Fruits and leaves with cuticle of Laurelia otagoensis sp. nov. (Atherospermataceae) from the early Miocene of Otago (New Zealand)

Conran, J.G., Bannister, J.M. & Lee, D.E., 2013. Fruits and leaves with cuticle of Laurelia otagoensis sp. nov. (Atherospermataceae) from the early Miocene of Otago (New Zealand). Alcheringa 37, 496–509. ISSN 0311-5518.

Laurelia otagoensis sp. nov. Conran, Bannister & D.E. Lee (Laurales: Atherospermataceae) is described from the earliest Miocene Foulden Maar diatomite deposit, Otago, New Zealand. The new species is represented by mummified fossil leaves with well-preserved cuticle and associated clusters of achenes bearing persistent, long plumose styles. This basal angiosperm family is of significance because of its classic southern disjunctions and ecological importance in extant Gondwana-type rainforests, but has a very sparse fossil record. The present study describes one of very few convincing leaf fossils for Atherospermataceae and the only definitive fossil fruits. The presence of fossil Laurelia in Oligo–Miocene New Zealand combined with fossil leaf impressions from the late Eocene, Miocene dispersed cuticle and pollen from the Oligocene to Holocene shows that the family has had a long history in Cenozoic New Zealand. These new fossils also support palaeoclimatic data suggesting warmer conditions in the earliest Miocene of New Zealand.

John G. Conran [john.conran@adelaide.edu.au], Australian Centre for Evolutionary Biology & Biodiversity and Sprigg Geobiology Centre, School of Earth and Environmental Sciences, Benham Bldg DX 650 312, The University of Adelaide, SA 5005 Australia; Jennifer M. Bannister [jennifer.bannister@xtra.co.nz], Department of Botany, University of Otago, PO Box 56, Dunedin, New Zealand; Daphne E. Lee [daphne.lee@otago.ac.nz], Department of Geology, University of Otago, PO Box 56, Dunedin, New Zealand. Received 17.12.2012; revised 9.4.2013; accepted 15.4.2013.     

Tertiary Nothofagus macrofossils from Cethana,Tasmania

Two new Tertiary species of Nothofagus from the Early Eocene-Oligocene deposit at Cethana represent the first reports of fossil species which are not closely related to the extant Australian species N. moorei and N. cunninghamii. N. cethanica sp. nov. is most closely related to the extant New Zealand species N. fusca and N. truncata and gives further evidence of the relatively slow evolution within this genus. The other specimen is indistinguishable from extant adult N. gunnii leaves, and has been assigned to that species. This fossil shows that the deciduous habit was probably already present in N. gunnii by the Oligocene, and this may have helped N. gunnii to survive the Late Tertiary/Quaternary glaciations. Juvenile N. gunnii foliage gives some insight into the possible origins of this species, which may have been from the same ancestral stock as N. fusca, N. truncata, and N. cethanica.     

Biodiversity and palaeoecology of Foulden Maar: an early Miocene Konservat-Lagerstätte deposit in southern New Zealand

Lee, D.E., Kaulfuss, U, Conran, J.G., Bannister, J.M. & Lindqvist, J.K., August 2016. Biodiversity and palaeoecology of Foulden Maar: an early Miocene Konservat-Lagerstätte deposit in southern New Zealand. Alcheringa 40, xxx–xxx. ISSN 0311-5518.

This paper highlights the biodiversity and palaeoecology of the 23 million year old Foulden Maar, the first Konservat-Lagerstätte deposit described from New Zealand and a key site for reconstructing early Miocene Southern Hemisphere terrestrial ecosystems. The 1000-m-diameter, ca 200-m-deep Foulden Maar volcanic crater lake was a closed system with anoxic bottom waters, capturing and preserving in exquisite detail organisms from the lake and adjacent rainforest. The fossils include numerous leaves, flowers with in situ pollen, fruits, seeds, fish and arthropods. Surrounding Foulden Maar was an evergreen, Lauraceae-dominated notophyll vine forest with a diverse understorey, lianes, epiphytes and mistletoes. Diverse pollination and seed dispersal modes are evident. Fish include larval to adult stages of articulated specimens of Galaxias, some with preserved soft tissue and a species of eel resembling Anguilla. The arthropod fauna comprises ca 20 families in the orders Araneae (spiders), Plecoptera (stoneflies), Odonata (dragonflies), Isoptera (termites), Hemiptera (true bugs), Diptera (true flies), Coleoptera (beetles), Trichoptera (caddis flies) and Hymenoptera (wasps, ants and bees), representing faunas typical of soil, leaf litter, forest floor or freshwater habitats. Many fossil taxa have close relatives in the extant New Zealand biota; others are now locally extinct. Coprolites containing quartz sands sourced from outside the lake indicate the presence of volant birds, presumably waterfowl. The Foulden Maar Lagerstätte is crucial for reconstructing Miocene lake and forest ecosystems in New Zealand, particularly the terrestrial arthropod component.

Daphne E. Lee [daphne.lee@otago.ac.nz], Department of Geology, University of Otago, PO Box 56, Dunedin, New Zealand., Uwe Kaulfuss [uwe.kaulfuss@otago.ac.nz] and Jon K. Lindqvist, [jonlind@ihug.co.nz], Department of Geology, University of Otago, PO Box 56, Dunedin, New Zealand; John G. Conran [john.conran@adelaide.edu.au], Australian Centre for Evolutionary Biology & Biodiversity and Sprigg Geobiology Centre, School of Biological Sciences, Benham Bldg DX 650 312, The University of Adelaide, SA 5005, Australia; Jennifer M. Bannister [jennifer.bannister@xtra.co.nz], Department of Botany, University of Otago, PO Box 56, Dunedin, New Zealand.     

Fossil arachnids from the earliest Miocene Foulden Maar Fossil-Lagerstätte,New Zealand

Selden, P.A. & Kaulfuss, U., May 2018. Fossil arachnids from the earliest Miocene Foulden Maar Fossil-Lagerstätte, New Zealand. Alcheringa XX, XX–XX. ISSN 0311-5518.

Fossil Arachnida from New Zealand are extremely rare and represented by some unidentifiable amber inclusions only. The first fossil arachnids from New Zealand to be described in detail are presented here, based on four compression fossils from the earliest Miocene Fossil-Lagerstätte at Foulden Maar, South Island. Two specimens are referred to Arachnida incertae sedis and Araneomorpha incertae sedis, respectively, whereas two specimens are mygalomorph spiders. One of these is placed in the Rastelloidina clade of Mygalomorphae, probably in the Idiopidae, which is represented in New Zealand by the extant Cantuaria.

P.A. Selden* [selden@ku.edu] Paleontological Institute and Department of Geology, University of Kansas, Lindley Hall, 1475 Jayhawk Boulevard, Lawrence, KS 66045, USA; U. Kaulfuss [uwe.kaulfuss@otago.ac.nz] Department of Geology, University of Otago, PO Box 56, Dunedin, New Zealand. *Also affiliated with: Natural History Museum, London, UK.     

A Devonian fish fauna from subsurface sediments in the eastern Officer Basin,South Australia

This paper reports the discovery of three of the most iconic New Caledonian endemic genera, Amphorogyne, Paracryphia and Phelline, as dispersed leaf cuticle fossils in the early Miocene of New Zealand. New Caledonia's endemic angiosperm families have given it a reputation as one of the most interesting botanical regions in the world, but unfortunately it has no known pre-Pleistocene Cenozoic plant fossil record. A once more widespread distribution of its key plants in the context of a cooling and drying Neogene world suggests the current vegetation of New Caledonia is the result of contraction, or even a migration, from more southerly landmasses. Thus, New Zealand may have been a source of at least some of New Caledonia's plants.     

An early Miocene ant (subfam. Amblyoponinae) from Foulden Maar: the first fossil Hymenoptera from New Zealand

Kaulfuss, U., Harris, A.C., Conran J.G. & Lee, D.E., 2014. An early Miocene ant (subfam. Amblyoponinae) from Foulden Maar: the first fossil Hymenoptera from New Zealand. Alcheringa 38, 568–574. ISSN 0311-5518.

The ant subfamily Amblyoponinae is presently represented in New Zealand by one endemic species in the cosmopolitan genus Stigmatomma and an introduced Australian species of Amblyopone. The fossil record of the group is restricted to two species of Stigmatomma from late Eocene Baltic Amber. Here, we describe the third fossil record, an Amblyopone-like specimen from the early Miocene of Otago, southern New Zealand, based on a winged male that resembles the extant A. australis Erichson in size, general habitus and characters of wing venation, but also shares features with the African amblyoponine genus Zymmer. This represents the first fossil record of Amblyoponinae from the Southern Hemisphere and the first example of Hymenoptera among the few pre-Quaternary insect fossils known from New Zealand. It suggests a long history of Amblyoponinae in New Zealand and Australia.

Uwe Kaulfuss [uwe.kaulfuss@otago.ac.nz] and Daphne E. Lee [daphne.lee@otago.ac.nz], Department of Geology, University of Otago, PO Box 56, Dunedin 9016, New Zealand; Anthony C. Harris [anthony.harris@otagomuseum.govt.nz], Otago Museum, PO Box 6202, Dunedin 9059, New Zealand; John G. Conran [john.conran@adelaide.edu.au], ACEBB & SGC, School of Earth and Environmental Sciences, The University of Adelaide, Benham Bldg, DX 650 312, Adelaide SA 5005, Australia. Received 18.3.2014; revised 15.5.2014; accepted 23.5.2014.     

Two new camerate crinoid species from the Permian of eastern Australia

The habitat and habit of Australia's first recorded Tertiary marsupial species, Wynyardia bassiana, found some 130 years ago at Wynyard on the northwestern coast of Tasmania, remain enigmatic (Aplin 1987, Aplin &; Rich 1990). Fossil pollen and spores preserved in a rafted clast of estuarine silts from the same sequence of earliest Miocene marine sandstones as the skeletal remains indicate the local vegetation was Nothofagus-gymnosperm evergreen rainforest, probably with a cryptogam-rich rather than woody subcanopy stratum. Comparisons with present-day Nothofagus rainforests suggest that, although the subcanopy would have been sufficiently open to allow the passage of a large ground-dwelling herbivorous marsupial, limited food resources are more consistent with Wynyardia being a generalist arboreal herbivore.     

Silicified wood of Early Miocene Nothofagus,Acacia and Myrtaceae (aff. Eucalyptus B) from the upper Lachlan valley,New South Wales

Three specimens of silicified wood, two transported and one probably in situ, have been found in association with Early Miocene basaltic lavas in the upper Lachlan valley, N.S.W. On the basis of their preserved structure, the three specimens have been identified as belonging to Nothofagus, Acacia, and the family Myrtaceae, respectively. The specimen of Nothofagus (one of the transported specimens) constitutes the first identification of a macrofossil of this genus from this interval in southeastern mainland Australia, while its conjunction with the myrtaceous specimens (one probably in situ) may indicate vegetation zonation according to elevation, as proposed on pollen evidence from Kiandra. Such zonation is not demanded by the upper Lachlan evidence, however, but if it is favoured, reconstruction of the Early Miocene geography of the upper Lachlan constrains the maximum elevation above the locality of the myrtaceous fossils to be 350 m.     

A new Cenozoic cyclostome bryozoan genus from Argentina and New Zealand: strengthening the biogeographical links between South America and Australasia

Taylor, P.D. & Brezina, S., February 2018. A new Cenozoic cyclostome bryozoan genus from Argentina and New Zealand: strengthening the biogeographical links between South America and Australasia. Alcheringa XX, xx–xx. ISSN 0311-5518.

Uniserial encrusting cyclostome bryozoans (‘stomatoporiforms’) are especially challenging to study taxonomically because of the paucity of their morphological characters. Here we introduce Axilosoecia gen. nov. for two previously undescribed species characterized by gonozooidal brood chambers located in the axils of branch bifurcations. The type species, Axilosoecia giselae sp. nov., comes from the upper, Danian part of the Roca Formation of La Pampa, Argentina; the second species, Axilosoecia mediorubiensis sp. nov., is from the lower Miocene of Southland, New Zealand. On account of its basal gonozooids, Axilosoecia is assigned to the family Oncousoeciidae despite similarities in colony form with Stomatoporidae. The two known occurrences of this new genus support previously suggested biogeographical links between southern South America and Australasia.

Paul D. Taylor, Department of Earth Sciences, Natural History Museum, Cromwell Road, London SW7 5BD, UK [p.taylor@nhm.ac.uk]; Soledad Brezina, Instituto de Investigación en Paleobiología y Geología, Unidad de doble dependencia, Universidad Nacional de Río Negro—Consejo Nacional de Investigaciones Científicas y Técnicas, Av. Roca 1242, R8332FDJ, General Roca, Río Negro, Argentina [sbrezina@unrn.edu.ar].     

A review of Australasian ichthyosaurs

Ichthyosaur fossils have been recorded from four landmasses in the Australasian region—Australia, New Zealand, New Caledonia and Timor—and occur in all three systems of the Mesozoic. Most of the remains are non-diagnostic, but at least three genera have been identified: Mixosaurus, from the Middle Triassic of Timor; Shonisaurus, from the Upper Triassic of New Caledonia; and Platypterygius, from the Lower Cretaceous of Australia and New Zealand. Of these, Platypterygius contains the only material that can be diagnosed to species level. However, current taxonomy of the specimens is controversial, with two synonyms, P. australis and P. longmani, persisting in the literature. An examination of cranial traits in the ‘quasi-holotype’ of P. australis vs P. longmani demonstrates that they represent the same taxon. Thus, P. longmani should be regarded as the junior synonym. A neotype is also here designated for P. australis to replace the original, which is presumed lost.     

Acaste (Trilobita: Phacopina) from the Early Devonian of Tasmania

Edgecombe, G.D. March 2007. Acaste (Trilobita: Phacopina) from the Early Devonian of Tasmania. Alcheringa 31, 59-66. ISSN 0311-5518.

The widespread Siluro-Devonian acastid trilobites Acaste and Acastella are known from few occurrences in Australasia, these being confined to Lochkovian strata in Victoria and New Zealand. A new species, Acaste andersoni, is described from a correlative of the Bell Shale near St Valentines Peak in north-western Tasmania. Acaste supplements the biogeographic links between the Early Devonian faunas of New Zealand, Victoria and Tasmania.

Gregory D. Edgecombe [greged@austmus.gov.au], Australian Museum, 6 College Street, Sydney, NSW 2010, Australia; received 2.3.2005, revised 6.4.2005.     

A new record of an aristonectine elasmosaurid (Sauropterygia,Plesiosauria) from the Upper Cretaceous of New Zealand: implications for the Mauisaurus haasti Hector, 1874 hypodigm

O’Gorman, J.P., Otero, R.A. & Hiller, N., 2014. A new record of an aristonectine elasmosaurid (Sauropterygia, Plesiosauria) from the Upper Cretaceous of New Zealand: implications for the Mauisaurus haasti Hector, 1874 Hector, J., 1874. On the fossil Reptilia of New Zealand. Transactions and Proceedings of the New Zealand Institute 6, 333–358. [Google Scholar] hypodigm. Alcheringa 38, 504–512. ISSN 0311-5518

An indeterminate aristonectine elasmosaurid is recorded from a lower Maastrichtian bed of the Conway Formation, Waipara River, South Island, New Zealand. The described specimen (CM Zfr 104), previously considered part of the hypodigm of Mauisaurus haasti, came from the upper part of the Alterbidinium acutulum biozone, the same zone from which the only well-known aristonectine from New Zealand, Kaiwhekea katiki, is recorded. The cervical vertebrae of CM Zfr 104 have the same distinctive features (i.e., with extremely broad rather than long centra) as those from previously recorded juvenile aristonectines from Argentina, Chile and Antarctica. This new record is congruent with the biogeographic relationships of Cretaceous marine amniotes from the Weddellian Palaeobiogeographic Province (i.e., Patagonia, western Antarctica, New Zealand and southeastern Australia). Therefore, this type of vertebra is regarded as a distinctive feature of the Weddellian aristonectine elasmosaurids.

José P. O’Gorman [joseogorman@fcnym.unlp.edu.ar], División Paleontología Vertebrados, Museo de La Plata, Universidad Nacional de La Plata, Paseo del Bosque s/n., B1900FWA, La Plata, Argentina; [CONICET (Consejo Nacional de Investigaciones Científicas y Técnicas)]; Rodrigo A. Otero [paracrioceras@gmail.com], Red Paleontológica U-Chile. Laboratorio de Ontogenia y Filogenia, Departamento de Biología, Facultad de Ciencias, Universidad de Chile, Av. Las Palmeras 3425, Santiago, Chile; Norton Hiller, [norton.hiller@canterbury.ac.nz], Department of Geological Sciences, University of Canterbury, PB 4800, Christchurch, 8001, New Zealand and Canterbury Museum, Rolleston Avenue, Christchurch, New Zealand, 8013.     

The record of Araucariaceae macrofossils in New Zealand

Pole, M., December, 2008. The record of Araucariaceae macrofossils in New Zealand. Alcheringa 32, 405–426. ISSN 0311-5518.

The Araucariaceae have a long record in New Zealand, extending back to the Jurassic at least, and Araucaria extends back to at least the Late Cretaceous. This paper reviews the macrofossil record of the family and presents new information based largely on the leaf cuticle record. Agathis, which is the only genus of the family currently growing in New Zealand, has no record before the Cenozoic. All specimens previously identified from pre-Cenozoic strata clearly belong to other taxa or do not show characteristic features of the genus. Araucariaceae macrofossils are virtually ubiquitous in the Cretaceous assemblages of New Zealand but are conspicuous by their absence or rarity in Palaeocene deposits. Their demise may be an expression of events at the Cretaceous–Palaeogene boundary.     

A palynological investigation of plesiosaur-bearing rocks from the Upper Cretaceous Tahora Formation,Mangahouanga, New Zealand

High-palaeolatitude plesiosaur, mosasaur and, more rarely, dinosaur fossils are well known from the Maungataniwha Sandstone Member of the Tahora Formation in Mangahouanga Stream, Hawkes Bay, New Zealand. A palynological investigation of strata exposed along Mangahouanga Stream and of transported boulders hosting vertebrate fossils reveals well-preserved assemblages dominated by terrestrial pollen and spores but also including marine dinoflagellate cysts in some samples. The palynofacies are strongly dominated by wood fragments including charcoal; one outcrop sample and the sample taken from a boulder hosting plesiosaur vertebrae contain entirely terrestrially derived palynoassemblages, suggesting a freshwater habitat for at least some of the plesiosaurs. The host unit spans the Santonian to lowermost Maastrichtian, while the key pollen taxa Nothofagidites senectus and Tricolpites lilliei, together with the dinocyst Isabelidinium pellucidum and the megaspore Grapnelispora evansii, indicate a late Campanian to early Maastrichtian age for the fossiliferous boulders. The palynoflora indicates a mixed local vegetation dominated by podocarp conifers and angiosperms with a significant tree-fern subcanopy. The presence of taxa with modern temperate distributions, such as Nothofagus (southern beech), Proteaceae and Cyatheaceae (tree-ferns), indicates a mild-temperate climate and lack of severe winter freezing during the latest Cretaceous, providing an ecosystem that most probably made it possible for polar dinosaurs to overwinter in this part of the world.     

A fossil sea turtle (Testudines: Pan-Cheloniidae) from the upper Oligocene Pomahaka Formation,New Zealand

Gard, H.J.L. & Fordyce, R.E., August 2016. A fossil sea turtle (Testudines: Pan-Cheloniidae) from the upper Oligocene Pomahaka Formation, New Zealand. Alcheringa 41, XX–XX. ISSN 0311-5518.

An isolated turtle xiphiplastron similar to that of Puppigerus sp. is described from the upper Oligocene (27.3–25.2 Ma) Pomahaka Formation near Tapanui, Otago, New Zealand. The bone is unlike any previously described turtle from the Cenozoic of New Zealand and is from a newly recognized estuarine vertebrate locality. It represents the first Oligocene cheloniid turtle bone described from the southwest Pacific.

Henry J. L. Gard [henrygard@hotmail.com] and R. Ewan Fordyce, [ewan.fordyce@otago.ac.nz], Department of Geology, University of Otago, PO Box 56, Dunedin, 9054, Otago, New Zealand.     

Nothofagus macrofossils from the Tertiary of Tasmania

Leaves assignable to Nothofagus from two fossil deposits in Tasmania represent the first macrofossils of this genus from the Tertiary in southeastern Australia. One fossil species, N. johnstonii, is closely related to the extant Australian species N. cunninghamii while the other fossil species, N. tasmanica, has very close affinites with the extant Australian species N. moorei. All four of these species are closely interrelated. The pollen type produced by the fossil species is unknown, since all three types are present in the microfloras. However, both N. cunninghamii and N. moorei produce N. menziesii-type pollen. The macrofossils confirm the conclusion from pollen studies that evolution in Nothofagus has occurred very slowly.     

Polarized light microscopy casts new light on plants used to make tapa

Information about plant materials of construction in artefacts advances knowledge about human history, agriculture, trade, migration and adaptation to new environments. Typically, materials identification in artefacts made from plants is problematic, since processing, age, dirt and surface treatments can mask identifying features, while ethical considerations relating to sampling limit the use of some analytical techniques. The study tests the usefulness of polarized light microscopy for identifying the New Zealand and Pacific plant species used to make tapa, indicating birefringent and morphological characteristics that can be used to differentiate fibres at the level Moraceae (Pacific; from genera Artocarpus, Broussonetia and Ficus) and Malvaceae (New Zealand; from genera Hoheria and Plangianthus).     

Mid-Cenozoic ostracod biostratigraphic range extensions and taxonomic notes on selected species from a new Oligocene (Duntroonian–Waitakian) fauna from southern New Zealand

AYRESS, M., ROBINSON, J.H. & LEE, D.E., March 2017. Mid-Cenozoic ostracod biostratigraphic range extensions and taxonomic notes on selected species from a new Oligocene (Duntroonian–Waitakian) fauna from southern New Zealand. Alcheringa 41, 487–498.

This paper discusses a highly diverse assemblage of Ostracoda, particularly biostratigraphically useful species, from a richly fossiliferous shallow-marine deposit of late Oligocene age on Cosy Dell farm, near Waimumu, Southland, New Zealand. The ostracod fauna, consisting of 126 species, is yet to be fully analysed, although it is clear that this new window into the nearshore fauna of the mid-Cenozoic provides the earliest records of many characteristic extant New Zealand genera and species, including several species hitherto only known as far back as the early Miocene (Otaian–Altonian New Zealand local Stages). Finding these younger species in an Oligocene deposit was unexpected and indicates that our current knowledge of mid-Cenozoic ostracod age ranges, particularly for nearshore species, is incomplete. Using nannofossils and molluscs, the Cosy Dell deposit is dated as late Oligocene (Duntroonian or lower Waitakian local Stages) in age; therefore, the new ostracod species occurrences described here are considered valid range extensions. Although the biostratigraphic application of the benthonic ostracod species discussed is limited owing to the effects of sedimentary facies and other palaeoenvironmental factors, the ostracod fossil record does include significant extinction and inception events that offer correlative potential for mid-Cenozoic stratigraphical studies. This study identified a need for modern illustration of many species, and we provide scanning electron microscope illustrations of those Cosy Dell taxa previously considered biostratigraphically useful, as support for species identification and associated biostratigraphic conclusions.

Michael Ayress [mike.ayress@rpsgroup.com], RPS ichron Group, Gadbrook Business Centre, Northwich CW9 7TN, UK; Jeffrey Robinson [jeffreyhrobinson@yahoo.co.nz] and Daphne Lee [daphne.lee@otago.ac.nz], Geology Department, University of Otago, Dunedin, New Zealand     

First record of Procariama simplex Rovereto, 1914 (Phorusrhacidae,Psilopterinae) in the Cerro Azul Formation (upper Miocene) of La Pampa Province; remarks on its anatomy,palaeogeography and chronological range

Vezzosi, R.I., June 2012. First record of Procariama simplex Rovereto, 1914 (Phorusrhacidae, Psilopterinae) in the Cerro Azul Formation (upper Miocene) of La Pampa Province; remarks on its anatomy, palaeogeography and chronological range. Alcheringa, 157–169. ISSN 0311-5518.

New records of Procariama simplex in central and northwestern Argentina are reported. The fossil material includes mandibular bones, cervical vertebrae and several elements of the fore and hindlimbs. After an exhaustive comparison of anatomical characters and morphometric analysis, the fossil specimens are assigned to the psilopterine Procariama simplex. The specimen from La Pampa Province represents the first record of a psilopterine in this central region of Argentina and derives from upper Miocene sediments of the Cerro Azul Formation. Lithological features and biostratigraphical data allow the host bed to be assigned to the Huayquerian faunal stage (late Miocene). The La Pampa record broadens the geographical distribution of Procariama simplex, which was previously restricted to northwestern Argentina.

Raúl Ignacio Vezzosi [vezzosiraul@gmail.com], Centro de Investigaciones Científicas y Transferencia de Tecnología a la Producción CICYTTP-CONICET, Materi y España s/n (3105), Diamante, Entre Ríos, Argentina. Received 1.4.2011, revised 28.5.2011, accepted 14.6.2011.