loc. 1372, Section A, Block 13, Waingaromia Survey District (type); Loc. 1349, Ihungia Mudstone, tributary Konganaru Stream, 300–500 feet, below top. Age: Hutchinsonian, Upper Oligocene. Remarks: This species is noticeable for the flatness of the inner side, and for the large excisura. Otolithus (Scopelus) tutamoensis sp. n. (Fig. 9, inner side; Fig. 10, outer side). Description: Dimensions 4 mm. × 3 mm. The shape resembles that of O. (Scopelus) marwicki, it differs in the truncation of the dorsal area, and in the outer side. Material: Six examples, including Holotype.

Distribution: Loc. 1301, Ihungia Mudstone, N.W. tributary Konganaru Stream (type); Loc. 1358, Ihungia Mudstone, Haupatua Stream, Tutamoe Survey District. Age: Hutchinsonian, Upper Oligocene. Otolithus (Scopelus) circularis Frost (Fig. 11, inner side; Fig. 12, outer side). 1924. Frost, Trans. N.Z. Inst., vol. 55, p. 608, Pl. 61, Fig. 2. Description: Dimensions 3 mm. × 3 mm. A circular flat form described by the author from the Awamoan Beds of Target Gully. Material: Four examples. Distribution: Loc. 1358, Ihungia Mudstone, Haupatua Stream, Tutamoe Survey District. Target Gully, Oamaru (type). Age: Hutchinsonian, Upper Oligocene; and Awamoan, Miocene. Remarks: The example figured is a much finer specimen than the original Holotype, especially in the clearer lines of the sulcus. Otolithus (Scopelus) concentricus sp. n. (Fig. 13, inner side; Fig. 14, outer side). Description: Dimensions 2 mm. × 2 mm. Shape circular, outer side raised and showing lines of growth, inner side flat, rostrum, antirostrum, and excisura present. Angle between dorsal and posterior rims, ventral rim strongly dentated. Sulcus typically scopelid, with double lines below the cauda. Material: Nine examples, including Holotype. Distribution: Loc. 1262, Tutamoe Beds, Island Creek (eight examples) (type); Loc. 1358, Ihungia Mudstone, Haupatua Stream, Tutamoe Survey District. Age: Awamoan, Lower Miocene; Hutchinsonian, Upper Oligocene. Otolithus (Scopelus) regularis sp. n. (Fig. 15, inner side; Fig. 16, outer side). Description: Dimensions 2 ½ mm. × 2 ¼ mm. Shape circular, outer side convex, furrowed on dorsal and ventral rims; inner side flat, angle between dorsal and posterior rims, rostrum broad, no antirostrum or excisura. Sulcus with angle of lower margin, ostium and cauda equal in length, cauda wider than ostium. Material: The unique Holotype. Distribution: Loc. 1372, Section A, Block 13, Waingaromia Survery District. Age: Hutchinsonian, Upper Oligocene. Remarks: Resembles the otolith of the recent species Myctophum caninianus (see Frost, 1926, XVIII, p. 468, Pl. XX, Fig. 13).

Otolithus (Scopelus) cassidiformis sp. n. (Fig. 17, inner side; Fig. 18, outer side). Description: Dimensions 2 ½ mm. × 2 ¼ mm. Shape circular, outer side convex, smooth, inner side flat; dorsal rim high anteriorly, with posterior angle, ventral rim curved and serrated, posterior rim curved, anterior rim oblique, rostrum and slight antirostrum present, no excisura. Sulcus with lower angle, ostium wide, with straight front, cauda narrow, ovate, lower margin with double lines. Material: The unique Holotype. Distribution: Loc. 1349, Ihungia Beds, tributary Konganaru Stream, Tutamoe Survey District. Age: Hutchinsonian, Upper Oligocene. Otolithus (Scopelus) konganaruensis sp. n. (Fig. 19, inner side). Description: Dimensions 5 mm. × 4 ½ mm. Shape circular, outer side convex, smooth, inner side very convex, angle between dorsal and posterior rims, ventral rim serrated. Rostrum, antirostrum, and an angular excisura present. Sulcus very wide, straight, division ill-defined. Material: Two examples, including Holotype. Distribution: Loc. 1301, Ihungia Mudstone, tributary to Konganaru Stream, 300 to 500 feet below top, Tutamoe Survey District. Age: Hutchinsonian, Upper Oligocene. Otolithus (Scopelus) incisus sp. n. (Fig. 20, outer side). Description: Dimensions 5 mm. × 4 mm. Shape ovate, umbonated and furrowed on all rims. Prominent rostrum and small pointed antirostrum, with deep short groove below it. Material: The unique Holotype. Distribution: Loc. 1351, three-quarters of a mile above Waingaki, Patutahi Survey District. Age: Awamoan, Lower Miocene. Remarks: Owing to this example being imbedded in the matrix, it is only possible to describe the features of the outer side. It differs from the remainder of the scopelid otoliths described in shape, and in the presence of the deep slit below the antirostrum. Otolithus (Scopelus) laminatus sp. n. (Fig. 21, inner side; Fig. 22, outer side). Description: Dimensions 3 ½ mm. × 3 mm. Shape circular, without rostrum, antirostrum, or excisura. Both sides flat, outer side furrowed, anterior part of ventral rim serrated. Sulcus wide, nearly straight, divided, extending entirely across the inner side, ostium longer than cauda. Material: The unique Holotype. Distribution: Loc. 1262, Tutamoe Beds, Island Creek, Tutamoe Survey District. Age: Awamoan, Lower Miocene.

Fig. 1–4.—Otolithus (Scopelus) marwicki sp. n. Fig. 5–6.—Otolithus (Scopelus) mammatus sp. n. Fig. 7–8.—Otolithus (Scopelus) excisus sp. n. Fig. 9–10.—Otolithus (Scopelus) tutamoensis sp. n. Fig. 11–12.—Otolithus (Scopelus) circularis Frost. Fig. 13–14.—Otolithus (Scopelus) concentricus sp. n. Fig. 15–16.—Otolithus (Scopelus) regularis sp. n. Fig. 17–18.—Otolithus (Scopelus) cassidiformis sp. n. Fig. 19.—Otolithus (Scopelus) konganaruensis sp. n. Fig. 20.—Otolithus (Scopelus) incisus sp. n. Fig. 21–22.—Otolithus (Scopelus) laminatus sp. n. Fig. 23–24.—Otolithus (Scopelus) mimiensis sp. n

Fig. 25–26.—Otolithus (Scopelus) ellipticus sp. n. Fig. 27.—Otolithus (Congridarum) carinatus sp. n. Fig. 28.—Otolithus (Congridarum) otapiriensis sp. n. Fig. 29–30.—Otolithus (Congridarum) mysticus sp. n. Fig. 31.—Otolithus (Macruronus) pukeuriensis Frost. Fig. 32.—Otolithus (Pleuronectidarum) orbicularis sp. n. Fig. 33–34.—Otolithus (Macrurus) sagittiformis sp. n. Fig. 35–36.—Otolithus (Macruridarum) latisulcatus sp. n. Fig. 37–38.—Otolithus (Macrurus) toulai Schubert. Fig. 39–40.—Otolithus (Physiculus) terakohensis sp. n. Fig. 41–42.—Otolithus (Sparidarum) sigmoidalis sp. n. Fig. 43–44.—Otolithus (Serranidarum) serratus sp. n. Fig. 45–46.—Otolithus (Cyclopterus) trifoliatus sp. n.

Otolithus (Scopelus) mimiensis sp. n. (Fig. 23, inner side; Fig. 24, outer side). Description: Dimensions 3 mm. × 2 mm. Shape ovate, outer side convex, furrowed, inner side flat. Dorsal rim low, forming angle with the posterior rim, ventral rim strongly serrated, rostrum and antirostrum prominent and equal, excisura large. Sulcus straight, divided, ostium long, constricted anteriorly, distended medianly, cauda short and narrower than the ostium. Material: The unique Holotype. Distribution: Loc. 1125, coast between Waikiekie Stream and Mangapuketea Stream, Mimi Survey District. Age: Taranakian, Upper Miocene. Otolithus (Scopelus) ellipticus sp. n. (Fig. 25, inner side; Fig. 26, outer side). Description: Dimensions 3 mm. × 2 ½ mm. Shape elliptical, both sides flat, a slight angle between dorsal and posterior rims, ventral rim curved, passing into the rounded anterior rim, which is indented above the sulcus. The sulcus is wide and straight, and has a ridge above it, the ostium is constricted anteriorly, and the cauda is large and rectangular. Material: The unique Holotype. Distribution: Loc. 1351, three-quarters of a mile above Waingake, Kauwaewaka Stream, Patutahi Survey District. Age: Awamoan, Lower Miocene. Order Apodes. Otolithus (Congridarum) carinatus sp. n. (Fig. 27, inner side). Description: Dimensions 3 mm. × 2 ½ mm. Shape irregular, biconvex, dorsal rim high, pointed, sloping forward; ventral rim deepest anteriorly, posterior rim pointed, anterior rim rounded. Sulcus oblique, with groove opening on dorsal rim. Material: The unique Holotype. Distribution: Loc. 1236, Ihungia Beds, Muddy Creek, Tutamoe Survey District. Age: Hutchinsonian, Upper Oligocene. Remarks: Resembles in shape that of the recent species Congromureana balaerica, and in the sulcus that of Uroconger lepturus (Richardson) (Frost, 1926, XVII, Pl. 11, Figs. 15–11). Otolithus (Congridarum) otapiriensis sp. n. (Fig. 4, inner side). Description: Dimensions 7 mm. × 5 mm. Shape ovate, biconvex, dorsal rim straight, ventral rim deepest anteriorly, posterior rim pointed, anterior rim consists of stout rostrum. Sulcus wide, open between dorsal rim and point of rostrum, end rounded. Material: Three examples, including Holotype.

Distribution: Loc. 256, Grey marls, Otapiri Creek, Southland. Age: Hutchinsonian, Upper Oligocene. Remarks: Resembles that of the recent species Uroconger lepturus (Rich.) (Frost, 1926, XVII, Pl. 11, Fig. 15–11). It differs in the less symmetrical shape, and in the width of the sulcus. Otolithus (Congridarum) mysticus sp. n. (Fig. 29, inner side; Fig. 30, outer side). Description: Dimensions 3 mm. × 2 mm. Shape ovate, biconvex, symmetrical. Sulcus filled in with colliculum, a slight depression leads to dorsal and anterior rims. Material: The unique Holotype. Distribution: Loc. 1272, Ihungia Beds, Island Creek, Tutamoe Survey District. Age: Hutchinsonian, Upper Oligocene. Order Anacanthini. Otolithus (Macruronus) pukeuriensis Frost (Fig. 31). 1924. O. (Merluccius) pukeuriensis Frost, Trans. N.Z. Inst., vol. 55, p. 609, Pl. 61, Fig. 7. Description: Dimensions 7 mm. × 3 mm. Material: Two examples. Distribution: Loc. 1262, Tutamoe Beds, Island Creek, Tutamoe Survey District; Loc. 1125, coast, Mimi Survey District; Oamaru series, Pukeuri. Age: Awamoan, Lower Miocene; Taranakian, Upper Miocene. Remarks: An example from Pukeuri was described by the author in 1924, and was referred to the genus Merluccius, but a later comparison shows that it resembles more closely the otolith of the recent species Macruronus novae-zeelandiae (see Frost, 1926, XVIII, 484, Pl. XXII, Fig 4–5), the anterior point being considerably higher than in Merluccius. Otolithus (Macrurus) sagittiformis sp. n. (Fig. 33, inner side; Fig. 34, outer side). Description: Dimensions 3 mm. × 2 mm. Shape that of an arrow head, outer side straight, furrowed, with a medianly placed umbo, from which a deep groove descends to the anterior rim. Anterior of otolith broad, posterior part pointed. Inner side flat, dorsal rim furrowed and serrated, a smooth surface is present above and below the sulcus. The sulcus extends across the inner side, lower margin with angle, ostium upwardly inclined, cauda curved, ending on ventral rim below the posterior point of the otolith; a small round colliculum is present in both the ostium and the cauda. Material: Two examples, including Holotype. Distribution: Loc. 1262, Tutamoe Beds, Island Creek, Tutamoe Survey District; Loc. 1293, Akiripuraho Stream, Waingaromia Survey District (type).

Age: Awamoan, Lower Miocene; Hutchinsonian, Upper Oligocene. Remarks: Resembles the otolith of the recent species Macrurus cœlorhynchus (see Frost, 1926, XVIII, 483, Pl. XXII, Fig. 3). Otolithus (Macruridarum) latisulcatus sp. n. (Fig. 35, inner side; Fig. 36, outer side). Description: Dimensions 4 mm. × 2 mm. Shape ovate, biconvex, anterior part high, posterior part pointed; anterior rim oblique, with two rounded projections. Sulcus enclosed, wide, with median upper angle, both ends rounded. Material: The unique Holotype. Distribution: Loc. 1262, Tutamoe Beds, Island Creek, Tutamoe Survey District. Age: Awamoan, Lower Miocene. Remarks: The shape is similar to those of the family Gadidae, but the sulcus resembles closely that of the recent species Trachyrhynchus trachyrhynchus of the family Macruridae (Frost, 1926, XVIII, Pl. 12, Fig. 1). Otolithus (Macrurus) toulai Schubert (Fig. 37, inner side; Fig. 38, outer side). E. Schubert, 1905, 55, Pl. XVI, Fig. 34–37. G. G. Bassoli, 1906, XII, Pl. 1, Fig. 13–14. G. Allan Frost, 1924, 55, p. 608, Pl. 61, Fig. 4. Material: One example (young), 2 ¼ mm. × 1 ½ mm. Distribution: Loc. 1262, Tutamoe Beds, Island Creek, Tutamoe Survey District. Age: Awamoan, Lower Miocene. Remarks: Described by Schubert from the tertiaries of Austro-Hungary, by Bassoli from the Miocene of Monte Gibio, Italy, and by the author from Pukeuri, Awamoa, Ardgowan, and Target Gully, New Zealand. Otolithus (Physiculus) terakohensis sp. n. (Fig. 39, inner side; Fig. 40, outer side). Description: Dimensions 2 ¼ mm. × 1 ½ mm. Shape irregular, outer side bulbous, anterior part raised, inner side flat; anterior part of dorsal rim high and rounded, ventral rim with three processes. Sulcus situated posteriorly to the dorsal elevation, contains angle on lower margin, ostium oval, cauda distended and open on posterior rim. Material: The unique Holotype. Distribution: Loc. 1286, marl, Terakohe, Takaka. Age: Hutchinsonian, Upper Oligocene.

Remarks: This otolith resembles that of the recent New Zealand species Physiculus bacchus (see Frost, 1926, XVIII, p. 488, Pl. XXII, Fig. 24) in the high dorsal projection and in the furcation of the posterior rim. It differs in the absence of similar processes anteriorly, also in the fore part of the sulcus. It resembles more closely the fossil form O. (Physiculus) bicaudatus Frost (1924, Trans. N.Z. Inst., 55, p. 608, Pl. 61, Fig. 5), described by the author from Pukeuri, New Zealand, but differs in the shape of the dorsal process, and in the sulcus. Order Heterosomata. Otolithus (Pleuronectidarum) orbicularis sp. n. (Fig. 32, inner side). Description: Dimensions 4 mm. × 3 ½ mm. Shape subcircular, biconvex; outer side smooth, inner side smooth, showing rings of growth. Sulcus open on anterior rim, ostium small, cauda not discernible. Material: The unique Holotype. Distribution: Loc. 1125, coast between Waikiekie Stream and Mangapuketa Stream, Mimi Survey District. Age: Taranakian, Upper Miocene. Remarks: Resembles the otolith of the recent species Peltorhamphus novae-zeelandiae (see Frost, 1930, vol. V, p. 234, Pl. IX, Fig. 12). Order Percomorphi. Otolithus (Sparidarum) sigmoidalis sp. n. (Fig. 41, inner side; Fig. 42, outer side). Description: Dimensions 3 mm. × 2 ½ mm. Shape ovate, outer side convex, furrowed on dorsal and ventral rims, inner side flat, with depression above sulcus. Dorsal rim highest anteriorly, posterior rim forms small angles with dorsal and ventral rims; a small pointed rostrum is present. Sulcus sigmoidal, direction horizontal, two angles on lower margin; ostium upwardly inclined, cauda longer than ostium. Material: Two examples, including Holotype. Distribution: Loc. 1236, Ihungia Beds, Muddy Creek; Loc. 1262, Tutamoe Beds, Island Creek, Tutamoe Survey District (type). Age: Hutchinsonian, Upper Oligocene; Awamoan, Lower Miocene. Remarks: This otolith resembles that of the recent berycoid species Polymixia japonica (see Frost, 1927, XIX, p. 440, Pl. VIII, Fig. 1), but differs in the upward inclination of the cauda, resembling in this respect that of the percoid species Micropterus salmonoides (see Frost, 1927, XX, p. 301, Pl. V, Fig. 19). The general characteristics appear to agree best with the otoliths of the family Sparidae, notably with those of Cantharus lineatus (see Frost, 1927, XX, p. 301, Pl. V, Fig. 17); it appears to be an early sparid form with berycid affinities.

Otolithus (Serranidarum) serratus sp. n. (Fig. 43, inner side; Fig. 44, outer side). Description: Dimensions 6 mm. × 3 mm. Shape long, ovate, outer side flat, furrowed on the dorsal and ventral rims, inner side flat. Dorsal rim curved and serrated, ventral rim deepest anteriorly, serrated, forms angle with posterior rim, anterior rim consists of broad rostrum. Sulcus straight, angle present on lower margin; ostium open on upper part of rostrum, cauda narrow, with curved and pointed end. Material: The unique Holotype. Distribution: Loc. 1262, Tutamoe Beds, Island Creek, Tutamoe Survey District. Age: Awamoan, Lower Miocene. Remarks: This species resembles in outline the otoliths of the recent species Centropomus undecimalis, it differs in the flatness of its form, and in the regular dorsal rim. The sulcus resembles those of the recent percoid fishes Pomatomus saltatrix and Arripis georgianus (see Frost, 1927, vol. XX, Pl. V, Figs. 5, 6, 7). Order Scleroparei. Otolithus (Cyclopterus) trifoliatus sp. n. (Fig. 45, inner side; Fig. 46, outer side). Description: Dimensions 2 mm. × 1 ½ mm. Shape subcircular, biconvex, outer side smooth, inner side with conchoidal depression above the sulcus. Sulcus biovate, enclosed and divided; ostium and cauda equal, in conjunction with the depression of the dorsal area they form a trefoil. Material: The unique Holotype. Distribution: Loc. 1262, Tutamoe Beds, Island Creek, Tutamoe Survey District. Age: Awamoan, Lower Miocene. Remarks: Resembles in all features the otolith of a young example of the recent species Cyclopterus lumpus (see Frost, 1929, vol. IV, p. 262, Pl. 111, Fig. 27). Literature. Schubert, R., 1905. Dic Fischotolithen des Osterr. Ungarischen. Tertiars, II. Macruriden und Beryciden. Jahrbuch der k.k. Geol. Reichsanst. LV, Wien., 1905, p. 613–638. Bassoli, G. G., 1906. Otoliti Fossili Terziari dell' Emilia. Rivista Ital. di Paleont. Anno XII, fasc. I, Perugia, 1906. Frost, G. Allan, 1924. Otoliths of Fishes from the Tertiary Formations of New Zealand. Trans. N.Z. Inst. 55, p. 605–614. —– 1926 (1). A Comparative Study of the Otoliths of the Neopterygian Fishes. Ann. and Mag. Nat. Hist., Series 9, XVII, p. 99–104. —– 1926 (2). Ann. and Mag. Nat. Hist., Series 9, XVIII, p. 465–482, p. 483–490. —– 1927. Ann. and Mag. Nat. Hist., Series 9, XX, p. 298–305. —– 1929. Ann. and Mag. Nat. Hist., Series 10. IV, p. 260–264. —– 1930. Ann. and Mag. Nat. Hist., Series 10, V, p. 231–239.

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Transactions and Proceedings of the Royal Society of New Zealand, Volume 63, 1934, Page 133

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Otoliths of Fishes from the Tertiary Formations of New Zealand. Transactions and Proceedings of the Royal Society of New Zealand, Volume 63, 1934, Page 133

Otoliths of Fishes from the Tertiary Formations of New Zealand. Transactions and Proceedings of the Royal Society of New Zealand, Volume 63, 1934, Page 133