New Caledonian Araucaria

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The main diversity among genus Araucaria is hosted in New Caledonia , where 14 species, all endemic, are described out of a total of 20 extant species. [1] These New Caledonian species are mainly found as dispersed populations in open areas, where competition is less intense. [2]

Contents

Araucaria humboldtensis growing in altitude shrubland, on the slopes of Mont Humboldt, New Caledonia Araucaria humboldtensis growing in altitude shrubland, on the slopes of Mont Humboldt, New Caledonia.jpg
Araucaria humboldtensis growing in altitude shrubland, on the slopes of Mont Humboldt, New Caledonia

New Caledonia, considered as the smallest of the most significant biodiversity hotspots in the world, [3] hosts a unique flora of which 75.1% is endemic. [4] Species of Araucaria trees can be found in every habitat that New Caledonia possesses. [2] However, almost all of them are growing on ultramafic substrate, characterized by low fertility (low N, P, K levels) and high levels of heavy metals (nickel, cobalt, etc.). [5]

Phylogenic history

Context

An alternative to the view that the island is a museum for plant relicts is the possibility that the diversity results from a recent radiation, subsequent to the island's emergence. [6] New Caledonia was part of the supercontinent Gondwana, and separated from Australia 80 Ma. [6] However, geological evidence suggests that New Caledonia was submerged during Paleocene (ca. 65 Ma) and Eocene (until ca. 37 Ma). [7] The actual New Caledonian biota may then result from a total recolonization since the Oligocene. [8]

Araucaria columnaris growing on calcareous substrate, in the South of New Caledonia Araucaria columnaris Prony 2.jpg
Araucaria columnaris growing on calcareous substrate, in the South of New Caledonia

The first Araucaria species were mentioned by Guillaumin in 1948: A. balansae, A. cookie, A. montana , A. muelleri and A. rulei . [9] In 1949, J. T. Bucholz added three new species: A. bernieri , A. biramuluta and A. humboldtensis . In 1972, D. J. de Laubenfels described five other species: A. luxurians , separated from A. cookie renamed A. columnaris , A. laubenfelsii , A. nemorosa , A. scopulorum and A. schmidii , [10] and divided these thirteen species in two groups. [2] The first includes species following the Massart model (plagiotropic branches, partial reiterations) while the second includes species following the Rauh model (orthotropic branches and no reiterations) with bigger leaves. [11] Veillon described how the morphology of New Caledonian Araucaria conforms to these models and noticed that species following the Rauh model have bigger leaves. He also designed a key to help in field identification, based on characters fixed in adults, so ecological factors impacting tree morphology won't interfere with identification. [2]

New Caledonian Araucaria species belong to the Eutacta section, one of the four sections defined by Wilde and Eames in 1952. [12] This section also includes A. heterophylla from Norfolk Island, and A. cunninghamii hosted by both Australia and New Guinea. [12]

Rejection of the Gondwanan hypothesis

In 1998, first genetic analyses based on RBCL gene sequence validated the recognition of the four sections within genus Araucaria. [13] Among Eutacta section, New Caledonian species formed a monophyletic group where A. cunninghamii (Papua New Guinea) was derived first, then A. heterophylla (Norfolk Island). The New Caledonian species revealed a strong homology for rcbL sequences (from 99.5 to 100%), where 10 out of 13 species are identical for this gene sequence. [13] This strong homology and A. heterophylla as a sister group of New Caledonian species (the Norfolk Island being relatively young, less than 3 million years old) are first elements suggesting a recent differentiation of Araucaria trees in New Caledonia. This hypothesis comes in opposition of an older Gondwanan origin.

Gaudeul et al. attempted in 2012 to better describe the evolutionary relationships and diversification of New Caledonian species by using AFLP markers and by performing Bayesian, genetic distances and cladistics analyses. [14] Ecological, morphological and geographical parameters were also considered in the study, which ended supporting a recent diversification of the genus in New Caledonia. Moreover, another genetic group was created: coastal species. However, no evidence were shown for more environment implication in driving speciation, which may be the result of both adaptation and allopatry. The concept of cryptic species, rare among such an iconic group, is mentioned, regarding several divergent populations. [14]

In the meantime, Escapa and Catalano published a phylogenetic analysis using parsimony. [15] For the first time in the family Araucariaceae, genetic (19 plastid, 2 nuclear and 2 mitochondrial genomic regions) and morphologic data (52 discrete and 10 continuous characters) were combined, confirming a strong monophyly for the 4 sections existing. However, relationships among New Caledonian species remain difficult to elucidate even if their relatively recent origin is confirmed by phylogeny based on combination of plastid and nuclear data, and the use of molecular clock. [6] While this study seems to corroborate with overwater dispersal consecutively to the submersion of New Caledonia, New Caledonian Araucaria species seems too old to originate from one single dispersal from Australia to Norfolk Island, and then to New Caledonia.

Cryptic diversity

Araucaria muelleri population, Yate, New Caledonia Yate - Chute de la Madeleine.jpg
Araucaria muelleri population, Yaté, New Caledonia

The complexity of species identification and delimitation in New Caledonian Araucaria is illustrated by Rusham et al. (2016), [16] underlying a cryptic diversity present between two relative species (A. rulei and A. muelleri). This work leads to the identification of a fourteenth endemic species, Araucaria goroensis, [1] confirming the difficulty to distinguish species among the Monkey Puzzle genus in the territory. This new species was initially confused with A. muelleri, but it appears to be more closely related to A. rulei. Distinctions with the latter hold in larger leaves, microsporophylls without a shouldered base and shorter female cone bracts. [1]

The origin of such a diversity among the genus Araucaria in New Caledonia still remain unclear, but appears to be relatively recent. Also speciation may be still be processed as suggested by the cryptic diversity observed among the genus. [14]

GroupSpeciesIUCN statusLeaf length (mm)Leaf width (mm)Architectural modelSubstrateHabitatAltitude (m)
Coastal A. columnaris LC5-74-5MassartCalcareousCalcareous platforms< 50
A. nemorosa CR6-101.5-3MassartUltramaficMaquis< 100
A. luxurians EN5-74-5MassartUltramaficHumid evergreen forest< 300
Small-leaved A. bernieri VU2-3.51.5-2.5MassartUltramaficHumid evergreen forest, deep valleys100 - 800
A. schmidii VU7-101.5-2MassartAcidicMontane cloud forest> 1400
A. scopulorum EN3-42.5-3MassartUltramaficMaquis50 - 800
A. subulata NT4-62-2.5MassartUltramaficHumid evergreen forest, deep valleys100 - 1100
Large-leaved A. biramulata VU7-95-6MassartUltramaficHumid evergreen forest, deep valleys300 - 1000
A. goroensisEN26-331-16RauhUltramaficMaquis150 - 550
A. humboldtensis EN5-64-5MassartUltramaficMaquis of high altitude600 - 1500
A. laubenfelsii NT12-208-10MassartUltramaficEvergreen forest or maquis700 - 1200
A. montana VU11-147-8MassartUltramafic and acidicDense humid forest or maquis400 - 1200
A. muelleri EN30-3515-20RauhUltramaficDense humid forest or maquis150 - 1200
A. rulei EN20-2511-14RauhUltramaficDense humid forest or maquis400 - 800

Related Research Articles

<i>Araucaria</i> genus of evergreen conifers in the family Araucariaceae

Araucaria is a genus of evergreen coniferous trees in the family Araucariaceae. There are 20 extant species in New Caledonia, Norfolk Island, eastern Australia, New Guinea, Argentina, Chile, Brazil, and Paraguay.

<i>Araucaria laubenfelsii</i> species of plant

Araucaria laubenfelsii is a species of conifer in the family Araucariaceae. It is found only on Grande Terre, the main island of New Caledonia, mainly on the southern mountains Mont Mou, Mont des Sources, Mont Dzumac and Mont Dou, though smaller populations also exist on Mont Kaala and Mont Canala in the north. It is one of the larger of New Caledonia's native araucaria species, sometimes reaching up to 50 meters in height in emergent rainforest specimens, and potentially living for as long as 500 years or more. As with other New Caledonian araucaria species, Araucaria laubenfelsii is threatened by habitat loss, though southern populations at least are considered to be healthy and it is not currently considered vulnerable or endangered. A population genetic study suggested that A. laubenfelsii is doubtfully distinct from the more widespread A. montana.

Solmsia is a genus containing one or two species of flowering plants belonging to the family Thymelaeaceae. It is endemic to New Caledonia. The genus was named to honor Hermann zu Solms-Laubach by Henri Ernest Baillon. It is related to Arnhemia, Deltaria, Gonystylus and Lethedon.

Oncotheca is a genus of tree endemic to New Caledonia. There are two species, Oncotheca balansae and Oncotheca humboldtiana.

Nephrodesmus is a genus of flowering plants in the legume family, Fabaceae. It belongs to the subfamily Faboideae. It contains c. 5 species, all endemic to New Caledonia. Its closest relatives is Arthroclianthus, also endemic to New Caledonia and their distinction has been challenged.

Achlydosa glandulosa is a species of orchid endemic to New Caledonia and the only species of the genus Achlydosa. This species has previously been placed in the genus Megastylis. Its closest relative is Pterostylis, the sole other genus of subtribe Pterostylidinae.

Amphorogyne is a genus of hemiparasitic trees and shrubs in the family Santalaceae. The genus is endemic to New Caledonia in the Pacific and contains three species. Its closest relative is Daenikera, also endemic to New Caledonia.

<i>Stromatopteris</i> genus of plants

Stromatopteris is a genus of fern in the family Gleicheniaceae, endemic to New Caledonia. The only species in the genus is Stromatopteris moniliformis. Its closest relative is the more widespread genus Gleichenia.

Nemuaron vieillardii is a species of trees in the Atherospermataceae family. It is endemic to New Caledonia and the only species of the genus Nemuaron. Its closest relative is the monotypic genus Atherosperma from Australia.

Salaciopsis is a genus of shrubs and small trees in the family Celastraceae. The genus is endemic to New Caledonia in the Pacific and contains six species.

Menepetalum is a genus of shrubs and small trees in the family Celastraceae. The genus is endemic to New Caledonia in the Pacific and contains four species. Its closest relative is Dinghoua from Australia.

Peripterygia marginata is a species of shrubs in the family Celastraceae. It is endemic to New Caledonia and the only species of the genus Peripterygia. Its closest relatives are Crossopetalum and Siphonodon.

Dicarpellum is a genus of shrubs and small trees in the family Celastraceae. The genus is endemic to New Caledonia in the Pacific and contains four species. Its closest relative is Hypsophila from Australia.

<i>Loxodiscus</i> genus of plants

Loxodiscus coriaceus is a species of shrubs in the family Sapindaceae. It is endemic to New Caledonia and the only species of the genus Loxodiscus. Its closest relatives are Cossinia, Diplopeltis, Dodonaea and Harpullia.

Daenikera corallina is a species of parasitic in the Santalaceae family. It is endemic to New Caledonia and the only species of the genus Daenikera. Its closest relative is Amphorogyne, also endemic to New Caledonia.

Elaphanthera baumannii is a species of hemiparasitic shrub in the Santalaceae family. It is endemic to New Caledonia and the only species of the genus Elaphanthera, probably related to Exocarpos.

<i>Thiollierea</i> genus of plants

Thiollierea is a genus of shrubs in the family Rubiaceae. The genus is endemic to New Caledonia in the Pacific and contains 14 species which used to be placed in the genera Bikkia or Morierina

Gastrolepis is a genus of shrubs and trees in the family Stemonuraceae. The genus is endemic to New Caledonia in the Pacific and contains two species.

Periomphale balansae is a species of shrub in the Alseuosmiaceae family. It is endemic to New Caledonia and the only species of the genus Periomphale. It has sometimes been included in the genus Wittsteinia.

Acianthus elegans, commonly known as the elegant acianthus, is a species of mosquito orchids, plants in the genus Acianthus. It is found in New Caledonia.

References

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