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New Jurassic Ammonitina from New Zealand: Bathonian-Callovian Eurycephalitinae

Article in New Zealand Journal of Geology and Geophysics · December 2002 DOI: 10.1080/00288306.2002.9514988 · Source: OAI

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All in-text references underlined in blue are linked to publications on ResearchGate, Available from: Neville Hudson letting you access and read them immediately. Retrieved on: 05 October 2016 WNewestermann Zealand et Journal al.—Eurycephalitinae of Geology & Geophysics, from New Zealand2002, Vol. 45: 499–525 499 0028–8306/02/4504–0499 $7.00/0 © The Royal Society of New Zealand 2002

New Jurassic Ammonitina from New Zealand: Bathonian–Callovian Eurycephalitinae

GERD E. G. WESTERMANN Zealand occurrence is much earlier than in Indonesia (Middle School of Geography & Geology Oxfordian). Furthermore, first occurrence is diachronous McMaster University according to ammonite biostratigraphy even in Auckland Hamilton, Ontario L8S 4M1, Canada Province, that is, between Faunas 3 and 4 in Awakino valley and in Fauna 2 at Kawhia Harbour. Regional and local facies NEVILLE HUDSON control is indicated. On the other hand, Fauna 2 includes R. stehni Damborenea known from the Early/earliest JACK GRANT-MACKIE Callovian of the Andes. Department of Geology University of Auckland Keywords Ammonitina; Sphaeroceratidae; Jurassic; Private Bag 92019 Eurycephalitinae; new taxa; Bathonian; Callovian; Auckland, New Zealand Oxfordian; Kimmeridgian; New Zealand; Kawhia Harbour; Awakino valley; Andean Bioprovince; chronostratigraphy

Abstract Several new ammonite assemblages are described from the uppermost Temaikan and basal Heterian INTRODUCTION regional stages of New Zealand, Auckland Province, consisting mainly of the east Pacific Eurycephalitinae A general introduction to the Jurassic geology of New (Family Sphaeroceratidae). Four successive faunas are Zealand was given by Westermann et al. (2000), which distinguished in the upper Awakino valley: all include the treated the Bajocian Ammonitina. This paper is concerned Andean genus Araucanites, here occurring with both sex- with the next-higher ammonite faunas that are known only morphs. Fauna 1 includes Araucanites awakino n. sp., from Auckland Province in the North Island. Iniskinites cf. crassus Riccardi & Westermann, and While this work on the Middle Jurassic Ammonitina was Xenocephalites grantmackiei Westermann & Hudson m/m in progress for the last several years, G. R. Stevens has ¥ Lilloettia aff. steinmanni (Spath) f/M—latest Bathonian; thoroughly revised the Upper Jurassic Ammonoidea of New Fauna 2 includes Araucanites postawakino n. sp. and Zealand (Stevens 1997). Due to the persistent uncertainty Xenocephalites cf. stipanicici Riccardi et al.—Early/earliest in the correlation of New Zealand regional stages and Callovian; Fauna 3 with Araucanites ponganui n. sp., uncorrected taxonomic errors of the past, however, Stevens’ Iniskinites gr. cepoides (Whiteaves), and Choffatia gr. furcula and our works include some of the same taxa from the Oraka (Neumayr)—Early Callovian; Fauna 4 with Araucanites Sandstone of the Kawhia Harbour section. The genera spellmani n. sp.—? Middle Callovian. Epicephalites and Subneumayria, poorly known The Oraka Sandstone at Kawhia Harbour is now placed Kimmeridgian perisphinctaceans from Mexico, were entirely in the uppermost Bathonian and Lower Callovian identified from isolated fragments almost half a century ago (i.e., with Faunas 1 and 2). Ammonites previously identified by W. J. Arkell (1956; in Fleming & Kear 1960), and these with Kimmeridgian taxa—that is, “Epimayaites”, identifications and age have been retained by Stevens (1997). “Epicephalites”, and “Subneumayria”—are now classified By contrast, we place these poorly preserved New Zealand as microconchs of Araucanites and with macroconchs of forms, including E. marwicki Stevens, in the genera Lilloettia and Iniskinites. The superjacent Ohineruru Araucanites and Lilloettia of the east Pacific sphaeroceratid Formation contains a typically Indo-Pacific fauna of Late subfamily Eurycephalitinae. Furthermore, the Oxfordian Oxfordian to Early Kimmeridgian age, based on Sulaites genus Epimayaites formerly identified from the Oraka heteriensis (Stevens) [ex Idoceras], a close relative of Sandstone (Hudson et al. 1987; Sukamto & Westermann S. gerthi Oloriz & Westermann from New Guinea, and, 1992; Westermann 1994), characteristic for the Himalayan above, Paraboliceras macnaughti (Stevens) [ex Kossmatia]. faunal province, and recognised as a eurycephalitine The interval Upper Callovian to Middle Oxfordian microconch by Westermann (1996a, b), is now placed in cannot be documented by ammonites in New Zealand, Araucanites awakino n. sp. New Zealand Eurycephalitinae suggesting a hiatus between Oraka and Ohineruru can be compared to latest Bathonian to basal Oxfordian taxa Formations, marked by the Captain King’s Shellbed. of the Andes but probably ranged no higher than the Middle Other useful index fossils are forms of the bivalve Callovian. The only ammonite indicating Oxfordian is the Retroceramus. The first occurrence of R. galoi (Boehm) Late Oxfordian to Early Kimmeridgian genus Sulaites Oloriz defines the base of the Heterian regional stage, but its New & Westermann (1998) formerly known from the Himalayan Province. S. heteriensis (Stevens) resembles the (?)Early Kimmeridgian S. gerthi Oloriz & West. from New Guinea. Significantly, this form was formerly also compared with G02002; published 6 December 2002 Kimmeridgian Idoceras of Mexico (Arkell 1956; and in Received 7 January 2002; accepted 9 September 2002 Fleming & Kear 1960; Stevens 1997). 500 New Zealand Journal of Geology and Geophysics, 2002, Vol. 45

The late Middle Jurassic ammonites of the sphaeroceratid 2. Totara peninsula, Kawhia Harbour (Fig. 1A, E) Subfamily Eurycephalitinae Thierry have been thoroughly The sequence exposed on the east coast of the Totara revised, taxonomically and stratigraphically, by Riccardi & peninsula has long been known for its rich invertebrate Westermann (1991). The recent finds and/or identifications faunas (Trechmann 1923) including ammonites. Fleming & of eurycephalitines in New Zealand (Westermann & Hudson Kear (1960) erected three formations for marine strata 1991; Westermann 1996a, b) have greatly extended our immediately above their Wharetanu Measures: Oraka knowledge of their biogeographic global distribution, from Sandstone, Captain King’s Shellbed, and Ohineruru the eastern Pacific into the southwestern Pacific. Explor- Formation. The Oraka Sandstone is 80 m thick in this section ations in the Himalaya of Nepal by Cariou et al. (1994) and and has yielded a rich fauna of eurycephalitine ammonites. the Andes of northern Chile by Hillebrandt & Gröschke Francis (1977) recorded a single “Epicephalites cf. (1995) have significantly enhanced our knowledge of epigonus” from the Captain King’s Shellbed (R15/f8027). eurycephalitine phylogeny and chronology (Westermann Eurycephalitine ammonites are unknown from the overlying 1996c). In particular, the time-correlation of the Xeno- Ohineruru Formation. Much of the pre-1960 material from cephalites grantmackiei-Lilloettia assemblage from the this section is poorly localised and, although much of it can Upper Temaikan Stage of Auckland Province (here called be placed in the correct formation based on lithology and/or Fauna 1), can now more firmly be dated as latest Bathonian detailed locality information, the exact horizon within these Discus Chronozone. The Araucanites faunas newly units is unknown. All of the pre-1981 collections from the described here form successive associations above. type section of the Oraka Sandstone were assigned to a single This New Zealand ammonoid fauna, however, is fossil locality (R15/f8550). The collections made from this extremely low in diversity. Associated with the Eury- section since 1981, with finer stratigraphic control, are as cephalitinae are chronostratigraphically insignificant follows: phylloceratids, sometimes dominant, and some lytoceratids, as well as rare perisphinctids. The benthic invertebrate fauna, R15/f47 0–0.15 R15/f215 25.45–26.6 on the other hand, is moderately diverse, but only the R15/f52* 0.15–8.0 R15/f216* 26.6 –28.05 retroceramid bivalves so far have any value for inter-regional R15/f211 8–14 R15/f217* 28.05–30.15 correlation (Damborenea 1990, 1993; Damborenea et al. R15/f51 9 R15/f218 30.15–33.25 1992). R15/f212 14–20 R15/f219* 33.25–37.25 R15/f213 20–23 R15/f268 60 R15/f214 23.0–25.45 R15/f350 3–5 m below top FOSSIL LOCALITIES, STRATIGRAPHY, (Heights above the base of the Oraka Sandstone in metres; MACROFAUNA *locality yielding ammonites cited in text.) All ammonites described in this paper were collected from To date, in this section the lowest known occurrence sandstones and mudstones of the Rengarenga and Kirikiri of Retroceramus galoi, and therefore the base of the Groups (Fleming & Kear 1960) outcropping in the Kawhia Heterian Stage, is at R15/f213 (Helby et al. 1988). Regional Syncline (Suggate in Suggate et al. 1978) of However, in January 2000, two poorly preserved speci- Southwest Auckland Province, North Island, New Zealand mens identified only as R. ex gr. galoi were collected from (Fig. 1, 2). The strata concerned range in age from Late the upper half of R15/f52. These suggest that the base of Temaikan to ?Middle Heterian. Eurycephalitine ammonites the stage may be significantly lower than currently are not known to occur in contemporaneous sedimentary accepted, perhaps almost at the base of the Oraka sequences exposed in the Southland Regional Syncline Sandstone. (Suggate in Suggate et al. 1978 ) of southern South Island, New Zealand, and Baie de Saint Vincent Group (Campbell 3. Whakapirau Road, near Taharoa, south Kawhia (Fig. 1A) et al. 1985) of western New Caledonia. A similar lithologic sequence is exposed along Whakapirau In Southwest Auckland, strata in four areas (Fig. 1) have Road, c. 300–400 m by road northeast of its junction with yielded the ammonites described herein. the Taharoa–Te Anga Road. The Oraka Sandstone in this section is 105 m thick, with all but the basal 20–30 m 1. Opuatia valley, Port Waikato (Fig. 1A) exposed in a high cliff from which in situ collection is Rare Araucanites are known from a small farm quarry (R13/ impossible. Material clearly derived from this cliff is f125* and f7012: R13/72022193†) beside Ponganui Road scattered at its base and contains a rich fauna (R16/f6048). and near Opuatia Stream. Here quarrying operations have Eurycephalitine ammonites have been collected from the exposed 5–10 m of grey mudstone of the Pakau Formation, Oraka Sandstone in this section but they are less numerous 160–170 m above its base. This locality is 130–140 m below and diverse than at the Totara peninsula. The Captain King’s the incoming of Retroceramus galoi (R13/f239) and 180– Shellbed and basal part of the Ohineruru Formation are not 200 m below the rich fauna of Moewaka Quarry. exposed along the road, but the presence of the former in the hillside above R16/f6048 has been documented by Francis (1977). Two well-localised ammonite-bearing fossil localities are known from the lower part of the Oraka Sandstone in this section, R16/f86 and f403, respectively 10 and 0–18 m above its base. *Fossil locality number f125 on 1:50 000 NZMS 260 map series, Sheet R13, recorded within the national archival Fossil Record Masterfile of the Geological Society of New Zealand. 4. Northern (“upper”) Awakino valley (Fig. 1A, C) †Grid reference on Sheet R13 of the metric NZMS 260 1:50 000 Eurycephalitine ammonites are known from four sections topographic map series. in the upper Awakino valley: Westermann et al.—Eurycephalitinae from New Zealand 501

Fig. 1 Locality maps showing collection sites of faunas studied herein. A, Southwest Auckland showing location of maps B, C, and four fossil localites. B, Totara peninsula, southern shore of Kawhia Harbour. C, Upper Awakino valley showing Whakapatiki Stream and Spellmans Cliff sections. Fossil localites bear their Fossil Record File “f” numbers, which should be prefixed with the relevant map sheet number as indicated. 502 New Zealand Journal of Geology and Geophysics, 2002, Vol. 45

Fig. 2 Stratigraphic columns showing lithologic sequences and fossil localities relevant to this study for parts of Southwest Auckland covered by Fig. 1 and Table 1. (Fossil locality numbers on each column are prefixed by the NZMS 260 map sheet indicated in the heading for each.)

(1) Waterfall Creek section. An unnamed, westward flowing rich, dominantly molluscan, fauna including abundant tributary which enters the Awakino River at R17/68350315, eurycephalitine ammonites. Part of this ammonite fauna has and is referred to informally as Waterfall Creek (Francis been documented previously (Westermann & Hudson 1991). 1977). A single locality (R17/f617) has yielded Araucanites Poorly preserved eurycephalitine ammonites are also in situ 30 m below the top of the Oraka Sandstone. Two common at R17/f375, 145–170 m higher in the same other specimens, most likely derived from the Oraka formation. A further 10–25 m higher in the formation (R17/ Sandstone, are recorded from float (R17/f613, f615) in this f670–f673), four eurycephalitine ammonites (Araucanites, creek. Lilloettia, and Xenocephalites) were collected in January (2) Whakapatiki Stream section. Exposures in and beside 1995 from a 30 m high cliff (R17/f376) which had been Whakapatiki Stream and on the eastern bank of the Awakino searched on nine previous occasions between 1982 and 1991 River 100–200 m upstream from the mouth of Whakapatiki without yielding a single eurycephalitine ammonite. Stream. The stratigraphically lowest locality (R17/f566), a (3) “Spellmans Cliff”—an informal name for prominent 2 m thick pebbly medium sandstone, 100 m above the base cliffs from R17/68039876 to 68059888, on the eastern side of an unnamed Kirikiri Group formation, has yielded a very of Awakino River (Fig. 3), opposite the house of John and Westermann et al.—Eurycephalitinae from New Zealand 503

Fig. 3 View of the Spellman Cliff section on the northern bank of the upper Awakino River. Strike is parallel to the slope, and dip c. 25∞N (i.e., away from the slope), so that the total thickness of the exposed section is over 100 m. River flows left to right and the better exposed lower cliff is on the right above the sheep; the upper cliff lies in the upper left, with a bushed area between.

Mary Spellman. Here, 100–105 m of medium-bedded to Late Temaikan to Middle Heterian, as defined by bio- massive (0.2–3.0 m thick), medium, and fine to very fine stratigraphic schemes erected by MacFarlan (1975) and sandstones and mudstones included in the Oraka Sandstone Hudson (1983) and modified by Westermann et al. (2000). by Hudson (1999) are exposed. The cliff consists of two Hudson (1999) utilised the first appearance of Retroceramus sets of exposures separated by a small waterfall. Exposure marwicki (Speden) to mark the base of the Late Temaikan. on the downstream portion of the cliff extends from near The base of the succeeding Heterian Stage is marked by the river level to c. 50 m above, and exposure in the upstream first appearance of Retroceramus galoi (Boehm), as portion extends from 30 to 100 m above river level. indicated by Marwick (1953) and defined by Fleming & Kear Collections from Spellmans Cliff made before 1990 were (1960). MacFarlan (1975) erected a three-fold subdivision not finely localised and only separated as two localities— scheme for the Heterian, with the base of the Middle and downstream portion or lower cliff (R17/f302), and upstream Late Heterian marked by the first appearances of Malayo- portion or upper cliff (R17/f619). Subsequent collections maorica malayomaorica (Krumbeck) and Retroceramus aff. have been made from narrower stratigraphic intervals as subhaasti, respectively. Hudson (1999) erected a four-fold follows: zonal scheme for the Late Temaikan, based on the first R17/f643* c. 65 R17/f648* 12–15 appearances of successive species of Retroceramus, as follows: zone 1 Retroceramus marwicki; zone 2 Retro- R17/f644* c. 75 R17/f649 25 ceramus n. sp. A.; zone 3 Retroceramus stehni Damborenea; R17/f645* c. 80 R17/f650 30–35 zone 4 Retroceramus spp. C or D (of Hudson 1999). R17/f646 c. 95–98 R17/f651 c. 40 The ages of the eurycephalitine ammonite localities R17/f647* c. 100–105 R17/f652* 50–55 based on the Late Temaikan to Middle Heterian bio- (Heights above the base of cliff and Oraka Sandstone in stratigraphic scheme outlined above are as follows: (1) small metres; *locality yielding ammonites cited in text; R17/ farm quarry, Opuatia valley—Late Temaikan zone 4; (2) f643–647 are from the upper cliff, R17/f648–652 are from Totara peninsula—Late Temaikan R15/f52, Early Heterian the lower cliff.) R15/f216, f217, f219, Middle Heterian R15/f8027; (3) (4) A single locality (R17/f620) near Gribbon Road and Whakapirau Road—Late Temaikan R16/f86, f403, Early Awakino River, 1.5 km northeast of the mouth of Palmer Heterian R16/f6048; (4a) Waterfall Creek—Early Heterian; Creek. Here, 5–10 m of weathered fine sandstone to sandy (4b) Whakapatiki Stream—Late Temaikan zone 2 R17/f566, mudstone, c. 250 m above the base of an unnamed formation zone 3 R17/f375, f670–673; (4c) Spellmans Cliff—Late of the Kirikiri Group, has yielded a single Araucanites. Temaikan zone 4 R17/f648, f652, Early Heterian R17/f643– The diverse invertebrate macrofauna of these ammonite- 645, f647; (4d) Gribbon Road—Late Temaikan zone ?1. bearing localities includes brachiopods, bivalves, gastropods, As discussed by Westermann et al. (2000), the lowest and belemnites (Table 1). zone in the Late Temaikan, characterised by Retroceramus marwicki, correlates with the early Late Bajocian based on ammonite correlation of its stratigraphic range in both New BIOSTRATIGRAPHY AND Zealand and Argentina. Retroceramus stehni is recorded CHRONOSTRATIGRAPHIC CORRELATIONS from the Discus to Gracilis Chrons of Argentina (Damborenea 1990). Retroceramus galoi and Malayo- Bivalve correlations maorica malayomaorica were described from the Oxfordian The ammonites described and discussed in this paper come of Indonesia (Boehm 1907; Krumbeck 1934). However, in from a narrow stratigraphic interval which ranges in age from New Zealand, Fleming & Kear (1960) dated the lowest 504 New Zealand Journal of Geology and Geophysics, 2002, Vol. 45

Table 1 Macrofauna of Late Temaikan to Early Heterian ammonite-bearing localities of Bathonian–Callovian age. Fossil localities are grouped by section and within each group in stratigraphic order, younging to the right. Westermann et al.—Eurycephalitinae from New Zealand 505

Fig. 4 Summary of ammonite and Retroceramus ranges in Late Temaikan and Early Heterian strata of Kawhia and Awakino areas, assuming that the zone of R. stehni at Awakino was con- temporary with part of the R. galoi range at Kawhia. The standard chronostratigraphy at the left is drawn on the assumption that the sedimentary record is tolerably complete, without major non- sequences on the scale of sub- stages. Lines between columns for the international and local stages indicate the degree of precision of correlations, with narrowest single line representing the most certain correlation.

occurrences of this bivalve in the Heterian stratotype as (cf. Arkell 1956), where the marine beds begin in the Kimmeridgian based on Arkell’s unpublished ammonite Temaikan regional stage (see also Fig. 4). identifications. Stevens (1997) concluded that Retroceramus 1. Oraka Sandstone. The earlier literature on the fauna of galoi in New Zealand ranges in age from Oxfordian to Early the Oraka was recently summarised by Meesook & Grant- Kimmeridgian. The disparity is largely due to Arkell’s and Mackie (1995), Stevens (1997), and Hudson (1999). subsequently Stevens’ misidentification of ammonites from According to Stevens, the relatively thin Oraka Sandstone the lower part of the Heterian stratotype. These mis- ranges in age from Late Bathonian/Early Callovian to Early identifications and consequent chronostratigraphic Kimmeridgian. Our taxonomic revision of the ammonite misinterpretation are addressed below. fauna, however, restricts the age to latest Bathonian to Early Ammonite faunas and correlations Callovian. The basal, yellowish-brown weathering sandstone/ Kawhia Harbour siltstone has yielded long-ranging infaunal bivalves, without The New Zealand Jurassic is well exposed in the thick Retroceramus galoi, and is therefore placed in the uppermost classical sequence along the south side of Kawhia Harbour Temaikan regional stage. Rare and usually poorly preserved 506 New Zealand Journal of Geology and Geophysics, 2002, Vol. 45 sphaeroceratid ammonites include Iniskinites cf. crassus Ohineruru Formation (Francis 1977), together with the Ricc. & West., Xenocephalites cf. grantmackiei West. & occurrence of Retroceramus galoi, would indicate an Hudson, and ?Eurycephalites cf. extremus (Tornq.). The two Oxfordian age for most of the Oraka. Furthermore, Stevens former clearly indicate the Late or latest Bathonian, I. crassus (1997) listed a single specimen (C392), the holotype, of the being known from the Steinmanni Zone of Argentina and supposedly Kimmeridgian “Idoceras” heteriense Stevens, X. grantmackiei from the coeval Discus Chron of the upper as coming “from the uppermost Oraka Sandstone” (p. 46). Awakino valley (see below); E. extremus is known from the This species is now placed in Sulaites Oloriz & Westermann Early Callovian Bodenbenderi Zone of the Andes (Riccardi (1998) and relatively well dated as Late/latest Oxfordian to & Westermann 1991), but even generic affinity is doubtful. Early Kimmeridgian. The holotype, however, came from an Latest Bathonian is strongly indicated (Fauna 1; see below). old collection made in 1945 by E. S. Richardson, and From the concretionary mudstones in the middle part of stratigraphic placing depends on similarly old unpublished the formation, Meesook & Grant-Mackie (1995) list the notes by R. A. S. Browne (see Stevens 1997, p. 46). bivalve Retroceramus galoi (Boehm), a species ranging in Extensive new collecting in these beds has yielded no Indonesia from the Middle Oxfordian to the Early additional specimens, and its lithology is the same as that Kimmeridgian (Sukamto & Westermann 1992; Damborenea of the calcareous concretions with S. heteriensis in the lower 1993) and defining the base of the Heterian regional stage. Ohineruru Formation from where the bulk of Sulaites are Damborenea & Manceñido (1992), however, illustrate from known to have come. Significantly, the other ammonite locality “R17/f219” (erroneously for R15/f219) the Andean, recorded by Stevens (1997, p. 42) as coming from the Late Bathonian R. cf. patagonicus (Phil.) (which occurs in uppermost Oraka Sandstone is the holotype of the New Zealand above R. stehni, not below as in Argentina, supposedly also Kimmeridgian “Epicephalites” marwicki and may therefore be Early Callovian), and Helby et al. Stevens, which is now placed in the typically Late Bathonian (1988) identified probable Callovian dinoflagellates. The to Early Callovian genus Lilloettia. poorly preserved, scarce ammonitine fauna here described 2. Captain King’s Shellbed. This c. 1 m thick highly is that of the lower part of the Awakino valley section, that fossiliferous formation, separating the Oraka Sandstone and is, Araucanites awakino n. sp. m/m and f/M and Lilloettia superposed Ohineruru Formations, can be dated by the rare gr. steinmanni (Spath) f/M. The holotype and only known ammonites as Late/latest Oxfordian to Early Kimmeridgian specimen of Lilloettia [“Epicephalites”] marwicki (Stevens) (see below), based on the first, rare occurrence of Sulaites came from either this or the upper part of the formation. heteriensis (Stevens 1997, p. 46). According to Stevens This “Andean” ammonite fauna suggests an Early Callovian (1997), this species possibly appears already in the topmost age, supporting the dating based on retroceramid bivalves Oraka Sandstone, as discussed above. The main occurrence and dinoflagellates. It predates the ?Late Callovian– of S. heteriensis is in the lower Ohineruru Formation (see Oxfordian date based on the identification of Retroceramus below). Francis (1977) also recorded “Epicephalites”, that galoi by Meesook & Grant-Mackie (1995) and Hudson is, Araucanites, besides rare phylloceratids. These rare and (1999), a species appearing above the awakino range zone usually fragmented ammonites, however, may have been (Faunas 1–2a) in the Awakino valley; and especially the reworked from earlier strata. Kimmeridgian date given by Stevens (1997), based on misidentification of ammonites (i.e., “Epicephalites”, 3. Ohineruru Formation. The few age-diagnostic “Subneumayria”). ammonites (Ammonitina) of the lower part of the Ohineruru Stevens’ (1997) dating followed Fleming & Kear’s belong mainly or entirely to Sulaites heteriensis (Stevens) classical study (1960), which included ammonite identi- of Late/latest Oxfordian to Early Kimmeridgian age. This fications by W. J. Arkell. Arkell (1956, p. 455) identified a species was originally named “Idoceras cf. humboldti few fragments sent to him with “Epicephalites cf. epigonus (Burckhardt)” [also as I. n. sp., aff. humboldti] (Arkell 1956 (Burckhardt)” and “Subneumayria sp.”, both originally and in Fleming & Kear 1960; Stevens 1997), a Mexican described from the Lower Kimmeridgian of Mexico form dated as late Early Kimmeridgian. Francis & (Burckhardt 1906). Whereas the holotype of “Epicephalites” Westermann (1994) compared these forms with marwicki Stevens (1997) belongs to Lilloettia, all or most “Perisphinctes” sularus-moluccanus, Boehm spp. (1907), other specimens allotted to that species (more or less crushed and “Pseudoparaboliceras aramaraii” Gerth (1965), that is, fragments) as well as “Subneumayria sp. indet. cf. respectively Sulaites sularus and gerthi of Oloriz & S. ordonezi (Burckhardt)” of Stevens (1997, fig. 30 and pl. Westermann (1998). S. sularus (Boehm) is known mainly 15, fig. 2, 3; fig. 30 shows a fastigate venter, which must be from the Upper Oxfordian (includes Divisum Standard Zone) due to crushing), belong to Araucanites (Westermann 1996a, of Indonesia. The species group of “P. aramaraii” 2001; Oloriz & Westermann 1988; see Systematics). Most (=S. gerthi) appears to be slightly younger, and was fragments, however, are indeed unidentifiable even at family tentatively dated from ex situ associations of Papua New level. Hudson et al. (1987) and Westermann (1994) identified Guinea as latest Oxfordian to Early/earliest Kimmeridgian a single, minute “Epimayaites” from the Oraka Sandstone, (Francis & Westermann 1994) and recorded from New a genus or subgenus (of Mayaites) known only from the Zealand (Westermann 1996a). Sulaites heteriensis Oxfordian of northeastern Gondwanaland. This was still (Stevens) closely resembles S. gerthi, which may even accepted as evidence for Oxfordian by Stevens (1997), be conspecific (and a junior synonym). Both in Papua although Westermann (1996a, b) had transferred it to the New Guinea and New Zealand (Meesook & Grant-Mackie Eurycephalitinae. The New Zealand form is much more 1995), the S. heteriensis-gerthi group is associated with the involute than juvenile mayaitines of similar diameter and is Middle Oxfordian to Kimmeridgian (Damborenea & now identified with Araucanites awakino microconchs. Manceñido 1992) or Kimmeridgian to ?Early Tithonian On the other hand, the apparent absence of stratigraphic (Stevens 1997) bivalve Malayomaorica malayomaorica breaks below the latest Oxfordian to Early Kimmeridgian (Krumbeck). Westermann et al.—Eurycephalitinae from New Zealand 507

Stevens (1997) records both Sulaites heteriensis rare A. postawakino n. sp., as well as rare representatives of (Stevens) and Paraboliceras macnaughti (Stevens) (ex the typically Early Callovian Lilloettia cf./aff. lilloetensis Kossmatia, see Systematics), as ranging throughout the Crickmay f/M and Xenocephalites cf. stipanicici Riccardi Ohineruru Formation and into the superjacent Waikutakuta et al. m/m. The bivalve Retroceramus stehni Damborenea, Siltstone, with dominance of S. heteriensis below (ranging typical for the Early Callovian of Argentina, has also been downward into the Captain King’s Shellbed and, possibly, identified from these localities (Damborenea & Manceñido the uppermost Oraka Sandstone) and of P. macnaughti above 1992). (and ranging somewhat higher within the Waikutakuta). From the incomplete set of recorded specimens available to Fauna 3. About 1.5 km downstream, beside the Awakino us, we can confirm the long range of Sulaites, from the lower River, but only 10–20 m stratigraphically higher, begin the Ohineruru into the Waikutakuta Siltstone (where S. n. sp. A high cliffs opposite the Spellman farmhouse (“Spellman differs from S. heteriensis in the more involute, compressed Cliff”), which expose c. 100 m section. A small Eury- whorls). The few specimens of P. macnaughti we were able cephalitinae assemblage (R17/f302, f648) with Iniskinites to examine were mostly from the upper Ohineruru and gr. cepoides (Whiteaves) f/M and Araucanites ponganui Waikutakuta, although Stevens (1997) recorded rare n. sp. m/m, as well as the rare perisphinctid Choffatia specimens (2) from the lower Ohineruru, confirming the (Homoeoplanulites) gr. furcula (Neumayr) m/m, and a few overlap of Sulaites with Paraboliceras. The occurrence of phylloceratids and lytoceratids, occurs 12–15 m above the the closely interrelated Kossmatia-Paraboliceras clade in base. the Kimmeridgian of New Zealand has been supposed previously (Arkell 1956 and in Fleming & Kear 1960; Fauna 4. Beginning 40 m higher in the Spellman Cliff Francis & Westermann 1994). Association with Sulaites is section, ranging from 50 m above the base through the good evidence that Paraboliceras appeared already in the remaining 50–55 m of the section of fine to very fine Early Kimmeridgian. This contrasts with the widely held sandstones and mudstones, occurs Araucanites spellmani opinion (e.g., “Treatise of Invertebrate Paleontology”) that n. sp. Both macroconchs and microconchs are relatively Paraboliceras and Kossmatia were restricted to the (higher) abundant at 65–105 m (R17/f619, f643–645, f647). The Tithonian. A Kimmeridgian appearance has recently been bivalve Retroceramus galoi (Boehm) is also found for the documented also for the Himalayas (Enay & Cariou 1996). first time at these levels, marking the base of the Heterian Perhaps Kossmatia and Paraboliceras should be placed into regional stage. the same genus, as subgenera, especially since some forms, Approximately 210 m of poorly exposed section, without for example, “Kossmatia” macnaughti, are morphologically known ammonite fauna, separates the top of the known range intermediate. This New Zealand species is here placed in of Fauna 4 at the top of Spellman Cliff from Captain King’s Paraboliceras because of the ventrally barely projected and Shellbed, which is located near the top of the ridge. clearly interrupted ribs. Stevens’ (1997) base for classi- fication are the highly variable and diagnostically unreliable Does Captain King’s Shellbed represent a parabolae. stratigraphic hiatus? The stratigraphic evidence from Kawhia Harbour On the Totara peninsula, Kawhia Harbour, stratotype of indicates that the Paraboliceras-Kossmatia clade branched Captain King’s Shellbed, the 1–2 m bed separates the off early Sulaites in the Early Kimmeridgian (or possibly subjacent Oraka Sandstone and the superjacent Ohineruru even the Late Oxfordian). This modifies the early hypothesis Formation. The relatively fossiliferous middle Oraka of a linear evolution from Sulaites in about the middle part Sandstone, up to within c. 15 m of the top, bears the Early of the Kimmeridgian, supposed by Westermann (in Francis Callovian eurycephalitine Fauna 2 of the Whakapatiki/ & Westermann 1994; Westermann 1996a). Awakino section. Fauna 1 occurs in the basal sandstones of the lower Oraka, but Faunas 3 and 4 are unknown from the Upper Awakino valley Kawhia Harbour area. The Ohineruru Formation is mainly Four Eurycephalitinae associations, here simply called of Early Kimmeridgian age, based on the Sulaites faunas, occur at separate levels (oldest first). heteriensis-Paraboliceras macnaughti association, although its basal beds and Captain King’s Shellbed, with Fauna 1. This assemblage occurs at a small tributary, Te S. heteriensis documented only, may be slightly older (i.e., Whakapatiki Stream, in a 2 m thick, pebbly sandstone (R17/ Late/latest Oxfordian). Hence, the shellbed only partly fills f566). Xenocephalites grantmackiei West. & Hudson m/m a Mid-Callovian to Mid-Oxfordian hiatus of 5–7 m.y. and Lilloettia gr. steinmanni (Spath) f/M [ex L. aff. boesei between deposition of the Oraka and Ohineruru Formations. (Burckh.)] have been described from here and dated as latest Such an extensive hiatus, however, contradicts the Bathonian (Westermann & Hudson 1991). These are lithostratigraphic record (Francis 1977). Captain King’s associated with abundant dimorphic Araucanites awakino shows no basal unconformity, erosional surface or n. sp. Unfortunately, many of the specimens are juvenile hardground, so that sedimentation of the sequence appears and/or incomplete and distorted. to have been uninterrupted, although deposition of the thin glauconite-rich shellbed must have been extremely slow. Fauna 2. This assemblage occurs c. 160 m stratigraphically Francis concluded that the slow deposition of the shallow- above Fauna 1, through a 25 m section at the confluence of water shellbed (20–40 m, from bivalves) was due to a bar Te Whakapatiki Stream and Awakino River (“Whakapatiki/ that prevented sediment reaching the area. Ballance & Awakino section”; R17/f375, f376, f670–673). Euryce- Campbell (1993), on the other hand, have argued that the phalitinae again dominate the small assemblage, with long-term, starved deposition of the Captain King’s was due microconchs of abundant Araucanites awakino n. sp. and to a sea-level highstand that drowned the sediment sources. 508 New Zealand Journal of Geology and Geophysics, 2002, Vol. 45

This, of course, implies increased depth, contrasting with Province, dated also as Late Bathonian (Hillebrandt et al. Francis’ conclusion of shallowing. The presence or absence 1992). Iniskinites cf. crassus f/M and Xenocephalites cf. of a time break associated with the base of the Captain King’s grantmackiei m/m have now been identified from the basal Shellbed remains unproven. Oraka Sandstone of Kawhia (R15/f8550, the specific interval Ammonite and retroceramid evidence concerning the from which it came is most likely to be R15/f52) and correlation of the Oraka Sandstone in the Awakino and Whakapirau Road (R16/f403), indicating the Late Bathonian Kawhia areas is conflicting. The upper Awakino valley here. Iniskinites cf. crassus has not been found with any section below the Captain King’s Shellbed is over 320 m other member of Fauna 1 but is tentatively included here thick, including roughly 105 m with Faunas 3 and 4 that because of its stratigraphic position with respect to A. cf. probably postdate the type Oraka Sandstone (GEGW). On awakino and the age overseas of I. crassus. the other hand, the ammonite evidence conflicts with that from the retroceramids which suggest that ammonite Faunas Fauna 2. [R17/f375, f376, f670–673] 3 (latest Temaikan—Retroceramus zone 4) and 4 (earliest Araucanites awakino n. sp. m/m and f/M, A. postawakino Heterian) are probably contemporaneous with the lowest 20– n. sp. m/m, L. cf./aff. lilloetensis f/M, Xenocephalites 30 m of type Oraka. The highest ammonite fauna (# 4) is cf. stipanicici m/m, Retroceramus stehni Damborenea. separated from the Captain King’s Shellbed by c. 220 m of Late Temaikan and Early/earliest Callovian. poorly exposed strata that have yielded no datable The listed Lilloettia and Xenocephalites “monosexual ammonites. Although the stratigraphic evidence clearly species” document the Lower Callovian. This is supported indicates that this thick part of the section is more complete by the Andean bivalve R. stehni identified by Damborenea than at Kawhia, the post-Araucanites spellmani beds cannot (in Damborenea & Manceñido 1992) and the moderate be dated. Thus, there is no way of knowing whether the mid- stratigraphic distance above association 1. Fauna 2 is also Callovian to mid-Oxfordian interval missing at Kawhia is found in the Lower Heterian portion of the type Oraka present here or not, until additional faunas have been found. Sandstone, where relatively common Araucanites awakino occur with Retroceramus galoi and very rare Lilloettia Port Waikato area: Ponganui Farm Quarry marwicki. The small farm quarry beside Ponganui Road (Ponganui Farm Quarry) and Opuatia Stream has yielded a mixed Fauna 2a. [R13/f125, f7012] eurycephalitine assemblage of Araucanites ponganui n. sp. Araucanites ponganui n. sp. m/m, A. awakino m/m and microconchs and similar numbers of dimorphic A. awakino f/M, Retroceramus sp. D. Late Temaikan and Early n. sp., together with one large Lilloettia (Imlayoceras) Callovian macroconch. This fauna includes elements of both Faunas This association is known only from the Ponganui Farm 2 and 3 of the Awakino valley, and is treated in an Quarry and its stratigraphic position is largely inferred from intermediate position below. the ammonite fauna being intermediate between Faunas 2 and 3. This fauna lies stratigraphically between the zone of Retroceramus stehni, and thus Fauna 2, and the base of the AMMONITE FAUNAS AND Heterian, and may be contemporaneous with Fauna 3. CHRONOSTRATIGRAPHIC CORRELATIONS The upward succession of ammonitine Faunas 1–4 is given Fauna 3. [R17/f302, f648] f below; Phylloceratina and, to a lesser degree, Lytoceratina Iniskinites gr. cepoides (Whiteaves) /M, Choffatia m become relatively abundant in the upper faunas (Table 1), (Homoeoplanulites) gr. furcula (Neumayr) /m, m but are not listed because of their low chronostratigraphic Araucanites ponganui n. sp. /m, Retroceramus sp. C. utility. The list is based mostly on data from the thick Latest Temaikan and Early Callovian. sequence of the upper Awakino valley, with the less extensive The species group of I. cepoides is known from the Upper data from the thinner and less “ammonitiferous” sequence Bathonian and basal Callovian of western North America at Kawhia Harbour incorporated. The main locality numbers (see Systematics and Callomon 1984); the Choffatia furcula are in brackets. species group has a similar stratigraphic range in the Mediterranean-Caucasian (“West Tethyan”) and Indo-Pacific Fauna 1. [R17/f566, ?R15/f52, ?R16/f403] (“Indo SW Pacific”) Subrealms, including the Gracilis Zone Xenocephalites grantmackiei West. & Hudson m/m ¥ of Indonesia (Westermann & Callomon 1988; Westermann Lilloettia cf. steinmanni (Spath) f/M, Araucanites 2000). Roughly at this level occurs also rare Araucanites awakino m/m and f/M, and possibly Iniskinites cf. ponganui m/m, found also together with A. awakino in Fauna crassus Ricc. & West. Late Temaikan and latest 2a. Retroceramus galoi has not been found. An Early Bathonian. Callovian age is indicated by the Iniskinites and Choffatia This association has in part been described previously species. from Te Whakapatiki Stream by Westermann & Hudson (1991). X. grantmackiei can be relatively securely dated as Fauna 4. [R17/f613, f617, f619, f643–647, f652] f m latest Bathonian (Westermann 1996c). It occurs in the A. spellmani n. sp. /M and /m, first Retroceramus Apertus Zone of Papua New Guinea (Westermann & galoi (Boehm). Latest Temaikan to earliest Heterian and Callomon 1988), which has recently been extended to Nepal ?Early–?Middle Callovian. where its indices are associated with Mediterranean- The close morphologic affinity between the eury- Caucasian (“West Tethyan”) faunas of the (lower) Discus cephalitines found here and those from the (latest Callovian Zone (Cariou et al. 1994). On the other hand, there is close and) basal Oxfordian beds of northern Chile described by affinity with the fauna of the Steinmanni Zone of the Andean Hillebrandt & Gröschke (1995) suggests a much later age Westermann et al.—Eurycephalitinae from New Zealand 509 than the previous faunas. Yet they are in close stratigraphic SYSTEMATIC DESCRIPTION OF AMMONITINA proximity (40 m) in this thick and rather uniform basinal (prepared by GEGW) sequence, with no sign of an hiatus between. Furthermore, Catalogue numbers and repositories eurycephalitine evolution was notoriously slow (cf. Riccardi & Westermann 1991), and Araucanites microconchs are C = Cephalopod specimen number, Department of unknown from the principal mid-Oxfordian Araucanites Geology, University of Auckland. localities in west-central Argentina (cf. Stipanicic et al. CE = Cephalopod specimen number, Institute of Geological 1976). On balance, we conclude that the most probable age & Nuclear Sciences, Lower Hutt, New Zealand. is Middle Callovian. Abbreviations Abbreviations for morphologic terms are: D, diameter; Wb, BIOGEOGRAPHY whorl breadth; Wh, whorl height; U, umbilical diameter; P, number of primaries per half-whorl; S, number of The Late Bathonian Fauna 1, consisting mainly of the secondaries per half-whorl; phr., phragmocone; b.ch., body Xenocephalites and Lilloettia pair together with the earliest chamber; apert., aperture; m/m, male shell/microconch; f/ Araucanites, belongs to the Andean Province, East Pacific M, female shell/macroconch. Measurements are recorded Subrealm. The extreme range of X. grantmackiei as being taken at the aperture (apert.), at the end of the Westermann & Hudson extended into the New Guinea area phragmocone, that is, at about the level of the last septum of the southern Himalayan Province, where it has been found (end phr.), or at an unspecified level of the body chamber, as an adjunct element in an Indo-Pacific ammonite fauna with the aperture broken away (b.ch.). (Westermann 1996b, 2000). The Early–Middle Callovian Eurycephalitinae faunas of New Zealand, which were dominated by the genus Araucanites and included Iniskinites Suborder AMMONITINA Hyatt, 1889 and Lilloettia, are also typically Andean in affinity, probably Superfamily STEPHANOCERATACEAE Neumayr, 1875 even partly conspecific. Only very rare “Tethyan” elements Family SPHAEROCERATIDAE Buckman, 1920 (e.g., the perisphinctid genus Choffatia), have been found. Subfamily EURYCEPHALITINAE Thierry, 1978 Some of these Eurycephalitinae were previously mis- identified with the “Tethyan” Epicephalites and Sub- Scope, phylogeny, taxonomy, and distribution of this neumayria from the Kimmeridgian of Mexico or with the subfamily with mainly East Pacific, especially Andean Indo-Pacific Epimayaites from the Oxfordian of East Africa distribution were discussed extensively by Sandoval et al. to New Guinea. (1990), Riccardi & Westermann (1991), and Westermann No Late Callovian to Middle Oxfordian ammonite faunas (1996a–c). Dimorphism is clearly present but corres- are presently known from New Zealand. Subsequent pondences are hard to establish even at generic level, because ammonite faunas are typically those of the Himalayan of the poor knowledge of the inner whorls in these involute Province, Indo-Pacific Realm/Subrealm, beginning with the sphaerocones, despite the often large samples. The perisphinctid Sulaites (Oloriz & Westermann 1998). This macroconchiate juvenile whorls and the entire phragmocone genus, formerly called “‘Perisphinctes’ gr. sularus- of the small and strongly differentiated microconchs need moluccanus (Boehm)”, is also known from the Late to be further investigated. A notable exception is the recent Oxfordian of the Himalayas (Enay & Cariou 1996) and documentation by Parent (1997) of the specific dimorphic Indonesia, and from the Late Oxfordian to Early correspondence between the macroconch Eurycephalites Kimmeridgian of Papua New Guinea (cf. Westermann vergarensis (Burckhardt) and the microconch Xenocephalites 1996b; Oloriz & Westermann 1998). The New Zealand gottschei (Tornquist) in Argentina. However, since species S. heteriensis (Stevens) may be conspecific with X. gotschei is not the type species of Xenocephalites and the relatively late, New Guinean species S. gerthi Oloriz several genera are currently distinguished as corresponding & Westermann. Other typically Indo-Pacific ammonite macroconchs, there is not sufficient reason to draw the name species also appear in New Zealand at this time—the Xenocephalites into synonymy. However, it is possible that Kimmeridgian–Tithonian Paraboliceras-Kossmatia clade the specific features are not yet developed in the minute inner with its earliest known member, P. macnaughti (Stevens). whorls which, according to the hypothesis of sexual The inoceramid bivalve fauna has similar biogeographic dimorphism, have to be morphologically identical. Thus, problems. The morphologic range of Retroceramus galoi current taxonomy by necessity treats several “macroconch needs to be better constrained than it currently is. But, genera” as equivalents to the single “microconch genus” presently, New Zealand material so identified appears to Xenocephalites. It is also possible that the mismatch has been be conspecific with type material from Indonesia, which due to excessive “splitting” of the macroconchs at mainly is of Middle Oxfordian–Kimmeridgian age. New Zealand the generic level, even in the most recent classifications specimens, however, are apparently older, occurring first (Riccardi & Westermann 1991). For example, the macro- in the probably Early–Middle Callovian spellmani conchs Eurycephalites, Lilloettia, and Iniskinites are ammonite association (Fauna 4). Significantly, pre- distinguished by the relative persistence of ornament and/ Middle Oxfordian inoceramid species are unknown or the degree of inflation only, but are clearly connected by from Indonesia-New Guinea, but abound in the Andes intermediate morphotypes. They may not represent lineages (Damborenea & Manceñido 1992), suggesting facies control and would perhaps be classified better as subgenera; some of distribution or absence due to an incomplete record. “species” (morphospecies) may be infraspecific variants. Perhaps a contribution to a solution will come from the Furthermore, dimorphism has not been established in the geographically intermediate Antarctic Peninsula locally dominant Andean genus Stehnocephalites, retroceramid sequence. presumably because of substantial morphological overlap 510 New Zealand Journal of Geology and Geophysics, 2002, Vol. 45

(for more on the species concept in ammonites see Callomon but ranges well into the Callovian along the northeastern 1985). Pacific. Its geographic range is here extended to New Microconchs have not been found in the principal, but Zealand. sparse, Andean occurrence of Araucanites, that is, the early Middle Oxfordian Perisphinctes-Araucanites beds of Lilloettia gr. steinmanni (Spath, 1928) f/M Fig. 5A, B Mendoza (Stipanicic et al. 1976; Riccardi & Westermann aff. 1928 Macrocephalites steinmanni Spath, p. 170. 1991; Parent 1997). This may be due to collection failure, 1991 Lilloettia aff. boesei (Burckhardt). Westermann & extreme sex-ratio or post-mortem transport. In contrast, the Hudson, p. 692, fig. 3.3–3.6. extended Araucanites lineage of New Zealand has yielded a sequence of microconchs which not only can be matched, MATERIAL: 2, C1424, C1426, from Te Whakapatiki Stream, more or less confidently, with the associated macroconchs, upper Awakino valley, R17/f566, previously identified as but that are also morphologically clearly distinct from Lilloettia aff. boesei are re-identified as L. gr. steinmanni. Xenocephalites (see under Araucanites, below). In Additional material: 1 fragment, C1709, of outer phrag- consequence, no separate “microconch genus” is distin- mocone with shell and umbilical wall of body chamber, float guished in Araucanites. from middle Oraka Sandstone at R15/f8550, Totara Diagnostic morphologic features of the eurycephalitine peninsula, Kawhia Harbour. shells, at the genus- and species-group levels, are diverse, so that their taxonomy is properly based on a combination DESCRIPTION: Phragmocone at D 40–60 mm is very of characters, including the type of dimorphism (for involute (U/D 10%) with compressed subtrapezoidal whorl morphologic terminology see Westermann 1996d). The section (H/W 1.2); body chamber remained involute with essential features are: (1) adult diameters of macro- and reconstructed D c. 80 mm. microconch (f and m) and the ratio between them; (2) degree of roundness (inflation) and coiling, here from involute to REMARKS: The L. steinmanni group includes several obvolute (umbilicus closed by callus); (3) in the microconch, discoconic, densely costate, macroconchiate “species” from the degree of variocostation (mature modification in ribbing) the Late Bathonian to Early Callovian of North and South that may be gradual or rapid and on different parts of the America that are closely interrelated, difficult to distinguish body chamber; (4) in the macroconch, whether ribs (costae) without knowledge of the inner whorls, and probably partly are maintained to aperture or disappear along the mature synonymous (Riccardi & Westermann 1991, p. 5). Included body chamber, and whether they occur on inner and/or outer are L. lilloetensis and mertonyarwoodi, Crickmay spp., and flank and/or venter; (5) presence or absence of primaries L. tipperi Frebold from British Columbia and South Alaska, (ribs on inner flank) on outer septate whorl(s), of macro- and L. steinmanni from Mexico, Argentina, and Chile. and microconchs. Because of the very incomplete record of Classification of our specimens is made difficult by the faunas, all following descriptions use “morpho- compaction and incompleteness. A larger, undistorted taxonomy”, not “biotaxonomy”, sensu Callomon (1985). specimen matches L. steinmanni from the Late Bathonian Highly probable dimorphs are, nevertheless, classified as a Steinmanni Zone of west central Argentina and northern single species (not different genera or subgenera). Chile, as richly illustrated by Riccardi & Westermann (1991, pl. 9, 10). The smaller of the two specimens formerly placed Genus Lilloettia Crickmay, 1930 in L. aff. boesei by Westermann & Hudson (1991) has only slightly fewer secondaries than typical Andean L. steinmanni TYPE SPECIES: L. lilloetensis Crickmay, 1930. (S c. 35 versus c. 40). Undistorted specimens appear to differ in the adult stage, with L. boesei being more inflated and This macroconchiate taxon is closely related to the partly more coarsely costate, but they could be mere variants. coeval genus Eurycephalites Spath, which is distinguished L. lilloetensis and the closely related or conspecific mostly by the slightly less tightly coiled and more subcircular L. mertonyarwoodi and L. buckmani, Crickmay spp., from to subquadratic whorls; the outer whorls have similar the Lower Callovian of British Columbia and South Alaska smoothening of the flanks. Iniskinites Imlay has inner whorls (cf. Imlay 1953, pl. 30, fig. 1, 3, 4, 6–11; Frebold & Tipper Eurycephalites similar to those of or relatively inflated forms 1967, pl. 1, fig. 7–9), have coarser secondaries and are more Lilloettia of , but its prominent primary costae persist to the inflated. adult aperture. Most were mesodomic discocones and essentially pelagic (includes oceanic and epeiric; see Lilloettia cf./aff. lilloetensis Crickmay, 1930 f/M Westermann 1996d), as supported by their typically Fig. 6B–D extensive geographic distribution, even at the species level. For example, the discoconic and finely costate L. steinmanni cf. 1953 Lilloettia lilloetensis Crickmay, Imlay, p. 77, pl. 30, fig. group is known along the eastern Pacific margin from South 1, 2, 4, 8. cf. 1964 Lilloettia lilloetensis Crickmay, Frebold, pl. 37, fig. 2a, Alaska and British Columbia to the Southern Andes, where b (holotype refigured), 3a, b. it is characteristic of the latest Bathonian Steinmanni Zone, cf. 1967 Lilloettia lilloetensis Crickmay, Frebold & Tipper, p.11, pl. 1, fig. 1–3 (topotypes and internal part of holotype).

Fig. 5 Faunas 1, 2 and 2a. All figures natural size. Fauna 1: A, B, Lilloettia gr. steinmanni (Spath) f/M, septate fragment C1709 from middle part of Oraka Sandstone, Totara peninsula, Kawhia Harbour (R15/f8550), left side (A) and venter (B). Fauna 2: C, Lilloettia marwicki (Stevens) f/M, holotype C391, probably from upper part of Oraka Sandstone, Totara peninsula (R15/f8550). Fauna 1: D–F, Iniskinites cf. crassus Ricc. & West. f/M, somewhat deformed phragmocone with body chamber fragment C1775, loose from basal c. 18 m of Oraka Sandstone, road cutting, Whakapirau Road, South Kawhia (R16/f403); D, venter; E, left side with 1/3 whorl body chamber fragment; F, right side without body chamber. Fauna 2: G, H, Araucanites cf. awakino f/M, from middle part of Oraka Sandstone, Totara peninsula; G, body chamber ➤ Westermann et al.—Eurycephalitinae from New Zealand 511

fragment C1733 (R15/f217); H, crushed 1/2 whorl body chamber C1730 (R15/f8550). Fauna 1: I, J, Xenocephalites cf. grantmackiei West. & Hudson m/m, deformed body chamber CE2106, from basal Oraka Sandstone, loose on shore platform, Totara peninsula, Kawhia Harbour. Fauna 2a: K–N, Araucanites cf. awakino n. sp. from Ponganui Farm Quarry, Port Waikato, (R13/f7012); K, f/M, crushed with incomplete body chamber CE2769; L–N. m/m, distorted but almost complete conch CE2768 in left, ventral, and right lateral views. 512 New Zealand Journal of Geology and Geophysics, 2002, Vol. 45

Fig. 6 Faunas 1 and 2. All figures natural size. Fauna 1: A, Xenocephalites cf. stipanicici West. & Ricc. m/m, C1736 with aperture, fragmentary and partially crushed, from Whakapatiki section, 17 m above base (R17/f671). B–D, Lilloettia cf./aff. lilloetensis Crickmay f/M, fragmentary phragmocone C1716 with part of inner whorl, from same section (R17/f670), 10 m above base. E–H, Araucanites postawakino West., n. sp. m/m, holotype, complete and undistorted (C1749), from same section (R17/f673), 23–25 m above base. I–K, A. awakino West., n. sp. m/m; I, crushed phragmocone with transposed, damaged body chamber C1755, from same section (R17/f375); J, complete but deformed specimen C1757, from Oraka Sandstone of Totara peninsula, Kawhia Harbour (R15/f219); K, crushed subadult C1756, from Oraka Sandstone of Totara peninsula, Kawhia Harbour (R15/f216). Fauna 2: L, Juvenile of Araucanites m/m or Lilloettia f/M, C1771, crushed, Te Whakapatiki Stream (R17/f375). Westermann et al.—Eurycephalitinae from New Zealand 513

Fig. 7 Epicephalites epigonus Burckhardt, holotype (plaster cast) from Kimmeridgian of northern Mexico.

MATERIAL: Large fragmented specimen C1716 with inner from the Tethyan family Aulacostephanidae to the Andean whorls and aperture, from R17/f670, 10 m above river level genus Araucanites of the East Pacific subfamily Eury- in Whakapatiki/Awakino section, Fauna 2. cephalitinae (Sukamto & Westermann 1992; Westermann 1996b). DESCRIPTION: Intermediate and outer septate whorls are The holotype has 1/4 whorl of the body chamber ovate or subtrapezoidal, roughly as high as wide, and preserved, beginning at D 60 mm. Assuming a 3/4 whorl involute. At D 35–40 mm ornamentation consists of distant body chamber, the estimated complete diameter was 80–90 primaries and 3 or 4 times as many, moderately dense and mm. Its left side (Stevens 1997, pl. 4, fig. 3) is strongly somewhat prorsiradiate secondaries. Primaries fade out at abraded, with the exception of the last 1/3 preserved whorl, D 60–70 mm, whereas secondaries are retained to at least where the inner flanks are indeed smooth and the whorl 90 mm. Aperture at D 100–110 mm is moderately oblique section involute and subtriangular. The remainder of the and smooth, with laterally constricted internal mould. exposed last whorl is essentially destroyed. The other side, however, has ribs preserved on the early part of the whorl, REMARKS: The primary costae on the intermediate whorls including primaries on the inner flank that we first noted on are markedly coarser than in the L. steinmanni group Stevens’ fig. 4. We have further developed this side by described above, but whorl section and lateral smoothening removing the inner flank of the outer whorl and exposed leave no doubt of generic affinity. L. lilloetensis from the prominent, coarse primaries on the penultimate preserved late Early Callovian of North America (Callomon 1984) whorl at D c. 30–40 mm. Epicephalites is smooth at this appears to be morphologically closest to our species stage (Fig. 7). (although the holotype from British Columbia appears to The whorls at D 30–65 mm are highly involute (internal be even more compressed). Note that this classification does mould at D 65 mm, U/D 0.075) and subtriangular, not imply that our New Zealand form evolved from the North approximately as wide as high, even somewhat depressed American species. at D 40 mm. Short, prominent, coarse primaries (P 11–12) are present on the umbilical shoulder and innermost flank Lilloettia marwicki (Stevens, 1997) f/M Fig. 5C at D c. 30–50 mm. They branch into about 3 rectiradiate 1997 Epicephalites marwicki Stevens, partim, p. 41, pl. 14, fig. secondaries (S c. 30) at 1/3 whorl-height and cross straight 3–5 and fig. 29 only (holotype). over the venter. The primaries disappear at or just before the beginning of the body chamber; its first quarter-whorl MATERIAL: Besides the holotype C391 (R15/f8550), only has 14 coarse and increasingly blunt, rectiradiate secondaries poorly preserved material from Oraka Sandstone of Kawhia on the outer flank that cross straight over the venter. The Harbour area that cannot be identified with confidence (e.g., last 2 or 3 septal sutures are approximated and corroded, so C1746 from R16/f86, Whakapirau Road). that it is difficult to follow an individual suture. There is a diminishing series of 4–5 pairs of lobes and saddles, with REMARKS: The holotype of “Epicephalites” marwicki came an apparently radial saddle envelope; the 5th minute pair is from the Oraka Sandstone at Kawhia Harbour, probably from on the umbilical shoulder and slope (Stevens 1997, fig. 29, its upper part (Stevens 1997, p. 42). According to Stevens erroneously shows the envelope crescented and with (1997), the species is closely related to the Kimmeridgian retracted umbilical lobe). Epicephalites epigonus (Burckhardt) of Mexico (Fig. 7, This apparently unique specimen resembles Lilloettia holotype refigured), from which it is said to be distinguished boesei (Burckhardt) from the uppermost Bathonian in the whorl section only; both allegedly have the same Steinmanni Zone of Mexico (Sandoval et al. 1990, pl. 5, ribbing that is confined to outer flanks and venter. We have, fig. 1a, b; pl. 6, fig. 1a, b, 6; Riccardi & Westermann 1991, however, earlier transferred the New Zealand “Epicephalites text-fig. 22), from which it differs in the subtriangular (not cf. epigonus” of Stevens (1968) and Hudson et al. (1987) broadly rounded) whorl section. 514 New Zealand Journal of Geology and Geophysics, 2002, Vol. 45

? Genus Eurycephalites Spath, 1928 Frebold, all of which are here placed in the species group of TYPE SPECIAL: Macrocephalites vergarensis Burckhardt, I. cepoides (Whiteaves). 1903. All Iniskinites species known previously were dated as Bathonian to ?basal Callovian (cf. Riccardi & Westermann ?Eurycephalites gr. extremus (Tornquist, 1898) f/M 1991, text-fig. 4), but the Andean record is rather poor. Probable Iniskinites specimens were recently illustrated, MATERIAL:1 large, orthogonally deformed (vertically however, from as high as the basal Oxfordian of northern embedded!) specimen consisting of inner whorls and Chile (Hillebrandt & Gröschke 1995, pl. 5, fig. 5a, b). fragment of (?)aperture (C1717), from basal Oraka Our two deformed specimens were originally compared Sandstone, at R15/f52, Totara peninsula, Kawhia Harbour. with the Early Bajocian sphaeroceratine Chondroceras defontii (McLearn) as well as with the Late Bathonian to DESCRIPTION AND REMARKS: Specimen was c. 100–120 Early Callovian bullatimorphid Kheraiceras sp., but they mm in diameter and probably almost as wide at the aperture. differ from both mainly by the sharp costae. At least the inner whorls up to c. 60 mm diameter were involute and inflated (depressed?), with very prominent Iniskinites cf. crassus Riccardi & Westermann, 1991 f/M primary and secondary costae (P c. 8; S c. 25) that crossed Fig. 5D–F straight over the venter. The apertural fragment has coarse, projected secondaries, but the flanks are missing. As far as cf. 1991 Iniskinites crassus Westermann & Riccardi, p. 63, pl. 12, can be made out, this fragment resembles large, broad- fig. 4; pl. 14, fig. 3. whorled eurycephalitines with coarse ribbing from the Andes, for example, the Upper Bathonian Iniskinites crassus MATERIAL:1 distorted phragmocone (C1775), from loose Ricc. et West. and, especially, the highly inflated and larger, block of c. 18 m of sandstones in basal Oraka Sandstone, Lower Callovian Eurycephalites? extremus (Tornq.) from R16/f403 at Whakapirau Road near Kawhia Harbour. (tentatively placed in Imlayoceras by Sandoval et al. 1990) This locality is of latest Temaikan to earliest Heterian age, as well as the poorly known Mexican E. cadoceroides with the former possibility indicated by the absence of the (Burckh.). Both Andean species are septate at c. 80 mm Heterian index (Retroceramus galoi), although its absence diameter (cf. Riccardi & Westermann 1991, text-fig. 27, pl. 7, from the associated collection may be a sampling effect. fig. 3a-c, pl. 13). The other possible genus our form may DESCRIPTION: The entirely septate shell was c. 70 mm in belong to is the mainly Late Bathonian Iniskinites. But the I. cf. crassus, described below from similar levels, is more diameter, but the last 1/3 whorl is preserved only with the finely ribbed. A Late Bathonian? to Early Callovian age is left flank. Whorls probably moderately inflated, with moderately small umbilicus (U/D c. 0.10), rounded umbilical indicated. shoulder, and probably ovate whorl-section with maximum Genus Iniskinites Imlay, 1975 width near 1/3 of height. Ornament consists of widely spaced, prominent primaries (P 10–12) that curve forward TYPE SPECIES: Kheraiceras magniforme Imlay, 1953. forming a shallow, adorally concave arc, and of almost 3 times as many secondaries that arise between 1/3 and 2/5 f Iniskinites gr. cepoides (Whiteaves, 1876) /M whorl-height and pass slightly projected over the venter. Fig. 8A–C Some of the secondaries arise by bifurcation, others by MATERIAL:2 distorted phragmocones (C1504, C1718) intercalation. There is no marked change in coarseness of from R17/f648, cepoides Association (Fauna 3), 12–15 m primaries or secondaries throughout the last 11/2 whorls of above base of Spellman Cliff section. the (complete?) phragmocone, except for possibly slightly wider spacing of the secondaries. DESCRIPTION: Shell globular with involute and strongly depressed ovate whorls, c. 40 mm in diameter and whorl REMARKS: The undiminished coarseness of the ornament width. Primary and secondary costae are sharp and dense, to c. 70 mm diameter strongly indicates that this form with 2 or 3 secondaries for each primary and division at belongs to the genus Iniskinites. In fact, this specimen closely maximum whorl width. Septate nearly to end, where resembles the two phragmocones of I. crassus illustrated primaries become rapidly coarser and longer. by Riccardi & Westermann (see synonymy), the others having their body chambers. Both agree in the similar REMARKS: There is close resemblance to highly inflated phragmocone diameter (D 70–80 mm), coiling (U/D .09), North American species, richly illustrated by Imlay (1953) and ribbing (P 10; S 28–30). In the absence of the body and Frebold (1978) from the Upper Bathonian of South chamber and because of the distortion, our specimen can Alaska and British Columbia, for example, I. magniforme, not be clearly identified specifically. I. cepoides, described I. martini, I. varicostatum, all Imlay spp., and I. robustus above, has much finer ribbing and is much more depressed.

Fig. 8 Eurycephalitinae from the upper Awakino valley; Faunas 1 and 3. All figures natural size unless otherwise indicated. ➤ Fauna 3 (see also Fig. 7B): A–C, I. gr. cepoides (Whiteaves) f/M, distorted and entirely septate specimen C1504, C with part of outer whorl removed, from 12–15 m above base of Spellman Cliff section (R17/f648). Fauna 1: D–T, All from R17/f566. D–G, X. grantmackiei West. & Hudson, topotypes; D, E, complete body chamber with crushed phragmocone C1423; F, crushed and damaged specimen C1734; G, X. cf. grantmackiei, juvenile, broken, somewhat crushed specimen C1735 showing coarse ribs on phragmocone. H–O, Araucanites awakino West., n. sp. m/m; H, holotype, C1732, complete and little distorted; I, J, somewhat distorted body chamber (?subadult) with damaged phragmocone C1741; K, crushed subadult, C1738; L, crushed subadult C1743; N, O, slightly crushed juvenile C1739. P–T, A. cf. awakino f/M; P, ultimate septum of subadult C1740; Q–T, juveniles; Q, distorted and crushed C1710; R, crushed C1711; S, T, small C1714 with obsolete primaries. Westermann et al.—Eurycephalitinae from New Zealand 515 516 New Zealand Journal of Geology and Geophysics, 2002, Vol. 45

I. crassus and relatives are known from the Late Bathonian a displaced fragment. The penultimate half-whorl has Steinmanni Zone of Argentina, Chile, and Mexico (Riccardi bifurcating subradial costae which pass straight across the & Westermann 1991). venter. The body chamber becomes somewhat more evolute The close resemblance to the single phragmocone and terminates at c. 70 mm diameter; costae rapidly become described from the basal Oxfordian (Dimorphus Zone) of extremely coarse, prominent and projecting. northern Chile by Hillebrandt & Gröschke (1995, pl. 5, This single specimen closely resembles the youngest fig. 3a, b) under “Eurycephalites? sp. A” is surprising and known Xenocephalites from the Southern Andes, puzzling. The specimen was said to occur in stratigraphic X. stipanicici from the Early Callovian Bodenbenderi and association with “Eurycephalites spp. B to F”, all of which Proximum Zones, which is associated in the Andes with are here placed in Araucanites. Was specimen “sp. A” X. involutus Ricc. & West. (?immature X. stipanicici) and misplaced from another (older) locality or does it belong to the macroconchs Eurycephalites rotundus (Tornqu.) and E.? a perfect homeomorph (in the phragmocone) of unknown extremus Ricc. & West. Only the latter, with highly depressed affinity? A range extension for Iniskinites from the Upper whorls and exceptionally coarse primaries, is large enough Bathonian into the Lower Oxfordian appears most unlikely. to be the macroconch of X. stipanicici. This suggests an Early No definite conclusion can be drawn, however, without Callovian age for the postawakino Association. knowledge of the body chamber. Genus Araucanites Westermann & Riccardi, 1985 Genus Xenocephalites Spath, 1928 TYPE SPECIES: Mayaites (Araucanites) stipanicici TYPE SPECIES: Macrocephalites neuquensis Stehn, 1923. Westermann & Riccardi (in Stipanicic et al. 1976).

Xenocephalites grantmackiei Westermann & Hudson, REVISED DIAGNOSIS: Macroconchs of medium to large 1991 m/m Fig. 5I, J, 8D–G size, extremely involute to obvolute and compressed, with 1988 Xenocephalites cf. neuquensis (Stehn, 1924) m, Westermann smooth flanks on all but innermost whorls. Microconchs & Callomon, p. 75, pl. 15, fig. 5a, b. roughly 1/2 sized, involute slender spherocones or 1991 Xenocephalites grantmackiei Westermann & Hudson, p. 690, platycones, strongly variocostate; phragmocone with fig. 3–1, 2. branching ribs that may fade away on inner flank; body chamber with gradually coarsening primaries and MATERIAL: Additional to the material described by secondaries, both becoming highly prominent toward Westermann & Hudson (1991): 1 (cf.) CE2106, distorted aperture. body chamber, probably from basal Oraka Sandstone, loose on shore platform, Oraka Bay, Totara peninsula, Kawhia REMARKS: Mainly on the grounds of its tight to obvolute Harbour (R15/f8550). Topotypes, 1, C1734, 1 (cf.) juvenile, (occluded) coiling and apparent biogeographic restriction C1735, from R17/f566. to the Southern Andes, the Araucanites macroconch was transferred from the Macrocephalitinae to the Eury- REMARKS: Body chamber CE2106 is a typical Xeno- cephalitinae as its latest (i.e., Oxfordian), member cephalites of the X. neuquensis-grantmackiei group, with (Westermann & Riccardi 1985). However, there remained a the deformation preventing determination of the direction long phylogenetic hiatus (“Lazarus effect”) between the Late of the primary ribs, distinguishing features for the closely Bathonian to Early Callovian macroconchs Eurycephalites interrelated Late Bathonian species. Three topotypes (C1423, and Lilloettia of the eastern Pacific margin and the Mid- C1734, C1735) are illustrated for comparison (Fig. 8D–G) Oxfordian Araucanites known only from west-central to show the coarse beginning of the body chamber of this Argentina, which could not be bridged (Westermann 1993). species that distinguishes it from the associated, more The only previously published occurrence of Araucanites abundant A. awakino n. sp. X. grantmackiei is of considerable was in the lower part of the La Manga Formation of chronostratigraphic importance, because it occurs rarely in southwest Mendoza and northeast Neuquen Provinces the Macrocephalites fauna of the Apertus Zone in Papua (Stipanicic et al. 1976). The ammonite-bearing Oxfordian New Guinea, which, in turn, has recently been extended to sequence there is only 4–16 m thick, in increasingly Nepal and found coeval with the European Discus Zone, regressive facies, and bound below by an unconformity latest Bathonian (Cariou et al. 1994; Westermann 1996b). It resting on strongly reduced Early and Middle Callovian. The is also closely allied with X. neuquensis (Stehn) from the Perisphinctes-Araucanites Assemblage Zone (Riccardi et al. Andean Steinmanni Zone. 1992, p. 140) is developed in carbonate facies with abundant clastics and oolites, and has yielded a possibly somewhat Xenocephalites cf. stipanicici Riccardi, Westermann & mixed assemblage of late Early and, mainly, early Mid- Elmi, 1989 m/m Fig. 6A Oxfordian Tethyan Ammonitina: Perisphinctes (Kranao- cf. 1989 Xenocephalites stipanicici Riccardi et al., p. 567, pl. 10, sphinctes) spp., P. (Arisphinctes) spp., Peltoceratoides cf. fig. 809. constantii (Orb.), Araucanites mulai, A. reyesi, A. stipanicici, cf. 1991 Xenocephalites stipanicici Riccardi & Westermann, p. 80, Westermann & Riccardi spp., and Euaspidoceras aff. pl. 18, fig. 3–6, pl. 19, fig. 1–2, text-fig. 29. waageni Spath. The main deposition presumably occurred in early Mid-Oxfordian time, when part of the assemblage, MATERIAL: Complete but crushed and damaged specimen including Araucanites spp., could have been redeposited C1736, from R17/f671, postawakino Association, from eroding Early Oxfordian strata. The recent finds of Whakapatiki/Awakino section, 17 m above base. similar eurycephalitines in the earliest Oxfordian of northern Chile (Hillebrandt & Gröschke 1995), however, dispel any REMARKS: The section of the last septate whorl is broadly doubt that the Araucanites are of Oxfordian age (not derived depressed and with rather dense costae, as discernible from from the Callovian). Westermann et al.—Eurycephalitinae from New Zealand 517

The Araucanites macroconch (f) is most closely related coarser costae on body chamber and phragmocone than the to the Late Bathonian to Early Callovian Lilloettia M/f from associated Araucanites postawakino m/m and could the east Pacific and New Zealand (see above), its probable correspond to the large macroconch ?Eurycephalites cf. ancestor. Araucanites species differ in the tighter umbilicus extremus (Tornq.), as in the Andes. with sharper margin, especially in the body chamber where Araucanites has previously been thought to have evolved the shell may occlude the umbilicus and, commonly, the from Eurycephalites rather than Lilloettia. The latter ranged greater whorl compression, as well as in the very early into the Early Callovian only in North America, but was disappearance or obscurity of primary costae. Lilloettia? replaced by Eurycephalites in South America in the earliest burckhardti Ricc. & West. M/f, from the Late Bathonian of Callovian (Riccardi & Westermann 1991, text-fig. 4; Mexico, and the related New Zealand form described above, Westermann 1993). The Eurycephalites clade, however, have similarly smooth flanks, but are much smaller than the developed from compressed to rotund forms, probably Oxfordian form and, at least in Mexico, with rounded, ending with the globular and coarsely costate E. extremus somewhat larger umbilicus. The poorly preserved New (Tornq.). Lilloettia is the much more likely ancestor, with Zealand form from the Late Bathonian grantmackiei phylogenetic increase of size, whorl compression, and Association, could, nevertheless, be the first member of the reduction of ornamentation. This is supported by the Araucanites lineage. The Eurycephalites macroconchs from occurrence in New Zealand of the macroconch L.? cf. the Lower Callovian of the Andes differ in the rounded whorl burckhardti Riccardi et al. and the associated microconch, section and the usual presence of primaries on most of the Araucanites awakino n. sp. The latest and largest New septate whorls. Some juvenile whorls of E. rotundus Zealand macroconch, A. spellmani, is also most closely allied (Tornq.), however, also resemble L.? cf./aff. burckhardti (cf. to the Andean, Mid-Oxfordian A. stipanicici type-species. Riccardi & Westermann 1991, pl. 4, fig. 4a, b). The latest Callovian to earliest Oxfordian eury- Based on several incomplete and subadult specimens cephalitines from the diverse Tethyan assemblage of northern from the Oraka Sandstone of Kawhia, the New Zealand Chile (Hillebrandt & Gröschke 1995) and the Early–Middle macroconchs have previously been misidentified with the Callovian species association of New Zealand, may poorly known genera Epicephalites Burckhardt and therefore, be the “missing link” bridging the former Mid- ?Subneumayria B. of the perisphinctacean family Aula- Callovian to Early Oxfordian phylogenetic hiatus in the costephanidae. Both are endemic genera from the late Early Eurycephalitinae clade (Westermann 2001). Curiously, this Kimmeridgian of northern Mexico (Arkell 1956; Arkell in Chilean assemblage includes also typical Iniskinites Fleming & Kear 1960; Stevens 1978; Hillebrandt et al. (Hillebrandt & Gröschke 1995, pl. 1, fig. 3a, b), a genus 1992). A plaster cast of the holotype of Epicephalites previously believed to have been confined to the Bathonian epigonus Burckhardt, type species and only illustrated to ?Early Callovian. However, the microconchs of the earlier specimen of the genus, is here again illustrated (Fig. 7). Araucanites species, including the large A. ponganui n. sp., E. epigonus differs from Araucanites macroconchs in the also have primaries on all septate whorls, in contrast to the more rounded whorls with larger umbilicus and the latest species, A. spellmani. rectiradiate, irregular secondary costae. In fact, E. epigonus is more closely homeomorphic to the Early Callovian Araucanites awakino Westermann n. sp. m/m and ?f/M Lilloettia burckhardti and Eurycephalites rotundus (Tornqu.) Fig. 5G, H, K–N, 6I–K, 8H–T from the Andes and Mexico (cf. Sandoval et al. 1990; Microconch: Riccardi & Westermann 1991). ?1956 Epicephalites cf. epigonus (Burckhardt), Arkell, p. 455. The clue to the distinction of Araucanites from Lilloettia, ?1960 Epicephalites cf. epigonus (Burckhardt), Fleming & Kear, however, lies in the microconchs. Those usually found in p. 24. most Araucanites associations, as well as with the possible 1987 Epimayaites sp., Hudson et al., p. 147, fig. 4. ancestral Lilloettia cf./aff. burckhardti, are distinct from 1997 Epicephalites marwicki Stevens, partim, p. 41, pl. 14, fig. Xenocephalites. Significantly, both occur together in the 6, 7 only, pl. 15, fig. 1. oldest, grantmackiei Association together with their Macroconch: presumed macroconchiate sexmorphs. There is a clear trend ?1997 Subneumayria (sp. indet.) cf. ordonezi (Burckhardt), of size increase throughout the Late Bathonian/Early Stevens, p. 44, pl. 15, fig. 2, 3, text fig. 30. Callovian to Middle Oxfordian Araucanites lineage, in the macro- and microconchs (see below). This includes the mid- DIAGNOSIS: Small, compressed. Microconch D 40–50 sized macro- and microconchs recently described under mm; phragmocone densely costate; mature Xenocephalites- “Eurycephalites” from the Callovian/Oxfordian boundary like modification confined to last quarter-whorl. beds of the Southern Andes (Hillebrandt & Gröschke 1995, Macroconch D 60–80 mm; secondaries very dense and pl. 5, fig. 4–6, pl. 6, fig. 3, 5), intermediate in age and size strongly projected, becoming smooth at D 20–25 mm. between the early, small New Zealand forms and the large, Oxfordian Argentine forms. However, as in other Eury- HOLOTYPE, TYPE LOCALITY, AND HORIZON: Microconch cephalitinae, this parallelism is imperfect, as exemplified C1732 (Fig. 8H), complete and little distorted on one side; by Lilloettia cf. burckhardti with smooth flanks and by from R17/f566, Te Whakapatiki Stream, upper Awakino A. ponganui that differs from A. spellmani only by the valley; Fauna 1. continuous presence of primaries. A similar phylogenetic size increase occurred in Xenocephalites, where the latest ETYMOLOGY: From Awakino River. species, X. stipanicici Riccardi et al., is also the largest one (cf. Riccardi & Westermann 1991). The X. cf. stipanicici OTHER MATERIAL: Microconch.Te Whakapatiki Stream: here described from the postawakino Association has much 12–15 (C1425, C1427, C1712, C1738, C1739 juvenile, 518 New Zealand Journal of Geology and Geophysics, 2002, Vol. 45

C1741–C1745, C1747, C1748), weakly to strongly distorted projected. The characteristically microconchiate orna- topotypes, in different states of completeness, from mentation is therefore only present on the ultimate 1/3–1/4 R17/f566, Fauna 1. Whakapatiki/Awakino section: 5 crushed whorl of fully grown specimens (i.e., above c. 30 mm diam.). and incomplete, C1737, C1751, C1753–C1755; 4 cf. Full adult size is 40–50 mm diameter. Many specimens (C1731, C1772–C1774) from R17/f375, Fauna 2. Kawhia appear to be completely preserved immatures with roughly Harbour area, middle Oraka Sandstone: 2 crushed, C1756 35 mm diameter. from R15/f216, C1757 from f219, Totara peninsula, Fauna The microconch differs from the associated Xeno- 2; cf. crushed fragments, C1758, C1759 from R16/f6048, cephalites grantmackiei Westermann & Hudson, by the finer Whakapirau Road, Fauna 2. Ponganui Farm Quarry, Port ornamentation of the ultimate whorl, especially in the less Waikato: cf. distorted but almost complete, CE2768 from strongly modified body chamber. R13/f7012, Fauna 2a. Two specimens (C1425, C1427) cited Comparison of this form with American species is above were previously included by Westermann & Hudson difficult because of the poor knowledge of the juvenile whorl (1991) in Lilloettia aff. boesei and are here re-allocated to in most species. For example, the only Andean species with this species. A concretion from the type Oraka Sandstone known juvenile whorls is Xenocephalites gottschei (Tornqu.) (R15/f8550 including C1781) containing at least four (cf. Riccardi & Westermann 1991, pl. 17, fig. 4a, b, 6a, b; specimens identified as Epimayaites sp. and figured by Parent 1997, fig. 1, 9–11), which has similar costation but Hudson et al. (1987) are also located here. is much more inflated and has been matched with Eurycephalites vergarensis (Burckh.) (Parent 1997). Macroconch (=A. cf. awakino). None of the specimens There also appears to be some resemblance to poorly is well enough preserved to be morphologically matched known North American forms. Interestingly, the Early with the microconch, which included the holotype. Species Callovian “hypotypes of Lilloettia lilloetensis Crickmay” determination of the macroconchs is based on association from British Columbia illustrated by Frebold & Tipper with the A. awakino microconch and analogy to closely (1967, ?pl. 1, fig. 7, 8; pl. 3, fig. 3) are either incomplete related species with better known dimorphism. All microconchs (i.e., Xenocephalites) or juveniles (and ?inner macroconchs are therefore named A. cf. awakino, and an whorls) of macroconchs. Another widespread North allotype cannot be designated. Te Whakapatiki Stream: at American species is X. vicarius Imlay, which, like our New least 10 juvenile and/or incomplete, more or less strongly Zealand form, may include immature or incomplete crushed specimens (all cf. including C1710, C1711, C1713– specimens and therefore lacks the adult modification (e.g., C1715) from R17/f566, Fauna 1. Totara peninsula, Kawhia X. hartsocki Imlay (1953, cf. pl. 28, fig. 1–8 versus Fig. 8H, Harbour, Oraka Sandstone: ?2 small, crushed fragments I, L)). The septate whorls differ from those of our species CE922, CE923 (=“Subneumayria sp. indet., cf. ordonezi” by the coarser primary costae and/or more densely spaced of Stevens 1997, incl. fig. 30); 3 incomplete body chambers, secondaries at comparative diameters. X. hebetus Imlay has one with remnants of phragmocone, C374, CE2725, CE2726 similar ornamentation but more inflated whorls. The new (=“Epicephalites marwicki” partim Stevens 1997); 3 cf. half- New Zealand species A. postawakino m/m differs mainly in whorl body chambers, crushed C1729, C1730, C1780, all the larger diameter. from R15/f8550, Fauna 2; 1 cf. body chamber fragment In recording Epimayaites from the Oraka Sandstone, C1733, middle part of Oraka Sandstone, R15/f217, Fauna Hudson et al. (1987) referred to the record by Thomson 2. Ponganui Farm Quarry: 1 cf., incomplete, crushed CE2769 (1982) of E. aff. transiens (Waagen) from the South Shetland and 1 cf., fragmental CE2770, R13/f7012, Fauna 2a. Islands. This record should also be re-examined as a possible Araucanites in the light of the New Zealand occurrences of DESCRIPTION AND REMARKS: Microconch. Phragmocone this latter genus. diameter 25–30 mm (undistorted in incomplete specimen Fig. 8P). Involute (U/D 0.10) with ovate whorl section, as Macroconch. Small (D 70–80 mm), compressed dis- high as wide, venter narrowly rounded and maximum width cocone. Ribs on body chamber very dense, projected, and at 1/3 whorl height, that is, a typical discocone (compressed on outer flank and venter only, withdrawing to outer third spherocone). Ornamentation of juvenile at c. 20 mm diameter of flank near aperture. Most specimens have incomplete consists of very dense, long, concave primaries (P 12–15) body chambers at <40 mm diameter, while others are that divide at 2/5 whorl height into dense, rectiradiate to adolescents. Whorls are very involute (U/D c. 10%) and have slightly prorsiradiate secondaries. Subsequently, primaries ovate sections, probably about as wide as high. Juvenile become more widely spaced (P 8–12) and divide at about ornamentation of internal moulds up to 20–25 mm diameter 1/3 whorl height; secondaries remain dense, c. 3 per primary, consists of thin and somewhat irregular primaries that and somewhat prorsiradiate. This change in primary costae become fainter at end of this stage, and dense, prorsiradiate occurs about one full whorl before adult aperture. Septum secondaries; subsequently, inner and, finally, middle flanks and suture are as in typical Sphaeroceratidae, with several become smooth (shell surface unknown) except for irregular complete saddle axes and a graded series of internal and striae, with very dense, projected costae restricted to outer external lobes; saddle envelope is radial to protracted flanks and venter. Septum and suture are poorly preserved. (markedly raised at umbilicus). The poorly preserved specimens from the middle Oraka Body chamber is mesodomic (c. 3/4 whorls) and egresses Sandstone of Kawhia Harbour include most of the specimens at the umbilicus strongly only with ultimate half-whorl, originally referred to as Epicephalites cf. epigonus and commonly producing elliptical coiling. On ultimate half- Subneumayria cf. ordonezi by Arkell (1956 and in Fleming whorl, section becomes subcircular and costation pro- & Kear 1960) and later partly named Epicephalites marwicki gressively more Xenocephalites-like: primaries grow shorter Stevens (1997), apart from its holotype which is here placed and more prominent, secondaries much coarser and in Lilloettia (see synonymies). Westermann et al.—Eurycephalitinae from New Zealand 519

The macroconch closely resembles the small Lilloettia (aperture U/D 0.15). Ornament becomes rapidly Xeno- with early smoothening of the inner flanks recorded from cephalites-like on ultimate half-whorl by shortening and the Steinmanni Zone of Mexico as L. steinmanni by coarsening of primaries and by extreme coarsening and Sandoval et al. (1990), and recently distinguished as forward-inclination of the secondaries. Peristome is oblique subspecies L. steinmanni burckhardti by Riccardi & but apparently not strongly modified. Westermann (1991). The latter authors maintained that the more abundant larger forms associated with it belong to the Macroconch is unknown. same, relatively inflated (geographic) subspecies. The larger forms, however, have primaries to the end of the phrag- REMARKS: The microconch is closest to that of A. mocone at c. 50 mm diameter (Sandoval et al. 1990), whereas ponganui, differing mainly by the smaller diameter. The in burckhardti the inner flank is already smooth at 30 mm. somewhat younger A. spellmani differs mainly in the The more subrectangular whorl section of the Mexican form narrower umbilicus and smooth inner flanks of the inner (which makes it morphologically intermediate to Eury- whorls. The older A. awakino is much smaller and has more cephalites), however, cannot be observed on our distorted rounded whorls. and smaller specimens. Yet Sandoval et al. (1990) did not recognise bimodality in the relatively large sample (39, but Measurements (mm): from different levels) between the large, more strongly DWbWhU costate forms and the small forms with reduced orna- Holotype, apert. 60 27.2 28.0 11.2 mentation. end phr. 39 20.5 20.0 6.0 It is interesting to note that internal moulds with early smoothening of the flanks have been reported together with more extensively costate forms also in other species, for Araucanites ponganui Westermann n. sp. m/m example, L. cf. lilloetensis from the Early Callovian of Fig. 9A, B British Columbia (Frebold 1978, pl. 8, fig. 2, 3), where these DIAGNOSIS: Microconch large (D 50–80 mm); primaries modifications have been considered as mere variants. present on ultimate septate whorl. Significantly, the early smoothening of the flanks of highly involute and compressed whorls is also the characteristic HOLOTYPE AND TYPE LOCALITY: C1519, complete but juvenile feature of the much later (Oxfordian), much larger somewhat crushed and damaged (Fig. 9A); from R13/f7012, Araucanites macroconchs, suggesting their evolutionary small farm quarry beside Ponganui Road and Opuatia pathway. Stream, Port Waikato. m Araucanites postawakino Westermann n. sp. /m ETYMOLOGY: After road at type locality. Fig. 6E–L DIAGNOSIS: Microconch diameter c. 60 mm; whorls OTHER MATERIAL: Crushed topotypes from Ponganui Farm compressed and moderately evolute; primaries well Quarry: 4 (C1518, C1520, cf., large fragment CE2771, from developed on phragmocone. R13/f7012, C1765 from R13/f125). 1 (C1767) from lower part of Spellman Cliff, R17/f302. HOLOTYPE, TYPE LOCALITY, AND HORIZON:The well- preserved, complete microconch (C1749) with remnants of DESCRIPTION:Relatively large microconch (D 52–80 mm). zeolitised shell (Fig. 6E–H); from Fauna 2, Whakapatiki/ Outer whorls are compressed discoconic, becoming rounded Awakino section, 23–25 m above base, R17/f673. ovate with body chamber; very involute (internal mould U/ D c. 0.05) with somewhat rounded margin. Ornament of ETYMOLOGY: Referring to the similar, but somewhat older widely spaced, moderately coarse primaries and dense A. awakino described above. secondaries continuous to end of adult phragmocone, somewhat prosocline and crossing straight over venter. On OTHER MATERIAL:1 crushed and incomplete microconch body chamber, primaries and secondaries become C1750 from R17/f672, ?1 crushed and incomplete increasingly prominent; widely spaced primaries (P c. 10) microconch C1752 from R17/f375, Whakapatiki/Awakino are often blade-like, dividing at mid-flank into 4 or 5 section. secondaries, which coarsen rapidly and become more strongly projected, with only 2 per primary. Near-adult DESCRIPTION: The adult diameter of the microconch is aperture ornamentation is extremely coarse and continuous c. 60 mm. Septate whorls are subtrapezoidal, slightly wider over flanks and venter. than high (W/H 1.05), and involute (U/D 0.10). Umbilical walls are steep and rounded. Ornamentation of phragmocone REMARKS: This microconch, when complete with body end consists of curved and moderately coarse primaries (P chamber, closely resembles A. spellmani m/m, from which 8–10) on the inner 2/5 of the flank, and 3–4 times as many, it can be distinguished only (or mainly) by the primary ribs somewhat prorsiradiate, dense secondaries which cross over present on the last septate whorl. Among the microconchs, the venter. Secondaries arise mostly by irregular intercalation the immature whorls of A. ponganui thus are as in the smaller between 2/5 and 1/2 whorl height. Septal suture is and older A. postawakino, and the body chamber as in the moderately complex with gradually reducing saddles along later A. spellmani, of equal size. straight envelope that rises slightly toward umbilicus. A fragment of a large macroconch (C1776) (D c. 120 Body chamber is 3/4 whorls, retains the same section as mm), found together with the holotype, has a densely costate phragmocone but egresses gradually and moderately venter while the flanks appear to be smooth, as in adult 520 New Zealand Journal of Geology and Geophysics, 2002, Vol. 45

Fig. 9 Faunas 2a–4. All figures natural size. Fauna 2a, 3 (see also Fig. 4A–C): A, B, Araucanites ponganui Westermann, n. sp. m/m; A, holotype, complete specimen C1519 with crushed body chamber and inner whorl, from Ponganui Farm Quarry, Port Waikato (R13/f7012), Fauna 2a; B, complete, distorted specimen C1767 from lower part of Spellman Cliff (R17/f302), Fauna 3. Fauna 4: C–M, Microconchs (m) of Araucanites spellmani ➤ Westermann et al.—Eurycephalitinae from New Zealand 521

Araucanites. But the whorl-section probably was much more subovate to trapezoidal, extremely involute to obvolute depressed, as in Early Callovian Lilloettia (Imlayoceras), (occluded). Umbilical wall vertical with narrowly rounded unless this is due to orthogonal deformation. Additional or sharp margin, becoming overhanging at end of body material is required to solve this taxonomic problem. chamber; minute umbilicus of body chamber elliptical with occlusion towards end; missing shell presumably closed Araucanites spellmani Westermann n. sp. f/M and m/m umbilicus. Flanks weakly convex and converging from Fig. 9C–M, 10A–G umbilical margin to rounded venter. Body chamber 3/4 to DIAGNOSIS: Macroconch a mid-sized Araucanites with full whorl. Innermost whorls up to D 20–25 mm, poorly very dense secondaries to aperture. Microconch relatively known. Ornamentation above D 25 mm consists only of large, with smooth inner flanks on mature phragmocone. strongly prosocline, dense secondaries (S 42–48) on outer half of whorl, withdrawing to Wh 1/3–1/4 on adult body HOLOTYPE, TYPE LOCALITY, AND HORIZON: The macro- chamber. Inner flank essentially smooth, except for faint, conch (f) C1521, complete body chamber, somewhat distant, curved riblets or fasciculate striae. Secondaries cross damaged and distorted, mostly internal mould (Fig. 10A, venter slightly convex. There is a remarkable difference in B); from R17/f619, Spellman Cliff section (40–100 m above rib prominence and acuteness between shell and internal base) in upper Awakino valley. mould. Septal suture poorly preserved. The macroconch resembles that of Mid-Oxfordian ALLOTYPE: The microconch (m) C1522, complete but Araucanites from Argentina, but differs in the retention of possibly not fully grown, partly with shell (Fig. 9C, D), from ventral costae. The closest is A. stipanicici West. & Ricc., R17/f645, Spellman Cliff section, c. 80 m above base; which differs in the coarser ribs and larger size. A. mulai attached to A. spellmani f/M, C1523 (Fig. 10G). W. & R. has subparallel flanks; and A. reyesi W. & R. is more inflated and larger. PARATYPES The subadult macroconch C1523 (Fig. 10E– The best match is probably to the single specimen of G) attached to allotype and complete but crushed microconch “Eurycephalites sp. B” of Hillebrandt & Gröschke (1995, C1527 (Fig. 9E, F), from R17/f619, 30–90 m above base, pl. 6, fig. 1a, b) from the earliest Oxfordian Dimorphus Zone Spellman Cliff section. The large, crushed microconch of northern Chile, which also is accompanied by similar C1768 (Fig. 9G, H), from R17/f617, float block, Waterfall microconchs (see below). This Chilean form (of which we Creek. have a plaster cast) is said to have a latest Callovian to Early Oxfordian range. It has a similar size and the typical, ETYMOLOGY: In honour and appreciation of Mr and Mrs occluded umbilicus, but may differ in the more developed John Spellman on whose farm the species has mainly been primaries of the septate whorls. The body chamber is more found. rounded and less tightly coiled in the internal mould, but its thick shell (partially preserved on the outer flank of the OTHER MATERIAL: Macroconch. Spellman Cliff: more or peristome) presumably occluded the umbilicus. This Chilean less strongly deformed and mostly incomplete and/or not specimen, however, is probably not fully grown, as is fully grown internal moulds: 4 adults (C1529 from 40–100 apparent in comparison to the microconch (see below). m above base of section, R17/f619; C1720, C1722, C1723 Alternatively, there may have been two species, with the from c. 80 m above base of section, R17/f645; and (cf.) 4 large and small microconchs representing species rather than juveniles (C1721, C1724–C1726), from c. 80 m above base growth stages and/or variants. Two fragmentary Chilean of section, R17/f645; 1 (C1727), from loose block (together specimens (Hillebrandt & Gröschke 1995, pl. 6, fig. 2, 4) with microconch C1728), R17/f302; (cf.) 1 (C1779) poorly differ in the larger umbilicus; one has smooth flanks and preserved, from 50–55 m above base of section, R17/f652; could be an exceptionally compressed Eurycephalites (cf.) 1 (C1719) almost complete, from Waterfall Creek, R17/ macroconch, the other has a flattened venter (tabulate) and f615. Microconch. More or less complete but crushed is of unknown affinity. M. Gröschke (pers. comm. 1995) specimens from Spellman Cliff: 5 (C1524–6, C1528, agrees with our assessment of the close affinity between the ?C1762), from 40–100 m above base of section, R17/f619 Chilean and New Zealand forms. and 1 (C1728) from loose block, R17/f302; (cf.) 1 (C1760), crushed, from 65 m above base, R17/f643; (cf.) 1 (C1778), Microconch. Complete diameter (D) is 52–80 mm, with incomplete, from c. 75 m above base, R17/f644; ?2 (C1763, holotype at lower limit (chosen for preservation). Whorls C1764), from c. 80 m above base, R17/f645; ?fragment moderately compressed subovate-trapezoidal, becoming (C1761), from c. 100–105 m above base, R17/f647. (cf.) 1 rounded-ovate on body chamber, very involute (internal (C1769), from Gribbons Road, R17/f620; (cf.) 1 (C1777), mould U/D c. 0.05), with subcylindrical umbilicus and float, from Waterfall Creek, R17/f613. somewhat rounded margin; egressing moderately at end of body chamber. Phragmocone with weakly convex, slightly DESCRIPTION AND REMARKS: Macroconch. Complete at converging flanks and broadly rounded venter. 100–130 mm diameter. Whorls moderately compressed, Ornamentation of innermost whorls to D 20–25 mm

West., n. sp.; C, D, allotype C1522, almost complete and little distorted (attached to A. spellmani macroconch/f, C1523, Fig. 10G), from Spellman Cliff section, c. 80 m above base (R17/f645); E, F, paratype C1527 with (E) and without (F) part of body chamber, from Spellman Cliff (R17/f619); G, H, paratype C1768, almost complete, lateral latex impression (G) and inner whorl with end of body chamber (H), from R17/f617, float, “Waterfall creek”, tributary to Awakino River; I, J, large, almost complete specimen C1528 with part of body chamber (transposed), from Spellman Cliff (R17/f619); K, L, undistorted, incomplete specimen C1524, from R17/f619; M, large and complete specimen C1526, but deformed with enlarged umbilicus, from R17/f619. 522 New Zealand Journal of Geology and Geophysics, 2002, Vol. 45

Fig. 10 Fauna 4. Spellman Cliff section. All figures natural size. A–G, Macroconchs (f) of Araucanites spellmani Westermann, n. sp.; A, B, holotype C1521, complete but distorted and somewhat crushed, from middle or upper part of section, 40–100 m above base (R17/f619); C, D, almost complete C1529, one side crushed, same locality; E–G, paratype C1523, complete subadult, mostly undistorted; ventral view (E), lateral view (F) and oblique view (G) together with allotype (m) C1522, from c. 80 m above base of section (R17/f645) (see Fig. 9C, D). Westermann et al.—Eurycephalitinae from New Zealand 523 unknown. Outer whorls have very dense secondaries (S c. has been noted by Grant-Mackie et al. (2000); it closely 43), somewhat prosocline on outer half of whorl and cross resembles the only pseudoperisphinctine species known from straight over venter; primaries are weak or obsolete on inner Indonesia, C. (H.) cf./aff. furcula from the late Early whorls. At or near end of phragmocone, widely spaced Callovian Macrocephalites keeuwensis Association of the primaries arise on umbilical slope and extend adorally Sula Islands (Westermann & Callomon 1988, p. 76, pl. 17, concave to middle of flank. Secondaries of uneven length fig. 2–4). Similar morphotypes, however, occur in Europe begin here by fasciculation and irregular intercalation. On in the Late Bathonian. body chamber are very prominent, often blade-like primaries (P c. 10); secondaries at first dense, subradial to somewhat prorsiradiate with 4 or 5 per primary, but progressively Subfamily PERISPHINCTINAE Steinmann, 1890 become coarser, more projected, with reduction to 2 per Genus Sulaites Oloriz & Westermann, 1998 primary. Near-adult aperture at D 60–80 mm, extremely TYPE SPECIES: Perisphinctes sularus Boehm, 1907, from prominent, projected ribs are continuous over flanks and the Sula Islands, Indonesia. venter. Peristome oblique and somewhat “constricted” by blunted ribs and followed by “collar”. Septal suture poorly COMMENTS: The genus includes evolute and serpenticonic, preserved; ultimate suture markedly protracted (raised at small perisphinctids. The ovate whorls are depressed in umbilicus) in C1761. juveniles and compressed in adults. Ribbing is mainly The microconch closely resembles several of the recently dichotomous and ventrally projected, but interrupted on illustrated fragments of “Eurycephalites ? n. spp.” from the internal mould of inner whorls, and forming chevrons on uppermost Callovian and earliest Oxfordian of north Chile outer whorls. The inner whorls tend to have parabolic ribs (Hillebrandt & Gröschke 1995). The authors have kindly and constrictions; the outer whorls incipient to variable sent us plaster casts. Of their large specimens, one is ventral parabolae. complete with damaged peristome and venter; the other is Sulaites, named for the “Perisphinctes sularus- an adult body chamber fragment (their pl. 5, fig. 4; pl. 6, moluccanus group” (Francis & Westermann 1994), occurs fig. 3). Both closely resemble our larger, fully grown in the Late/latest Oxfordian and Early Kimmeridgian specimen (C1528) from Spellman Cliff (Fig. 9I, J). Three throughout the Himalayan Bioprovince (e.g., southern of their smaller specimens (their pl. 5, fig. 5, 6a, b; p. 6, Himalayas, Indonesia, Papua New Guinea, and New fig. 5) are close to our smaller specimens (C1522, C1524, Zealand; Oloriz & Westermann 1998). C1527; Fig. 9C–F, K, L), some of which appear not to be fully grown. Sulaites heteriensis (Stevens, 1997) The body chamber of the microconch closely resembles 1997 Idoceras heteriense Stevens, p. 46–48, pl. 15, fig. 4–9, pl. 16, that of A. ponganui n. sp. in the late development of the fig. 1, text-fig. 31. characteristic modification, but differs by the smooth inner flank preceding the body chamber. COMMENTS: Identification of supposed Idoceras with Mexican affinity from New Zealand goes back to Arkell Measurements (mm): (1956). This opinion, together with its important bio- DWhWbU geographic implications, has been followed by several New Macroconchs: Zealand authors until recently (Stevens 1997 and record Holotype, end b.ch. c. 115 c. 60 c. 50 7.5/occl. therein). Some years ago, Westermann (e.g., in Francis & Paratype C1523, subadult 61 36 32 3.0 Westermann 1994), however, recognised the close affinity Microconchs: of the great majority of New Zealand “Idoceras” (i.e., “I.” Allotype, apert. 53 25.4 25.3 5.7 heteriense), with “Perisphinctes gr. sularus-moluccanus”, end phr. c. 30 16.8 – c. 2.5 especially Sulaites gerthi Oloriz & West. from New Guinea C1528, b.ch. 62 31.5 c. 30 7.0 (=“Pseudoparaboliceras aramaraii” Gerth, 1965; nomen C1524, b.ch. 48 26 23 4.3 nudum, no type species or holotype designated). Compared to the “Tethyan” Idoceras, the whorls of Sulaites, including S. heteriensis, are more evolute and the ribs more projected, Superfamily PERISPHINCTACEAE Steinmann, 1890 with chevrons and parabolae. A recent review of Jurassic PERISPHINCTIDAE Family Steinmann, 1890 biogeography (Grant-Mackie et al. 2000) has also included PSEUDOPERISPHINCTINAE Subfamily Schindewolf, Sulaites in the New Zealand fauna. 1923 The exact locality and stratigraphic position of the Genus Choffatia Siemiradzki, 1898 holotype are unknown and its reported probable origin in Subgenus Homoeoplanulites Buckman, 1922 the Oraka Sandstone (Stevens 1997) is unlikely. The species TYPE SPECIES: H. homoeomorphus Buckman, 1922. occurs quite abundantly, however, in the higher Ohineruru Formation of Kawhia Harbour. Choffatia (Homoeoplanulites) ex gr. furcula (Neumayr, 1871) MATERIAL: Incomplete whorls C1770, from R17/f648, (or ?Family NEOCOMITIDAE Salfeld, 1921) c. 15 m above base of Spellman Cliff, Fauna 3. Genera Kossmatia Uhlig, 1907 and Paraboliceras REMARKS: The outer whorls are evolute with compressed Uhlig, 1910 ovate section and dense, regular costation. The primaries divide in twos and threes at mid-flank and smoothen COMMENTS: Kossmatia (type sp. Am. tenuistriata Gray) somewhat at mid-venter. The occurrence of this fragment typically has medium-coiled, compressed whorls with dense 524 New Zealand Journal of Geology and Geophysics, 2002, Vol. 45 ribbing ending in ventral chevrons, without parabolae, but Damborenea, S. E.; Manceñido, M. O. 1992: A comparison of includes more evolute and coarsely ribbed forms. The latter Jurassic marine benthonic faunas from South America and resemble morphotypes of Paraboliceras (type sp. Am. jubar New Zealand. Journal of the Royal Society of New Zealand 22: 131–152. Blanford) without parabolae, except for the absence of a ventral groove. Particularly when the stratigraphic record is Damborenea, S. E.; Polubotko, I. V.; Sey, I. I; Paraketsov, K. V. 1992: Bivalve zones and assemblages of the circum-Pacific poor, it then comes down to a more subjective choice region. In: Westermann, G. E. G. ed. The Jurassic of the between the diagnostic features. Stevens (1997) stressed the circum-Pacific. New York, Cambridge University Press. absence of parabolae in his generic classification of Pp. 300–307. Kossmatia macnaughti, while most authors disregard the Enay, R.; Cariou, E. 1996: Identification du Kimméridgien du incipient parabolae and instead stress the presence of a domaine Indo-Sud-Ouest Pacific: la faune à Paraboliceras ventral groove between weakly projected ribs, and place this (Ammonitina) de l’Himalaya à la Nouvelle-Zélande. species in Paraboliceras. Clearly, these genera are closely Compte rendu de l’Académie des Sciences Paris Ser. HII, interrelated and could be classified as subgenera only. 322: 449–474. Fleming, C. A.; Kear, D. 1960: The Jurassic sequence at Kawhia Harbour, New Zealand. New Zealand Geological Survey ACKNOWLEDGMENTS Bulletin 67. Francis, D. A. 1977: The Upper Jurassic Captain King’s Shellbed John H. Callomon, London, has given valuable advice. Brian in the Kawhia–Mahoenui region, south-west Auckland. Challinor, Hamilton, New Zealand, has contributed a valuable Unpublished MSc thesis, University of Auckland, specimen. Mr and Mrs John Spellman, Mahoenui, New Zealand, Auckland, New Zealand. 106 p + map. have repeatedly permitted our work on their farm property and have also been generous hosts during those visits. Essential Francis, G.; Westermann, G. E. G. 1994: The Kimmeridgian financial support for travel and fieldwork was granted to G.E.G.W. problem in Papua New Guinea and other parts of the Indo- by the National Science and Engineering Research Council of South-west Pacific. In: Carman, G. J.; Carman, Z. ed. Canada. We also thank Louise Cotterall, University of Auckland, Petroleum exploration and development in Papua New for photography; Isabel Sutherland and Keith Johnston, University Guinea. Proceedings of the 2nd Papua New Guinea of Auckland, for making collections available to us and coping Petroleum Conference, Port Moresby, 1993. Pp. 75–93. with numerous questions; Craig Jones and Graeme Stevens, Frebold, H. 1964: Illustrations of Canadian fossils—Jurassic of Institute of Geological & Nuclear Sciences, for the loan of western and Arctic Canada. Geological Survey of Canada specimens. Paper 64-3. 106 p. Frebold, H. 1978: Ammonites from the Late Bathonian Iniskinites fauna of central British Columbia. Geological Survey of REFERENCES Canada Bulletin 307. 26 p. Arkell, W. J. 1956: Jurassic geology of the world. London, Frebold, H.; Tipper, H. W. 1967: Middle Callovian sedimentary Edinburgh, Oliver & Boyd. rocks and guide ammonites from southwestern British Columbia. Geological Survey of Canada Paper 67-21. Ballance, P. F.; Campbell, J. D. 1993: The Murihiku arc-related 29 p. basin of New Zealand (Triassic–Jurassic). In: Ballance, P. F. ed. South Pacific sedimentary basins. Sedimentary Gerth, H. 1965: Ammoniten des mittleren und oberen Jura und basins of the world, 2. Amsterdam, Elsevier. Pp. 21–33. der alteren Kreide vom Nordabhang des Schneegebirges in Neu Guinea. Neues Jahrbuch der Geologie und Boehm, G. 1907: Die sudkusten der Sula-Inseln Taliabu und Paleontologie Abhandlungen 121: 209–218. Mangoli. 3-Abschnitt: Oxford des Wai Galo. Beitrage zur Geologie von Niederlandisch-Indien 1, Paleontographica Grant-Mackie, J. A.; Aita, Y.; Balme, B. E.; Campbell, H. J.; Suppl. 4, Abt.1: 61–120. Challinor, A. B.; MacFarlan, D. A. B.; Molnar, R. E.; Stevens, G. R.; Thulborn, R. A. 2000: Jurassic palaeo- Burckhardt, B. E. 1906: Le faune jurassique de Mazapil. Boletin biogeography of Australasia. Memoir of the Australasian del Instituto Geologico de Mexico 23. Association of Palaeontologists 23: 311–353. Callomon, J. H. 1984: A review of the biostratigraphy of the post- Helby, R; Wilson, G. J.; Grant-Mackie, J. A. 1988: A preliminary Lower Bajocian Jurassic ammonites of western and study of Middle and Late Jurassic dinoflagellate northern North America. In: Westermann, G. E. G. ed. assemblages from Kawhia, New Zealand. Memoir of the Jurassic–Cretaceous biochronology and paleogeography of Association of Australasian Palaeontologists 5: 125–166. North America. Geological Survey of Canada Special Paper 27: 143–174. Hillebrandt, A. v.; Gröschke, M. 1995: Ammoniten aus dem Callovium/Oxfordium Grenzbereich von Nordchile. Callomon, J. H. 1985: The evolution of the Jurassic ammonite Berliner Geowissenschaftliche Abhandlungen Reihe A 169. family Cardioceratidae. Special Papers in Palaeontology 40 p. 33: 49–90. Hillebrandt, A. v.; Smith, P.; Westermann, G. E. G.; Callomon, J. Campbell, H. J.; Grant-Mackie, J. A.; Paris, J.-P. 1985: Geology H. 1992: Ammonite zones of the circum-Pacific region. of the Moindou–Teremba area, New Caledonia. Strati- In: Westermann, G. E. G. ed. The Jurassic of the Circum- graphy and structure of Teremba Group (Permian–Lower Pacific. New York, Cambridge University Press. Pp. Triassic) and Baie de St. Vincent Group (Upper Triassic– 247–260. Lower Jurassic). Geologie de la France 1: 19–36. Hudson, N. 1983: Stratigraphy of the Ururoan, Temaikan and Cariou, E.; Enay, R.; Bassoulet, J.-P.; Colchen, M. 1994: Heterian Stages; Kawhia Harbour to Awakino Gorge, Biochronologie du Jurassique moyen de la Thakkhola South-west Auckland. Unpublished MSc thesis, University (Nepal central). Compte rendu de l’Academie Paris Serie of Auckland, Auckland, New Zealand. 168 p. + maps and 318: 93–100. columns. Damborenea, S. E. 1990: Middle Jurassic inoceramids from Hudson, N. 1999: The middle Jurassic of New Zealand: a study of Argentina. Journal of Paleontology 64: 736–759. the lithostratigraphy and biostratigraphy of the Ururoan, Damborenea, S. E. 1993: Early Jurassic pectinaceans and circum- Temaikan and Lower Heterian Stages (?Pliensbachian to Pacific paleobiogeography. Palaeogeography, Palaeo- ?Kimmeridgian). Unpublished PhD thesis, University of climatology, Palaeoecology 100: 109–123. Auckland, Auckland, New Zealand. 329 p. Westermann et al.—Eurycephalitinae from New Zealand 525

Hudson, N.; Grant-Mackie, J. A.; Helby, R. 1987: Closure of the Sukamto, R.; Westermann, G. E. G. 1992: Indonesia and Papua New Zealand “Middle Jurassic Hiatus”? Search 18: New Guinea. In: Westermann, G. E. G. ed. The Jurassic 146–148. of the Circum-Pacific. New York, Cambridge University Imlay, R. W. 1953: Callovian (Jurassic) ammonites from the United Press. Pp. 181–183. States and Alaska, Part II: Alaska Peninsula and Cook Inlet. Thomson, M. R. A. 1982: Late Jurassic fossils from Low Island, United States Geological Survey Professional Paper 249B: South Shetland Islands. British Antarctic Survey Bulletin 41–108. 56: 25–35. Krumbeck, L. 1934: Die Aucellen des Malms von Misol. Neues Trechmann, C. T. 1923: The Jurassic rocks of New Zealand. With Jahrbuch für Mineralogie, Geologie und Paläontologie 71 an Appendix on ammonites from New Zealand by L. F. (B): 422–469. Spath. The Quarterly Journal of the Geological Society of MacFarlan, D. A. B. 1975: Mesozoic stratigraphy of the Marokopa London 79 (3): 246–312, pl. XII–XVIII. area. Unpublished MSc thesis, University of Auckland, Westermann, G. E. G. 1993: Limits of global bio-event correlation: Auckland, New Zealand. 185 p. + maps and section. diachronous ammonite “extinction” across Jurassic Marwick, J. 1953: Divisions and faunas of the Hokonui System bioprovinces. Revista Asociation Geologica Argentina 47: (Triassic and Jurassic). New Zealand Geological Survey 353–364. Paleontological Bulletin 21. 141 p. 16 pl. Westermann, G. E. G. 1994: Correlating Australasian regional Meesook, A.; Grant-Mackie, J. A. 1995: Upper Jurassic stages: New Zealand Jurassic. Australasian Palaeon- stratigraphy, south Kawhia region, New Zealand. New tological Convention, February, 1994, Abstracts and Zealand Journal of Geology and Geophysics 38: 363–375. Programme. P. 63. Oloriz, F.; Westermann, G. E. G. 1998: The perisphinctid ammonite Westermann, G. E. G. 1996a: Correlating New Zealand regional Sulaites n. gen. from the Upper Jurassic of the Indo- stages by ammonites. In: Riccardi, A. C. ed. Advances in Southwest Pacific. Alcheringa 22: 231–240. Jurassic research. GeoResearch Forum 1–2: 93–100. Parent, H. 1997: Ontogeny and sexual dimorphism of Eury- Westermann, G. E. G. 1996b: New mid-Jurassic Ammonitina from cephalites gottschei (Tornquist) (Ammonoidea) of the New Zealand: implications for biogeography and Andean Lower Callovian. Geobios 30: 407–419. oceanography. In: Riccardi, A. C. ed. Advances in Jurassic research. GeoResearch Forum 1–2: 179–186. Riccardi, A. C.; Westermann, G. E. G. 1991: Middle Jurassic ammonoid fauna of the Argentine–Chilean Andes, Part IV: Westermann, G. E. G. 1996c: Circum-Gondwanan ammonite Bathonian–Callovian Eurycephalitinae, Stephan- correlation at the Bathonian-Callovian boundary. In: ocerataceae. Palaeontographica A 216. 110 p. Riccardi, A. C. ed. Advances in Jurassic research. GeoResearch Forum 1–2: 485–492. Riccardi, A. C.; Westermann, G. E. G.; Elmi, S. 1989: The Bathonian–Callovian ammonite zones of the Argentine– Westermann, G. E. G. 1996d: Ammonite life and habitat. In: Chilean Andes. Geobios 22: 553–597. Landman, N.; Tanabe, K.; Davis, C. ed. Ammonoid paleobiology. New York, Plenum. Pp. 607–707. Riccardi, A. C.; Gulisano, C. A.; Mojica, J.; Palacios, O.; Schubert, C.; Thomson, M. R. A. 1992: Western South America and Westermann, G. E. G. 2000: Marine faunal realms of the Mesozoic: Antarctica. In: Westermann, G. E. G. ed. The Jurassic of review and revision under the new guidelines for the Circum-Pacific. New York, Cambridge University biogeographic classification and nomenclature. Palaeo- Press. Pp. 123–161. geography, Palaeoclimatology, Palaeoecology 163: 49–68. Sandoval, J.; Westermann, G. E. G.; Marshall, M. C. 1990: Westermann, G. E. G. 2001: Modes of extinction, pseudo- Ammonite fauna, stratigraphy and ecology of Bathonian– extinction and distribution in Middle Jurassic ammonites: Callovian (Jurassic) Tecocoyunca Group, South Mexico. terminology. Canadian Journal of Earth Sciences 38: Palaeontographica A 210: 93–149. 187–195. Spath, L. F. 1928: Revision of Jurassic cephalopod fauna of Kachh Westermann, G. E. G.; Callomon, J. H. 1988: The Macro- (Cutch). Palaeontologica Indica New Series 9 (2). cephalitidae and associated Bathonian and early Callovian (Jurassic) Ammonitina of the Sula Islands and New Guinea. Stevens, G. R. 1968: The Jurassic System in New Zealand. New Palaeontographica A 203: 1–90. Zealand Geological Survey Report 35. Westermann, G. E. G.; Hudson, N. 1991: The first find of Stevens, G. R. 1978: Jurassic palaeontology. In: Suggate, R. P.; Eurycephalitinae (Jurassic Ammonitina) in New Zealand Stevens, G. R.; Te Punga, M. T. ed. The geology of New and its biogeographic significance. Journal of Paleon- Zealand. Wellington, Government Printer. Pp. 215–228. tology 65: 689–693. Stevens, G. R. 1997: The Late Jurassic ammonite fauna of New Westermann, G. E. G.; Riccardi, A. C. 1985: Middle Jurassic Zealand. Institute of Geological & Nuclear Sciences ammonite evolution in the Andean Province and emigration Monograph 18. 217 p. to Tethys. In: Bayer, U.; Seilacher, A. ed. Sedimentary and Stipanicic, P. N.; Westermann, G. E. G.; Riccardi, A. C. 1976: evolutionary cycles. Lecture Notes in Earth Sciences 7: The Indo-Pacific ammonite Mayaites in the Oxfordian of 6–34. the Southern Andes. Ameghiniana 12: 281–305. Westermann, G. E. G.; Hudson, N.; Grant-Mackie, J. A. 2000: Suggate, R. P.; Stevens, G. R.; Te Punga, M. T. ed. 1978: The Bajocian (Middle Jurassic) Ammonitina of New Zealand. geology of New Zealand. Wellington, Government Printer. New Zealand Journal of Geology and Geophysics 43: 820 p. 2 vol. 33–57.