Journal of the Royal Society of New Zealand Early ...

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Early Miocene flora of the Manuherikia Group, New Zealand. 6. Lauraceae Mike Pole

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Department of Geology , University of Otago , P.O. Box 56, Dunedin , New Zealand Published online: 14 Jun 2013.

To cite this article: Mike Pole (1993) Early Miocene flora of the Manuherikia Group, New Zealand. 6. Lauraceae, Journal of the Royal Society of New Zealand, 23:4, 303-312, DOI: 10.1080/03036758.1993.10721228 To link to this article: http://dx.doi.org/10.1080/03036758.1993.10721228

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©Journal of The Royal Society of New Zealand, Volume 23. Number4. December 1993, pp 303-312

Early Miocene flora of the Manuherikia Group, New Zealand. 6. Lauraceae

Downloaded by [121.75.216.62] at 17:59 13 October 2014

Mike Pole*

Three species of Lauraceae are described from Miocene sediments of the Manuherikia Group, Central Otago, New Zealand. One species is mummified, with anatomical detail, and is placed into the organ-genus Laurophyllum. It is regarded as the same taxon as Cryptocarya longfordiensis Holden, which is removed to Laurophyllum in the absence (for now) of a means to distinguish it from some other Lauraceae genera. The other two species are impressions only, and are regarded as Lauraceae on architectural detail. Keywords: Central Otago, Cryptocarya, Laurophyllum, paleobotany

INTRODUCTION The plant-bearing Manuherikia Group was deposited during the Miocene as a complex of fluvial, lacustrine, and associated facies on top of a deeply weathered, and possibly peneplained, schist basement. Douglas (1986) has produced a detailed analysis of Manuherikia Group sedimentology, stratigraphy and paleogeography. Palynological evidence (Mildenhall, 1989) does not constrain the age of the plant fossils in the lower Manuherikia Group closer than Early to Middle Miocene (Otaian to Lillbumian local stages). The leaves are preserved as impressions, or as mummified compressions, in fine silt and carbonaceous mud. Illustration technique follows Pole ( 1992a).

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Fig. l - Locality map. *Department of Geology, University of Otago, P.O. Box 56, Dunedin, New Zealand. Present address: Department of Plant Science, University of Tasmania, G.P.O. Box 252C, Hobart 7001, Australia.

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All angiosperm leaf taxa described in this study have been allocated a non-Linnean parataxon codename (the prefix MANU- followed by a number), in addition to any Linnean system name which may have been applied. Such a system is useful with fossil leaf material, where the affinities of most specimens are unknown and where it is advantageous to be able to discuss particular taxa without formally identifying them or setting up a form generic scheme. They are essentially species without a genus. In lieu of a specific diagnosis, a combination of architectural characters distinguishing the parataxon from other Manuherikia Group taxa is given. The parataxon equivalent of a holotype is a nominated reference specimen. Leaf architecture terminology follows Hickey ( 1973, 1979) and Pole (1991 ). Specific fossil localities (Fig. 1) are referred to by the registered N.Z. Fossil Record File locality number, based on the metric N.Z.M.S. sheets. All fossil material is catalogued and stored in the Department of Geology, University of Otago. Herbarium material is stored in the Department of Plant Science, University of Tasmania. BOTANICAL AFFINITY Leaf impressions of this general type have long been referred to the Lauraceae with very little supporting evidence. Generic placing is extremely tenuous, and the practice has been discarded by some paleobotanists in favour of recognising organ genera. Hill ( 1986), summarising the work of Wolfe (1977), Krausel and Weyland (1950), and Sturm (1971), advocates the use of the organ genus Laurophyllum Goeppert 1853 for all entire-margined fossil leaves with anatomical detail confirming placement in the Lauraceae. Leaf impressions are here placed in Lauraceae on the basis of having of entire margins, externodromous or eucamptodromous venation, and strong percurrent venation. Furthermore, small, closed, isodiametric areoles having a width of only about four times the thickness of the enclosing veins are characteristic of some Lauraceae. Mummified leaves are identified as Lauraceae on the basis of the epidermal characters listed by Hill (1986), i.e., hypostomatic, paracytic stomata with embedded guard cells and cuticular scales. I agree with workers previous to Hill on the difficulty or naivety of placing leaf remains (with or without cuticle) into lauracean genera. Frequently, venation patterns seem to cut across currently-accepted taxonomic boundaries. This may indicate a taxonomic problem, or that further definition is needed within the venation terminology. However it is my contention that full anatomical detail may not always be necessary for placement at family, or even generic level. The combination of entire-margin, externodromous or eucamptodromous first order pattern, with a strong percurrent second order pattern, is distinct in existing rainforests of Australasia and Papua New Guinea and appears to be limited to the Lauraceae. Holden (1982) described four species of Cryptocarya from the Mid to Upper Miocene Longford Formation, New Zealand. The remains she described were fragmentary and lacked cuticle; however I agree with her placement of these species in the family Lauraceae. (C. longfordiensis, C. murchisoniensis, and C. bulleriana). Holden's four species appear to be separated only on the character of lamina width and, comparing them with normal variation within similar living Cryptocarya species and populations of MANU-1 in the Manuherikia Group, I regard them as belonging to a single species. I feel that while Holden's species are quite likely in Cryptocarya (and probably closest to the acrodromous C. triplinervis R. Br. (Fig. 7B) as she suggested), this cannot yet be affirmed, and placement in say, Cinnamomum or Neolitsea ruled out. Additionally I believe that this species is identical to the Manuherikia Group MANU-I. It is unsatisfactory that well-preserved specimens are placed into a species based on poor type material, but I justify this decision in that it reduces rather than increases the number of species. As the cuticular details of MANU-1 confirm placement in the Lauraceae, I suggest the non-commital genus Laurophyllum Goeppert would be appropriate at this stage. Using priority of page-order, this is Laurophyllum longfordiensis. At present fourteen well described species of Laurophyllum (sensu Hill) exist in Australia (Hill, 1986, 1988; Pole, 1992b). 66

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A key to the Australian genera of Lauraceae based on cuticular characters has been developed (A. Rowett and D. Christophel, pers. comm. 1992) and it would be prudent to retain the specimens in Laurophyllum until this key is published. MANU-2 is perhaps only a variant of MANU-I, its only difference being the notched apex. MANU-3 has never been recovered with anatomical detail. It is considered as Lauraceae because of its close similarity in shape and in strong and regular percurrent second order venation with Cryptocarya species found in New Caledonia, C. macrophylla Guill. (Fig. 7A). Identification to generic level is not attempted but Cryptocarya is considered likely. I have seen patches of forest in New Caledonia where the leaf litter is dominated by the large, thick, and strongly veined leaves of this genus.


SYSTEMATICS Family LAURACEAE Laurophyllum Goeppert 1853 Laurophyllum longfordiensis (Holden) Pole (Parataxon MANU-I) Figs 2, 3, 6. Cryptocarya longfordiensis Holden 1982 Cryptocarya murchisoniensis Holden 1982 Cryptocarya bulleriana Holden 1982 Diagnosis Differs from all other Australasian Laurophyllum in having basal laterals which are almost paired, either at the base of the lamina, or just above, and cuticular scales which are nearly as long as the stomata. Parataxon Reference specimen OU30306 H4llf053 St Bathans Occurrence and referred specimens F4llf218 OU12646 F4llf222 OU 12339 F4llf227 OU12207 OU12179 0Ul2189 0Ul2178 0Ul2184 OU12190 OU12191 0Ul2197 OU12199 OU12193 0Ul2220 0Ul2221 0Ul2226 0Ul2227 OU12221 0Ul2228 OU12239 OU12246 0Ul2241 0Ul2248 OU12432 F41/f235 OU30031 F4llf237 OU29022 OU29023 OU29024 OU29025 OU29026 OU29027 OU29029 OU29030 OU29033 OU29034 OU29036 OU29037 H4llf053 OU30306 Description Size: length 41-90 mm, width 21-46 mm, area 7.2 cm 2 (OU29033)- 26.3 cm2 (OU29025). Microphyll =4 (80%): Notophyll = 1 (20%). Shape: wide-elliptic or ovate-elliptic, apex obtuse to acute, base acute-normal to obtusenormal, base sometimes asymmetrical. Petiole normal, length 6-7 mm. Margin entire. Development greatest towards base, acrodromous. Venation: first order venation pattern eucamptodromous. Midrib moderate, course straight to slightly curved. Basal first order laterals paired just above base, although one vein may be more decurrent along midrib, appearing sub opposite, strongly developed and curve up for

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Fig. 2- Laurophyllum longfordiensis (MANU-I), (a) OU29029, F41/f237, (b) OU29600, F41/f245, (c) OU29025, F41/f237, (d) OU29023, F41/f237, (e) OU29023, F41/f237, (t) OU29033, F41/f237.

Fig. 3- Laurophyllum longfordiensis (MANU-I), (a-b) Enlargement and whole leaf (OU30306), (c) Lower epidermis (0U30306, transmitted light), (d) Group of stomata (OU30306, transmitted light), (e) Single stomata (OU30306, SEM), (t) Lower epidermis, hair bases visible, stomata not visible (OU30306, SEM).

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em Fig. 4 - (a) Lauraceae sp. A (MANU-2), OU29167, F41/f235, (b) Laurophyllum longfordiensis MANU-I, OU 29574, F41/f227, (c) L. longfordiensis (MANU-I) enlargement of OU 29574.

about two thirds of lamina length before losing identity. Four or fewer less strongly developed first order laterals present in distal half of lamina, not decurrent on midrib, not parallel, angle of divergence moderate to wide acute, apical first order laterals diverge more widely than basal ones. External veins present as external loops. Second order venation pattern percurrent. Angle of origin of joining and interangular veins RR, approximately at right angles to midrib, arrangement alternate, simple and forked. Basal lateral segments have 17-18 interangular veins and only 3-4 joining veins, higher segments may have only 2-3 interangular veins and no joining veins. Third order venation pattern unresolved. Marginal ultimate venation looped; in lower portion of lamina loops flush with margins, forming sort of fimbrial vein. Areoles well developed, random, generally isodiametric, quadrangular to pentagonal, size small, width frequently only around four times thickness of enclosing veins. Veinlets not observed. Lower epidermis: Stomatal complex hypostomatic, paracytic stomata with embedded guard cells and cuticular scales,13-17 ~m long. Non-venous cells irregular, with variable number of sides, straight or curved walls, generally larger than stomatal complex, 13-20 ~m breadth. Venous cells markedly longer than non-venous cells, typically 10 x 25 ~m. Stomata oriented at random. Trichome bases with small, circular, thickened foot cell. Trichomes not preserved. Upper epidermis: cells isometrical, 12.5-22.5 ~m breadth, stomata absent, venous cells not distinguished. Lauraceae sp. A (Parataxon MANU-2) Fig. 4 Distinguishing features Differs from all other Manuherikia Group taxa in having acrodromous development with a retuse or emarginate apex.

Reference specimen OU29167, F41/f235 Bannockburn Occurrence and referred specimens F41/f235 OU13186 OU29167 Description Size: length 32 mm, width 26 mm, area 5.6 cm 2 (0029167). Microphyll = 2 (100%). Shape: wide ovate, apex retuse or emarginate, base obtuse-normal, or rounded, balance of 70


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base slightly asymmetrical. Petiole normal, length 2 mm (OU 29167). Margin entire. Development greatest towards base, acrodromous, basal perfect. Venation: first order venation pattern eucamptodromous. Midrib moderate, straight. One pair of strongly developed first order laterals paired at base of lamina, and one, or possibly two more in distal half of lamina, but on one side only of midrib, not decurrent on midrib, unevenly spaced; angle of divergence moderate acute, basal first order laterals as thick as midrib, others normal. Basal first order laterals curve up all way to emarginate apex where they apparently curve back down to unite with tip of midrib. Other first order laterals curve out from midrib to unite with basal first order laterals. External veins present as external loops. Second order venation pattern percurrent. Largest basal lateral segment consists entirely of interangular veins, while smaller one dominated by interangular veins but has two joining veins. More apical segments have only interangular veins. Angle of origin RR, strong, relationship to midrib mostly right-angles, arrangement alternate, simple and branched. Third order venation pattern unresolved. Marginal ultimate venation fimbrial. Areoles well developed, pattern unresolved, quadrangular to pentagonal, size moderate to small, veinlets possibly present, but not clear. Lauraceae sp. B (Parataxon MANU-3) Figs 5, 6. Distinguishing features

Differs from all other Manuherikia Group taxa in having regular and strong percurrent second order venation pattern. Reference specimen 0029163, F4l/f235 Bannockburn Occurrence and referred specimens F4l/f214 0012983 F41/f218 OU12451 0012452 OU12464 F41/f227 OU12235 F4l/f235 0Ul3177 0Ul3603 0013614 0013615 OU13618 0013619 0013630 0013631 0013632 0013633 0Ul3635 OU13638 0013639 0013640 0013643 OU 13644 0Ul3645 OU 13656 OU 13659 0Ul3668 OU13754 0013758 0029163 0029405 F4l/f236 0013224 OU13225 OU13227 0013228 0013231 0029505 F4l/f241 0029218 0029219 0029220 0029221 0029222 0029224 F42/f006 0029299 Description Size: length 26-120 mm, width 18-64 mm, area 18.9 cm 2 (0029163)- 52.3 em (0Ul3659). Microphyll = I (33% ): Notophyll = I (33% ): Mesophyll = 1 (33% ). Shape: ovate to narrow ovate, apex acuminate, base normal acute, balance of base slightly asymmetrical. Petiole normal, length 5 mm (OU 29163). Margin entire. Development more or less even over whole lamina except for unexpanded apex. Usual peak of development in mid-lamina. Venation: first order venation pattern variable, externodromous and eucamptodromous. Midrib moderate, straight to zig-zag. First order lateral veins not decurrent on midrib, unevenly spaced, alternate, non parallel, angle of divergence moderate acute, relatively thick, strongly curved, basal laterals not paired. External veins present as external loops. Second order venation pattern percurrent. Angle of origin of joining and interangular veins RR,

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...E

a

c Fig. 5 - Lauraceae sp. B (MANU-3), F41/f235, (a) OU29405, (b) OUI3177, (c) OUI3659, (d) OU29163.

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Lauraceae sp. B {MANU-3)

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lamina width/mm lamina width/mm Fig. 6- Lamina length versus width scatter-diagrams for Laurophyllum longfordiensis and MANU-3. Lines approximating the standard leaf size classes of Webb (1959) derived using Area =0.667 x length x width.

Fig. 7 - Extant Lauraceae for comparison with the fossils: (A) Cryptocarya macrophylla, OPH 1384, New Caledonia, (B) Cryptocarya triplinervis, D.L. Jones 186, Lord Howe Island. Scale in centimeters.

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relatively thick and regular, straight between first order laterals but convex when passing between lateral and midrib, sometimes branched, relationship to midrib varies from right angles to oblique, arrangement alternate. Third order venation pattern unresolved. Marginal ultimate venation looped. Areoles not known. CONCLUSIONS Three species of Lauraceae are now known from the Miocene of the Manuherikia Group and this number is likely to grow with future research. Due to the poor preservation potential of Lauraceae pollen, this family is generally absent from the palynological record. Its presence in the Manuherikia Group is detectable only from macrofossils, and, like other Tertiary localities worldwide (Hill 1986), it appears to have been an important element.


ACKNOWLEDGEMENTS I thank my supervisor, J.D. Campbell, and also R.S. Hill, J.D. Lovis, M.E. Dettmann and D.C. Christophel, who reviewed the manuscript at various stages, and the Patterson family of Bannockburn who allowed access on their property. I am grateful to B.P.M. Hyland who supplied several specimens of Cryptocarya triplinervis. This work was financed by a UGC scholarship and University of Otago Bridging Finance.

REFERENCES Douglas, B.J., 1986. Lignite resources of Central Otago. New Zealand Energy Research and Development Committee. Publication Pl04. Hickey, L.J., 1973. Classification of the architecture of dicotyledonous leaves. American Journal of Botany 60: 17-33. Hickey, L.J., 1979. A revised classification of the architecture of dicotyledonous leaves. in Metcalfe C.R. and Chalk L. (Eds): Anatomy of the Dicotyledons. Vol. I. second edition, pp. 25-39. Clarendon Press, Oxford. Hill, R.S., 1986. Lauraceous leaves from the Eocene of Nerriga, New South Wales. Alcheringa I0: 327351. Hill, R.S., 1988. A re-investigation of Nothofagus muelleri (Ett.) Paterson and Cinnamomum nuytsii Ett. from the Late Eocene of Vegetable Creek. Alcheringa 12: 221-231. Holden, A.M., 1982. Fossil Lauraceae and Proteaceae from the Longford Formation, Murchison, New Zealand. Journal of the Royal Society of N.Z. 12: 79-80. Hyland, B.P.M., 1982. A revised card key to rainforest trees of north Queensland. CSIRO, Melbourne. Hyland, B.P.M., 1989. A revision ofLauraceae in Australia (excluding Cassytha). Australian Systematic Botany 2: 135 -367. Klucking, E.P., 1987. Leaf venation patterns. Volume 2, Lauraceae. J. Cramer, Stuttgart. Kostermans, A.J.G.H., 1974. Flore de Ia Nouvelle-Caledonie et Dependances. 5. Lauracees. Museum National D'Histoire Naturelle, Paris. Krause], R., and Weyland, H., 1950. Kritische Untersuchungen zur Kutikularanalyse tertiare Blatter. I. Palaeontographica B 91: 7-92. Mildenhall, D.C., 1989. Summary of the age and paleoecology of the Miocene Manuherikia Group, Central Otago, New Zealand. Journal of the Royal Society of N.Z. 19: 19-29. Pole, M. S., 1991. A modified terminology for angiosperm leaf architecture. Journal of the Royal Society of N.Z. 21: 297-312. Pole, M.S., 1992a. Early Miocene flora of the Manuherika Group, New Zealand. I. Ferns. Journal of the Royal Society of N.Z. 22: 279-286. Pole, M.S., 1992b. Eocene vegetation from Hasties, north-eastern Tasmania. Australian Systematic Botany 5:431-475. Sturm, M., 1971. Die Eozane Flora von Messel bei Darmstadt. I. Lauraceae. Palaeontographica Abt B 134: 1-60. Wolfe, J.A., 1977. Paleogene floras from the Gulf of Alaska region. Geological Survey Professional Paper997.

Received 25 November 1991; accepted 23 February 1993

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