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Because of the T. The oldest record T. Fur- panian, c. In contrast to most other organism groups with a better understanding of their relations L. Dettman, in prep. The choice of continent where pollen of T. Dettman, Africa and Madagascar separated already Our goal with the discussion below is to give hypotheses in the Late Jurassic, c. This is long before the on the biogeography of Gunnera by using our cladogram on time calculated for the split between Gunnera and Myrot- the genus as the main startpoint.

The of endemism in biogeographical studies Humphries, ; sister position of Africa relative to southern South America Patterson, This practice was claimed to be one of the and Australasia, found in Gunnera has earlier been retrieved causes of the ambiguous results sometimes obtained in these in other biogeographical studies. It was first discovered in studies, where South America appeared at different levels in some groups of chironomid midges by Brundin and the cladogram.

On the basis of the the Falklands Plateau was severed near the transition distributions of different organisms, Crisci et al. Southern South America and Australasia north of c. In Gunnera, the basal split separates the area of G. Gunnera herteri inhabits and geological patterns the Atlantic coast in southern Brazil and Uruguay between lat. We interpret the distribution of G. This pattern is well-known in biogeography 2 extends the border identified by Crisci et al. Southern South clade including subgenera Misandra and Panke, on the other America is inhabited by a clade formed by subgenus Mis- hand, inhabits southern and Andean South America andra and Panke.

This clade is sister to a group including the according to Crisci et al. Gunnera macro- positions of G. The two phylla, the only representative of subgenus Pseudogunnera species of the genus Myrothamnus, sister group of Gunnera, inhabits New Guinea and the Malayan Archipelago reaching are distributed, in southern Africa and Madagascar, Sumatra, Borneo and the Philippines in the North.

We respectively. It Geologically, New Guinea is part of the same continental seems reasonable that the present distribution of G. For many organisms, large part of New Guinea and The present distribution of G. Gunnera perpensa also changes during the Tertiary. If so, the fossils of Gunnera occurs in a few mountain localities in central Madagascar. The rophylla or its close relatives.

Wanntorp than to southern South America Fig. Several area clad- be of direct Gondwana origin but immigrated by dispersal ograms have shown the same pattern Humphries, ; from Australia. A dispersalist Manos, , where New Zealand and Australia are explanation would have to be more complicated given the repeatedly found as more closely related to each other than relationship and distribution of subgenus Milligania Fig. The geological history of Gondwana breakup indicates a closer relationship between South America and Australia.

One species of Milligania, South America and Australia, on the other hand, G. In the cladogram, the Tasmanian species Mid Tertiary, c. If the split and the Antarctic Peninsula were separated and also the between G. There have been two strategies New Zealand, the same event could not also be used to towards the resolution of this apparent conflict between the explain the geographical pattern within Milligania, at history of vicariance events identified by several phylo- another level in the cladogram. We therefore interpret this genetic analyses and the sequence of geological breakup of pattern as caused by dispersal of the ancestor of G.

New Zealand shares a considerable number of Others, relying heavily on the geological history doubted the angiosperm genera with Australia and dispersal between patterns obtained from biogeographical analyses. This is not necessarily the case. The Winkworth et al. This formidable distance may well have held biogeographical barriers, climatic and In the area of southern South America, subgenus Misandra, others, as daunting as seas or deserts to organisms. Our poor consisting of two species only, is widely sympatric with knowledge of Antarctic paleobiogeography surely introduces subgenus Panke Bader, This is mostly the result of an element of uncertainty but it seems simplistic, just to the widespread G.

The second species of subgenus Misandra, According to the pollen records of T. Subgenus was present in both New Zealand and Australia in the Late Panke is distributed with many species along the rainy slopes Cretaceous, at the time of the separation of the Tasmantian of the Andes from southern Chile to Colombia and beyond, plate from Australia. In the absence of any evidence to the to the mountains of Central America.

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Outside the Andes contrary, we therefore accept the biogeographical pattern there are three species of Panke in the Juan Fernandez Is- found in Gunnera as representing a vicariance event between lands off Chile, and one species in the Serra do Mar in New Zealand and Australia. If this interpretation is correct, it eastern Brazil. Moreover, one species of Panke grows in greatly strengthens a vicariance interpretation also for other Mexico in the southernmost part of the North American organisms, including Nothofagus, extant in these areas landmass and two species are endemic to Hawaii.

Pole argued, partly from geolo- There is, unfortunately, not much resolution within gical evidence, that the entire flora of New Zealand might Panke to draw any well-founded conclusions about the have suffered from more or less wholesale extinctions during relationship and the biogeography of its South American the Tertiary.

Most of the New Zealand flora would thus not species. However, the positions of G. The submerged Kerguelen Plateau, however. In the Cretaceous present Hawaiian islands are not older than 5 Myr but the Kerguelen plate was situated close to Antarctica and there may have been a continuous sequence of emergent Australia and also close to the Indian plate McLoughlin, volcanoes on the Hawaiian seamount chain from at least The entire Hawaiian occurrence should reflect a relationship to Australia and flora has originated through long-distance dispersal from subgenus Pseudogunnera rather than to Africa and subge- different areas, mainly eastern Asia and North America nus Gunnera.

Skottsberg, ; Vargas et al. Gunnera, however, was widespread and abun- Gunnera belongs to the flora that was present in Gondwana dant in North America from the Late Cretaceous to the before its final breakup. Much of its biogeographical history Eocene 80—50 Ma. A sister group rela- size of T. This made Jarzen with the southern South American species more distantly hypothesize a South American relationship of the North related also echoes earlier patterns from botanical analyses, American species of Gunnera.

Our analysis demonstrates but is in conflict with some zoological analyses and also, at that Gunnera is of Gondwanan origin and Misandra from least apparently with geological history. In contrast to some southern South America is the sister group of Panke. The earlier studies, we interpret this pattern as due to vicariance, basal position within Panke of the North American claiming that all barriers were not necessarily ocean straits, G.

We maintain that clad- the Campanian, 80 Ma, leading to the colonization of ogram structure in conjunction with geological evidence North America. Subgenus Panke might then well be an supports a continuous presence of Gunnera in New Zealand originally North American taxon originated by early long- since the Cretaceous.

This is in conflict with the view that distance dispersal from South America leading to the most of the biota of New Zealand was extinguished by separation of Misandra in southern South America and Tertiary geological events. The position of the sole Tasma- Panke in North America.

Dispersal from North America to nian species nested among the New Zealand clade is best Hawaii during the Tertiary then gave origin to the explained by dispersal from New Zealand. In subgenus Hawaiian clade. According to the fossil record, Gunnera Panke, the basal position of the single North American retreated from higher latitudes in North America during the species as well as the Hawaiian clade indicates early dis- Tertiary, where no pollen records exist beyond the Eocene persal from southern South America. This also explains the Jarzen, leaving only a single species, G. Hawaii may the southern extreme.

At the emergence of the Panama land have been reached by dispersal from North America. Finally, bridge or earlier, the Andes may have been recolonized we suggest that Panke may actually have colonized South from the north, leading to extensive sympatry of the sub- America relatively late by long-distance dispersal from genera Misandra and Panke in South America. In contrast North America.

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Benham Auckland Regional Botanic the species. We are grateful to M. The Indian Gunnera: African or Australian origin? Cantrill in India during the Tertiary. We also thank two anonymous reviewers who con- It is, therefore, unlikely that the Indian fossil of T. Wanntorp acrodont lizard phylogenetics. Bader, F. Manos, P. American Bremer, K. Cladistics, Journal of Botany, 84, — Mattfeld, J.

Ostenia, — Evolution, 19, — McLoughlin, S. Australian ficance as evidenced by chironomid midges. Kungliga Svenska Journal of Botany, 49, — Mora-Osejo, L. In: Flora de Colombia Carson, H. Pinto and P. Ruiz , Vol. Instituto de of the Hawaiian Islands. In: Hawaiian biogeography ed. Wagner and V. Funk , pp. Smithsonian Parenti, L. Nelson and D. Rosen , pp. Cookson, I. Patterson, C. In: Vica- Australian Journal of Botany, 2, — Cooper, R.

Columbia University Press, New York. Trends in Ecology Pole, M. Journal of Biogeography, 21, — Crisci, J. Praglowski, J. Grana, 10, — Systematic Zoology, 40, — Schindler, A. In: Das Pflanzenreich Eriksson, T. Department of ed. Engler , IV, Vol. Verlag von Botany, Stockholm, Sweden. Wilhelm Engelmann, Leipzig. Hill, R. Trends in Ecology and Evolution, 7, — Special Hill, R.

In: The ecology and biogeography of Skottsberg, C. Veblen, R. Hill and J. Read , Atlantic and Antarctic vegetation. Geography Journal, 24, pp. Hoot, S. In: Essays in basal eudicots based on three molecular data sets: atpB, rbcL geobotany in honour of William Albert Setchell ed. Annals of the Goodspeed , pp. Humphries, C. In: Advances in southern cold temperate zone.

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