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ABSTRACT
ABSTRACT
The systematic taxonomy, paleogeography, paleoecology, and paleoenvironment of the first macrofossil from the Caples Terrane in Northland and second Permian crinoid from New Zealand is described. Monobrachiocrinus waipapaensis n. sp. was collected as float material at the mouth of the Waipapa Gorge, Northland. Possible paleobiogeographic origins for the specimen are suggested and influence of the global distribution of Permian taxa pertinent to the New Zealand occurrence is discussed.
Introduction
In November 2016 Auckland Geology Club members toured Northland visiting examples of different outcropping rocks. Accessible Caples Terrane (Figure 1) rock types were observed in the partly-dry gravel bed of the Waipapa River adjacent to the reserve area at the end of Forest Road, north of Okaihau. Co-author, Liz Hoskin, discovered amongst gravel in the river bed, a small water-worn, rounded pebble containing fossilised crinoidal elements. The fossil locality (Figure 2) is sited at the mouth of the Waipapa River gorge where it emerges from the Omahuta-Puketi Range. Catchment of the Waipapa River drains from the Puketi Range, which is predominantly composed of late Paleozoic-early Mesozoic Caples Terrane basement, comprising a volcanic/greywacke suite, with minor Late Cretaceous-Cenozoic sedimentary and volcanic cover-rocks (Edbrooke and Brook 2009).
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15 August 2018Figure 1. Map of the Northland-Auckland region indicating the area of exposed Caples Terrane and adjacent Waipapa Terrane (representational only).
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Figure 1. Map of the Northland-Auckland region indicating the area of exposed Caples Terrane and adjacent Waipapa Terrane (representational only).
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15 August 2018Figure 2. Geological map of the vicinity of the Waipapa River crinoid fossil locality (red dot), showing the area of Caples Terrane in the Waipapa horst, comprising an eastern Puketi Forest block and western Omahuta Forest block. Late Cretaceous and Cenozoic cover rocks, mainly Late Eocene Te Kuiti Group, overlie parts of the horst and extend further north. Pliocene Kerikeri Volcanics are located east and Northland Allochthon rocks north-west (modified from Spörli et al. 2007).
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Figure 2. Geological map of the vicinity of the Waipapa River crinoid fossil locality (red dot), showing the area of Caples Terrane in the Waipapa horst, comprising an eastern Puketi Forest block and western Omahuta Forest block. Late Cretaceous and Cenozoic cover rocks, mainly Late Eocene Te Kuiti Group, overlie parts of the horst and extend further north. Pliocene Kerikeri Volcanics are located east and Northland Allochthon rocks north-west (modified from Spörli et al. 2007).
A confluence of the Waipapa River and Mangapa River in the middle of the range flows through the uplifted Omahuta-Puketi basement block from north to south. North of the uplifted Caples block are in-place Late Eocene sedimentary rocks of the Te Kuiti Group, overlain by various Cretaceous to Oligocene displaced rocks of the Northland Allochthon and isolated remnants of Pliocene basalt flows (Kerikeri Volcanic Group) (Brook et al. 1988). By comparison of rock types it is clear that only the Caples Terrane could be the source of the indurated crinoid-bearing pebble. No macrofossils have previously been recorded from the Caples Terrane in Northland (Adams and Maas 2004), but rare macrofossils of Permian to Jurassic age have been recorded from adjacent Waipapa Terrane (e.g. Hornibrook 1951; Spörli and Grant-Mackie 1976).
Caples Terrane Northland
Studies of the geology of the uplifted Omahuta-Puketi forest basement blocks (Waipapa Horst) (Figure 2), including geochemical, petrographic and detrital zircon studies (Jennings 1991; Black 1994; Adams et al. 2009, 2013) have shown that the rocks of the Waipapa horst differ significantly from the Waipapa Terrane basement rocks of east Northland and Auckland, and led to their assignment to the Caples Terrane. Subsequent Rb – Sr whole-rock isochron age (t) and initial
The Omahuta-Puketi rocks lie between the Junction Magnetic Anomaly (Dun Mountain Ophiolite Belt) to the west and Waipapa Terrane to the east – a tectonostratigraphic setting analogous to that of the Caples Terrane in the South Island, with which they have been correlated (Black 1994; Isaac 1996; Edbrooke and Brook 2009). The Waipapa horst in Northland is the northernmost outcrop of the Caples Terrane in New Zealand. Jennings (1989) mapped the area and divided it into a western sedimentary association (Omahuta unit) of interbedded, greywacke sandstone and argillite, with rare conglomerate and breccia horizons. The eastern Puketi unit consists of massive basalt, basalt pillow lava, red and green tuffs, siliceous argillite, chert, minor mudstone and sandstone – a dominantly ocean-floor basalt association with multicoloured chert and argillite (Jennings 1989; Edbrooke and Brook 2009). The Waipapa River gorge cuts through both Omahuta and Puketi units (Figure 2), therefore the crinoid fossil could have come from either. Waipapa horst rocks have been metamorphosed in the Jurassic to prehnite-pumpellyite and pumpellyite-actinolite facies (Black 1994), probably during deep burial associated with collision and suturing of the Caples Terrane to south-eastern Gondwana.
Age determination of the Caples Terrane in Northland has been hampered by the lack of previously recorded macrofossils. Cherts within the Puketi unit of the Caples Terrane have yielded poorly preserved radiolaria of late Paleozoic (Permian) to possibly early Mesozoic age (R. Hori in Black 1994). Studies of the ages of detrital zircons in one sedimentary rock sample from the Omahuta unit on Kauri Sanctuary Road showed a dominance of zircons of Late Triassic age (220–230 Ma.) and a few of Late Permian age (290 Ma.) (Adams et al. 2009). The younger zircons were inferred to be derived from contemporaneous volcanism and to indicate an age no older than Late Triassic for these rocks. Most Caples Terrane sedimentary rocks in New Zealand that have been dated by detrital zircons appear to be Triassic (Adams et al. 2009), but the Caples Terrane is still generally considered to range in age from Permian to Triassic (e.g. Mortimer and Campbell 2014; Mortimer et al. 2017).
Systematic Paleontology
Taxonomy mainly follows: Wanner (1924, 1926, 1940), Lane (1967), Moore and Teichert (1978), Webster (1987, 1990), and Simms and Sevastopulo (1993).
Diagnosis
Dicyclic inadunate crinoid having two circlets of plates below radials; anal sac [cone] prominent; theca small, typically globose or cone shaped; with one arm-bearing radial; other radials present or absent, commonly unequal and reduced in size; infrabasals three; basals five; anal X plate absent; anal opening bounded by basals and radials, or by radial; tegmen low, composed of five orals, posterior oral large, separating BC and DC orals; orals small, commonly not preserved; arms uniserial, isotomous, non-pinulate, rarely preserved; stem columnals circular (Moore et al. 1978).
Age range
Lower Carboniferous (Tournasian); Early Permian (Asselian) – Late Permian (Changhsingian).
Discussion
Five genera comprise the extinct family Sycocrinitidae: Sycocrinities Austin and Austin 1842; Allosycocrinus Wanner 1924, Metasycocrinus Wanner 1920; Monobrachiocrinus Wanner 1916; and Parasycocrinus Marez-Oyens 1940. Lane (1967) postulated that a natural phylogeny occurred within the Sycocrinitidae commencing with Sycocrinites in the Carboniferous (Mississippian (Tournasian)), giving rise to Metasycocrinus, then Parasycocrinus, and subsequently Monobrachiocrinus (all Permian). Sycocrinitidae possess a tegmen (disc) composed of 5 small tessellate plates or a membrane, flexible uniserial non-pinnulate arms that are free above radial plates comprised mostly of rounded to sub-rounded ossicles. A short, semi-flexible column comprised of round columnals supported a symmetric or asymmetric small theca. The family was sessile, being attached to a hard benthic substrate by a cemented holdfast, morphology unknown (e.g. Lane 1967; Moore et al. 1978; Ausich et al. 1999; Ausich and Webster 2008).
Type species
M. ficiformis Wanner 1916; by original description.
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15 August 2018Figure 3. Plate diagram of the main morphological features of a Monobrachiocrinus theca: infrabasals (A ray); basals (BC, CD, DE, AE); D radial (after Moore et al. 1978) (representational only).
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Figure 3. Plate diagram of the main morphological features of a Monobrachiocrinus theca: infrabasals (A ray); basals (BC, CD, DE, AE); D radial (after Moore et al. 1978) (representational only).
Type locality
Basleo beds, West Timor, South East Asia.
Diagnosis
Dicyclic inadunate crinoid; cup inverted pear-shape; infrabasals 3, forming conspicuous part of cup, small infrabasal in A ray; basals 5, higher than wide; D radial enlarged, rounding off top of DE basal and limited laterally by CD and AE basals, other radials absent; arms non-pinnulate; anal opening bounded by CD and BC basals; orals 5, small, CD one largest; arms non-pinnulate (Wanner 1916; Moore et al. 1978).
Age range
Early Permian (Artinskian) – Middle Permian (Wordian).
Distribution
Europe (Russia (Krasnoufimsk)); Italy (Sicily); West Timor (Basleo region), New Zealand (Northland, North Island – this paper).
Discussion
Monobrachiocrinus is a small sedentary crinoid possessing morphological characters fundamental in function and form, and differentiated from other genera in the family by the number of theca radials. A single non-pinnulate arm emanating from a single radial plate characterises the cup.
Monobrachiocrinus waipapaensis Eagle n. sp. (Figures 4–7).
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15 August 2018Figure 4. A. Monobrachiocrinus waipapaensis; B. diagram showing: infrabasals (including A-ray), parts of basals, and oral plates (scale in mm).
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Figure 4. A. Monobrachiocrinus waipapaensis; B. diagram showing: infrabasals (including A-ray), parts of basals, and oral plates (scale in mm).
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15 August 2018Figure 5. A. Monobrachiocrinus waipapaensis: B. diagram showing: infrabasals, parts of basals, and oral plates (scale in mm.).
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Figure 5. A. Monobrachiocrinus waipapaensis: B. diagram showing: infrabasals, parts of basals, and oral plates (scale in mm.).
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15 August 2018Figure 6. A. Monobrachiocrinius waipapaensis: B. diagram showing: anal cone, arm ossicles, parts of basals, and D radial (scale in mm.).
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Figure 6. A. Monobrachiocrinius waipapaensis: B. diagram showing: anal cone, arm ossicles, parts of basals, and D radial (scale in mm.).
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15 August 2018Figure 7. A. Monobrachiocrinus waipapaensis; B. diagram showing: arm ossicles, D radial, parts of basals, and anal cone (scale in mm.).
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Figure 7. A. Monobrachiocrinus waipapaensis; B. diagram showing: arm ossicles, D radial, parts of basals, and anal cone (scale in mm.).
Material
Holotype: crinoidal skeletal elements preserved within a water-worn, rounded pebble (meristics at extremities in mm: 30 long, 24 wide, and 13 diameter) comprised of pale green indurated volcarenite matrix. Specimen type catalogue number: EC 1174 lodged: National Palaeontology Collections, GNS Science.
Type locality
Waipapa River, at the mouth of Waipapa Gorge, adjacent to picnic area (reserve), at end of Forest Road, Puketi, Northland; GNS Science fossil record number: P05/f0298; New Zealand map grid ref.: P05/2572947 6658090; NZGD49: 35.27884˚S 173.68298˚E. Float material collected by Liz Hoskin from Waipapa River bed gravels clearly sourced from Caples Terrane rocks upstream, on November 25, 2016.
Etymology
Named for the Waipapa River draining through the Waipapa Horst.
New Zealand distribution
Known only from the type locality.
New Zealand age
Being ‘float’ material the specimen lacks biostratigraphic zonation, precluding the species being an ecostratigraphic marker providing geological and chronologic evidence. Additionally, the Permian genus is not diagnostic of any particular age, so that an attempt at assigning an age to Monobrachiocrinus waipapaensis n. sp. is speculative. Supporting evidence of Late Permian radiolaria (R. Hori in Black 1994) and detrital zircons of Late Permian age (Adams et al. 2009) from the Puketi unit infer a Late Permian (Wuchiapingian or Changhsingian) age. However, distribution and evolutionary traits based on the occurrence of Monobrachiocrinus species suggest an Early to Middle Permian (Artinskian to Wordian) age.
Diagnosis
A dicyclic inadunate cladid of the Cyathocrinina with inverted pear-shaped infrabasals, five anal plates in cup, possessing a single D radial narrower than width; tegmen with massive, short, imperforate non-porous anal cone opening at distal end; anal opening bounded by basals and radial; oral plates prominent, posterior plate imperforate, commonly with hydropore; arms narrowly rounded, uniserial, non-pinnulate; stem transversely round, with large axial canal; columnal stem supporting cup anchored by a cemented holdfast.
Description
Crushed and partially distorted dicyclic inadunate crinoid; theca bowl-shaped, with 3 elongate, inverted pear-shaped infrabasals, 2 large, one small, forming A ray as adoral part of calyx; 5 adoral, broad, robust basal plates higher than wide; single D radial plate narrower than width aboral of calyx supporting a disarticulated, autonomous, single, unbranched, narrowly rounded uniserial, non-pinnulate arm, coiling capable; arm ossicles circular proximally to sub-circular distally, with small, nerve axial canal central; arm ossicle articulation synostosial, graduated in size proximal to distal extremities; 5 prominent, small, tessellate oral plates, posterior plate larger with hydropore; X plate absent; a short, imperforate, non-porous anal cone prominent bounded by basals and radial; small terminal anal opening distal end of cone; stem ossicles absent, presumed transversely round due to infrabasal dorsal termination; cemented holdfast absent; skeletal meristics mm: proximal arm ossicle maximum diameter 2.8; distal arm ossicle maximum diameter 0.7; largest infrabasal length 14.8; largest basal length 10.3; largest oral plate 0.6; D radial plate diameter 7.8; anal sac diameter 6.0.
Discussion
There are seven Permian species or subspecies of Monobrachiocrinus known globally: Monobrachiocrinus siciliensis Yakovlev 1930, p. 909, pl. 1, fig. 2; 1934, p. 275, pl. 19, fig. 15, from the Sosio Limestone, Sicily, Italy, Middle Permian (Wordian); M. waitzi Marez-Oyens 1940, p. 324, pl. 3, fig. 10, text- fig. 7 from the Besleo [Basleo] beds of Basleo, and from Noko (between Basleo and Noil Toke), West Timor, Early Permian (Artinskian); M. oviformis Yakovlev 1926, p. 57, text-fig. 1; 1930, p. 102, pl. 1, figs. 2a,b, from Mt. Divia, near Krasnoufimsk, southern Ural Mountains, Russia, Early Permian (Late Artinskian); M. ficiformis ficiformis Wanner 1916, p. 104, pl.102, figs. 6–9, text- fig. 28 in greenish tuff from the Besleo beds, Basleo, Noil Tonini (Toeninoe), Noil Fatoe, Koeka near Baung, Landschaft Amarassi, between Niki Niki and Noil Fatoe, Kiumoko, and Toenioen Eno, West Timor, Early Permian (Artinskian); M. ficiformis carinatus Wanner 1929, p. 62, pl. 3, figs. 11–12 from the Besleo beds, Basleo, West Timor, Early Permian (Artinskian); M. ficiformis elongatus Wanner 1916: pl. 108, pl. 102, fig. 10 from the Besleo beds, Basleo, and Dorf Sebot, West Timor, Early Permain (Artinskian); M. ficiformis granulatus Wanner 1924, p. 51, pl. 8, figs. 13–15 from the Besleo beds, Basleo, and Nefotassi near Soefa, Landschaft Beboki, West Timor, Early Permian (Artinskian). Initial interpretations of a Late (now Middle) Permian range (Wordian) for the West Timor Monobrachiocrinus species by Wanner and others, and a Guadalupian (Capitanian) age by Erwin (1993), was corrected by Webster (1998a): 45, as Early Permian (Artinskian). Reconstructed New Zealand Monobrachiocrinus waipapaensis (Figure 8) is of robust construction and mature size. Of the aforementioned Monobrachiocrinus species, the most comparable in morphological character to M. waipapaensis are M. f. ficiformis (Figure 9(A)) and M. f. granulatus (Figure 9(B)). The inverted pear-shaped infrabasals in M. f. ficiformis and M. f. granulatus possess a similar configuration to M. waipapaensis but are not as narrowly elongate; those of M. f. ficiformis are not as flared distally, but those in M. f. granulatus are. However, infrabasals in M. f. ficiformis are long, but not as long as in M. waipapaensis, whereas those of M. f. granulatus are much shorter in length and appear as reduced distally where columnal ossicles join the cup. Basals are similarly robust and broad in both M. f. ficiformis and M. f. granulatus, but those of M. f. ficiformis are longer than those of M. waipapaensis and M. f. granulatus which appear of similar length; all three species possess basal plates slightly inwardly curved at sutures. The D radial in M. waipapaensis appears to be of the same configuration as M. f. ficiformis, but is larger than that of M. f. ficiformis or M. f. granulatus as are the 5 oral plates. However this observation may also be due to compression and lateral expansion due to diagenesis. The five oral plates are of a similar configuration as those possessed by M. f. ficiformis and M. f. granulatus, with the CD oral plate appearing to be the largest, normally separated from the CD basal but presently by being splayed-out in series by compression. Oral plates in M. waipapaensis are naturally slightly larger than those of both comparative species, but probably more so because of flattened extension. M. waipapaensis, has a larger anal cone than M. f. ficiformis and possibly M. f. granulatus (no example of that species could be found to compare; the anal cavity seen in several specimens suggests this). Arm ossicles of M. waipapaensis are presumed to be of a similar shape and size as M. f. ficiformis and M. f. granulatus, as no examples could be obtained for comparison, either materially or published; arms (or elements thereof) of the Codiacrinoidea are rarely preserved, as are oral plates (Bather 1890; Lane 1967; Moore et al. 1978). The D radial in M. waipapaensis is larger than that of M. f. granulatus as are the 5 oral plates, forgiving compression and/or lateral expansion. Irrespective of the skeletal elements of M. waipapaensis having been crushed, partially distorted and broken, a number separated, and with some skeletal elements overlaid – sufficient diagnostic morphology exists to support differentiation from that of other Monobrachiocrinus species.
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15 August 2018Figure 8. A reconstruction of sessile, non-pinnulate Monobrachiocrinus waipapaensis in life situ showing skeletal morphology and cemented holdfast (representational only).
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Figure 8. A reconstruction of sessile, non-pinnulate Monobrachiocrinus waipapaensis in life situ showing skeletal morphology and cemented holdfast (representational only).
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15 August 2018Figure 9. Graphic illustrations of Monobrachiocrinus species comparable to M. waipapaensis; both species are Early Permian (Artinskian) from the Besleo beds of Basleo, Timor: A. M. ficiformis ficiformis dorsal, ventral CD-interray, and aboral views of theca (dorsal and ventral redrawn from Wanner 1916; scale: X 1.5); B. M. ficiformis granulatus (dorsal, ventral BC-interray, and aboral views of theca (redrawn from Wanner 1924; scale: X 1.5).
![]({"type":"image","src":"/na101/home/literatum/publisher/tandf/journals/content/tnzg20/2018/tnzg20.v061.i04/00288306.2018.1506484/20181008/images/medium/tnzg_a_1506484_f0009_ob.jpg"})
Figure 9. Graphic illustrations of Monobrachiocrinus species comparable to M. waipapaensis; both species are Early Permian (Artinskian) from the Besleo beds of Basleo, Timor: A. M. ficiformis ficiformis dorsal, ventral CD-interray, and aboral views of theca (dorsal and ventral redrawn from Wanner 1916; scale: X 1.5); B. M. ficiformis granulatus (dorsal, ventral BC-interray, and aboral views of theca (redrawn from Wanner 1924; scale: X 1.5).
Monobrachiocrinus waipapaensis is the second Permian crinoid known from New Zealand, and possibly the second cladid specimen. Waterhouse and Vella (1965) described a Permian fauna from Parapara Peak, North-West Nelson, New Zealand. The faunal list comprised mostly of brachiopods and molluscs occurring in the clastic Parapara Group, Cross Member, but included a paragraph on the first possible cladid crinoid to be recorded from New Zealand as follows:
Tribrachiocrinus? sp. Several dissociated basal and infrabasal plates and fragments of rounded columnals could belong to this genus. One basal plate has an ornament of tubercles like that of T. corrugatus Ratte 1885. Part of the arms of one specimen is seen on a mould [Victoria University, Wellington collection number: V1032]. (Waterhouse and Vella 1965, p. 79)
Paleobiogeography, Paleoecology and Paleoenvironment
The dominant Late Triassic detrital zircons in the Omahuta unit sample have been inferred to be derived from contemporaneous Caples volcanism along the plate boundary at the edge of Gondwana (Adams et al. 2009). Geochemistry of the Puketi unit basalts shows they are part of a tholeiitic oceanic andesite association, quite distinct from basalts in other basement rocks in northern New Zealand and are inferred to have accumulated in an oceanic rift environment similar to the present day Galapagos Rift (Jennings 1991; Black 1994).
Global evidence suggests that the Crinoidea sustained a catastrophic decline in the Late Carboniferous–Permian prior to the Permian–Triassic extinction event (Wanner 1937, 1940; Moore and Teichert 1978; Hess et al. 1999), compared with older Paleozoic periods, when they were abundant both in species diversity and population numbers. This is certainly true for New Zealand where few Carboniferous rocks are known and where no crinoidal Carboniferous or Early Permian rock has been recorded. Interestingly, Monobrachiocrinus is unknown from Australia (see: Webster 1998b). Though Late Permian species exist globally and are extremely rare, the genera that they are associated with are often long-ranging. The Permian was a critical time in crinoid evolution as the end-Permian extinction event (251 Ma.) eradicating almost all Paleozoic crinoid groups including the Cladida, Disparida, Flexibilia, and Camerata (Moore et al. 1978; Hess et al. 1999).
Assemblages of Permian crinoids globally are rare as are isolated specimens. The Northland, New Zealand Monobraciocrinus specimen, though only a portion of the skeletal structure and deformed, is a rarity. It is suggested that the specimen’s skeletal disarticulation is nominal and not caused by post mortem transportation or predation because of its state of preservation; proximity of theca elements and arm ossicles are associated and local. It appears that skeletal disarticulation was due to natural disintegration after tissue decay and the skeletal elements were quickly covered by sediment ensuring preservation; continued sediment accumulation led to deep burial and compression. No pinnules are found in association with Monobrachiocrinus waipapaensis, suggesting that the genus is non-pinnulate. Being mainly benthic and facies-bound, sessile crinoids like Monobrachiocrinus are not particularly suited as index fossils. However, their phylogenetic history, mostly producing short-lived species with rapid morphological change makes them invaluable for biostratigraphic zonation and biogeographic migratory correlations (Eagle 2014). Additionally, benthic crinoids are excellent ecostratigraphic markers providing geological and chronologic evidence of certain habitats, biotopes, or facies types when found in-situ (Eagle 2014). Elapsed time from the first reported occurrence of Monobrachiocrinus (Artinskian (284.4 Ma.)), to the last recorded occurrence (Wordian (265.8 Ma.)) equates to an 18.6 Ma. period, plenty of time to migrate and evolve different species in different places.
Throughout the Late Permian a regional migration of sessile benthic faunas (among them crinoids) following changing climate patterns, and water column conditions, colonised suitable substrates in warm water environments (Dickens 1961, 1973, 1978, 1985; Archbold 1987, 1991; Yeates et al. 1987; Meyerhoff et al. 1996; Hess et al. 1999) off shore from Gondwana. Mid-ocean basalts that erupted at a spreading centre or rift could have built a seamount that would have provided hard rock for settlement by M. waipapaensis.
Discussion
The limited age information suggests that the Omahuta unit may be younger (Late Triassic) than the Puketi unit (Late Permian) and if this is correct then the Permian crinoid most likely was derived from the older Puketi unit. This age dichotomy is similar to the situation in the adjacent Waipapa Terrane of northern Northland, where the ocean-floor basalts and associated chert is Late Permian-Early Triassic in age and the surrounding terrigeneous greywackes and argillites are Middle-Late Triassic (Spörli et al. 2007). Both situations are suggestive of eruption of the basalts at mid-ocean spreading centres or rifts with the slow accumulation of oceanic sediment (chert, multi-coloured argillite) on top of them as they were rafted along by seafloor spreading towards the margin of Gondwana and a proto-Zealandia. As these rocks came close to the eroding periphery of Gondwana in the Late Triassic, terrigeneous sediment (Omahuta unit) accumulated on top of them before they collided with and were sutured onto the margin of the supercontinent. Therefore, the possibility of a Permian-Triassic boundary exists within the Waipapa Horst, Caples Terrane rocks.
In the adjacent Waipapa Terrane all the chert, multi-coloured argillite and terrigeneous greywacke and argillite are accepted as accumulating at considerable water depths – bathyal or abyssal (probably >500 m water depth). The one known anomaly is that the Marble Bay pillow basalt is associated with carbonate sediment containing fossil reef corals and large fusulinid foraminifera (Hornibrook 1951). These two fossil groups are believed to have hosted commensal photobionts (photosynthesising algae) like their modern counterparts, and therefore would have lived in the photic zone at water depths no greater than 100 m. Thus we infer that some of the ocean rift basalts in the Northland Caples Terrane erupted to form seamounts that rose to shallow depths or even formed islands (like the Galapagos, Azores or Iceland do today). A similar setting may explain the presence of M. waipapaensis in with the basalt lava flows of the Puketi unit (although marble or limestone have not been recorded from the Omahuta-Puketi Range), and was likely living in 100–200 m-deep water, tens to thousands of kilometres away from Gondwana in the Panthalassan Ocean. This environmental situation may explain why the new species is different from those found elsewhere.
The Caples Terrane has also been recognised geochemically and petrographically from other Northland and Auckland localities at the western end of a number of basement greywacke blocks west of Whangarei, at Taipuha, Waipu Gorge and Pokeno (Adams et al. 2009). No obvious major terrane boundary or fault has been mapped or hypothesised separating these localities from Waipapa Terrane. Now with the similarity in age range and lithologic associations of the Caples and Waipapa Terranes (e.g. Adams et al. 2017), one might ask whether the two terranes in Northland may not be combined, with the Triassic Caples terrigeneous association being deposited more proximal to volcanism on the margin of Gondwana than the Waipapa terrigeneous sediments were. An alternative explanation that cannot be fully discounted is that the Permian-aged crinoid is contained within a pebble that has recently eroded out of a conglomerate bed within the Triassic Caples Terrane and thus the crinoid does not date the Caples but was reworked from Permian rocks into a Caples Terrane conglomerate during the Triassic. We favour the former, simpler explanation but further field work is needed to test these competing hypotheses.
| PHYLUM | ECHINODERMATA | Klein 1754 |
| CLASS | CRINOIDEA | Miller 1821 |
| SUBCLASS | INADUNATA | Wachsmuth and Springer 1885 |
| ORDER | CLADIDA | Moore and Laudon 1943 |
| SUBORDER | CYATHOCRININA | Bather 1899 |
| SUPERFAMILY | CODIACRINOIDEA | Bather 1890 |
| SUBFAMILY | THETIDICRININAE | Wanner 1930 |
| FAMILY | SYCOCRINITIDAE | Lane 1967, p. 12 |
Related Research Data
Acknowledgements
We thank an anonymous reviewer and Steve Edbrooke for their helpful comments and suggestions and Gary Webster for reviewing the identification and taxonomy of the Permian crinoid, all of which greatly improved the final MS.
Disclosure statement
No potential conflict of interest was reported by the author.
ORCID
Michael K. Eagle http://orcid.org/0000-0002-5569-3057
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