Journal archives for October 2024

October 7, 2024

A plant that seems to defy biogeographical and evolutionary rules: Dodonaea viscosa (Sapindaceae), part 1

THE PUZZLE

Dodonaea viscosa (https://www.inaturalist.org/taxa/122711-Dodonaea-viscosa and https://prota.prota4u.org/protav8.asp?g=pe&p=Dodonaea+viscosa and https://www.anbg.gov.au/gnp/interns-2007/dodonaea-viscosa.html and https://anpsa.org.au/APOL27/sep02-1.html) is biologically puzzling in at least six ways.

Firstly, it is uniquely cosmopolitan for a woody plant (https://www.mq.edu.au/__data/assets/pdf_file/0005/1275341/Plant-of-the-week-Dodonaea-viscosa-Hop-Bush-ed-2.pdf). Long before human influence, it spread naturally across several oceans and then far inland on several continents.

Secondly, this wide natural distribution was achieved despite the fact that 60 other members of the same genus remain restricted to the same continent - and in many cases the same landscapes - from which D. viscosa originated.

Thirdly, its original occurrence was on a landmass, namely the 'island continent' of Australia/Sahul (https://en.wikipedia.org/wiki/Sahul), that has never been connected to any continent by a land-bridge.

Fourthly, its diaspores seem not to be particularly adapted for long-distance dispersal.

It is true that the seed-capsules have papery wings, suggesting anemochory (https://www.dictionary.com/browse/anemochory), and the seeds survive immersion in sea water (West, 1984, https://www.publish.csiro.au/sb/BRU9840001).

However,

Fifthly, D. viscosa is 'polymorphic', in a way incongruent with subspeciation/raciality/ecotypy (https://en.wikipedia.org/wiki/Ecotype and https://digital.library.adelaide.edu.au/dspace/handle/2440/119801 and https://onlinelibrary.wiley.com/doi/10.1111/j.1365-2699.2009.02176.x).

Please see https://www.inaturalist.org/posts/99976-a-plant-that-seems-to-defy-biogeographical-and-evolutionary-rules-dodonaea-viscosa-sapindaceae#activity_comment_81def12b-fd99-44ea-9b4a-a264e1b52b8c.

In a given population, the leaves may take a distinctive form (e.g. https://wtlandcare.org/details/dodonaea-viscosa-subsp-cuneata/) without any geographical isolation from other forms (see 'key to the subspecies' in https://plantnet.rbgsyd.nsw.gov.au/cgi-bin/NSWfl.pl?page=nswfl&lvl=sp&name=Dodonaea~viscosa).

By the same token, the leaves may remain true to form despite having been subjected to environmental pressures and 'genetic drift' on a completely different continent for hundreds of thousands of years.

Sixthly, D. viscosa has remarkably dense wood (https://www.inaturalist.org/journal/milewski/100396-a-plant-that-seems-to-defy-biogeographical-and-evolutionary-rules-dodonaea-viscosa-sapindaceae-part-2-wood-density#).

DISCUSSION

The literature lacks any satisfactory hypothesis on how D. viscosa achieved its unique spread, either ecologically or mechanistically.

The following review of fruit-form and seed-dispersal in the family Sapindaceae, more generally, highlights the puzzle.

No sapindaceous plant seems to be adapted for long-distance dispersal.

This is because

  • the typical fruit-form in this family is arillate (https://en.wikipedia.org/wiki/Aril), involving seed-dispersing animals ranging from ants through birds to mammals,
  • those fruits adapted - by virtue of papery wings on the mature, dry capsule - to dispersal by wind tend not to move farther than a few hundred metres from the parent individual, and
  • the seeds are not particularly small in any member of the Sapindaceae.

In Dodonaea, the aril is an outgrowth of the funicle (West, page 33). It is

  • always so small that it is better-described as an elaiosome (https://en.wikipedia.org/wiki/Elaiosome) than as an aril, and
  • so small in D. viscosa that some authors have claimed that it is absent.

Two relevant mysteries are

Please also see https://www.inaturalist.org/journal/milewski/100103-a-comparison-of-sapindaceae-in-the-mediterranean-and-adjacent-arid-climates-of-australia-and-southern-africa#.

to be continued in https://www.inaturalist.org/journal/milewski/100396-a-plant-that-seems-to-defy-biogeographical-and-evolutionary-rules-dodonaea-viscosa-sapindaceae-part-2-wood-density#...

Posted on October 7, 2024 10:19 AM by milewski milewski | 39 comments | Leave a comment

October 13, 2024

A comparison of Sapindaceae in the mediterranean and adjacent arid climates of Australia and southern Africa

INTRODUCTION

In both Australia and southern Africa, the southwestern tips of the landmasses have rainfall mainly in winter (https://en.wikipedia.org/wiki/Mediterranean_climate). Adjacent to this mediterranean-type climate at temperate latitudes are semi-arid climates with rainfall mainly/partly in winter (https://en.wikipedia.org/wiki/Climate_of_Australia#/media/File:Australia_K%C3%B6ppen.svg and https://www.wits.ac.za/news/latest-news/opinion/2019/2019-02/scientists-split-on-south-africas-winter-and-summer-rainfall-zones.html).

In Australia, the relevant mediterranean regions occur disjunctly in Western Australia and South Australia. In southern Africa, the mediterranean region is restricted to Western Cape province, while the adjacent arid region extends to the westernmost part of Northern Cape province (https://en.wikipedia.org/wiki/Northern_Cape).

AIMS

In this Post, I make an intercontinental comparison of the sapindaceous indigenous floras of the two landmasses.

RESULTS: AUSTRALIA

Alectryon oleifolius https://www.inaturalist.org/taxa/783975-Alectryon-oleifolius and https://cdn.environment.sa.gov.au/landscape/docs/saal/alectryon-oleifolius-fact.pdf and https://plantnet.rbgsyd.nsw.gov.au/cgi-bin/NSWfl.pl?page=nswfl&lvl=sp&name=Alectryon~oleifolius and https://bie.ala.org.au/species/https://id.biodiversity.org.au/node/apni/2900874

Diplopeltis huegelii https://www.inaturalist.org/taxa/1035979-Diplopeltis-huegelii
Diplopeltis intermedia https://www.inaturalist.org/taxa/1303004-Diplopeltis-intermedia
Diplopeltis petiolaris https://www.inaturalist.org/taxa/1412630-Diplopeltis-petiolaris

Dodonaea adenophora https://www.inaturalist.org/taxa/1353014-Dodonaea-adenophora
Dodonaea amblyophylla https://www.inaturalist.org/taxa/1442734-Dodonaea-amblyophylla
Dodonaea aptera https://www.inaturalist.org/taxa/1242403-Dodonaea-aptera
Dodonaea baueri https://www.inaturalist.org/taxa/928143-Dodonaea-baueri
Dodonaea bursariifolia https://www.inaturalist.org/taxa/1074467-Dodonaea-bursariifolia and https://keys.lucidcentral.org/keys/v3/scotia/key/Plants%20and%20Fungi%20of%20south%20western%20NSW/Media/Html/Dodonaea_bursariifolia.htm
Dodonaea caespitosa https://www.inaturalist.org/taxa/1417556-Dodonaea-caespitosa
Dodonaea ceratocarpa https://www.inaturalist.org/taxa/1004722-Dodonaea-ceratocarpa
Dodonaea concinna https://www.inaturalist.org/taxa/1442510-Dodonaea-concinna
Dodonaea ericoides https://www.inaturalist.org/taxa/1533196-Dodonaea-ericoides
Dodonaea hackettiana https://www.inaturalist.org/taxa/1162883-Dodonaea-hackettiana
Dodonaea hexandra https://www.inaturalist.org/taxa/962235-Dodonaea-hexandra
Dodonaea humifusa https://www.inaturalist.org/taxa/1442735-Dodonaea-humifusa
Dodonaea humilis https://www.inaturalist.org/taxa/958294-Dodonaea-humilis
Dodonaea inaequifolia https://www.inaturalist.org/taxa/958967-Dodonaea-inaequifolia
Dodonaea larreoides https://www.inaturalist.org/taxa/1413464-Dodonaea-larreoides
Dodonaea lobulata https://www.inaturalist.org/taxa/353810-Dodonaea-lobulata
Dodonaea microzyga https://www.inaturalist.org/taxa/940720-Dodonaea-microzyga
Dodonaea pinifolia https://www.inaturalist.org/taxa/1257565-Dodonaea-pinifolia
Dodonaea procumbens https://www.inaturalist.org/taxa/1089543-Dodonaea-procumbens and https://spapps.environment.sa.gov.au/SeedsOfSA/speciesinformation.html?rid=51
Dodonaea ptarmicifolia https://www.inaturalist.org/taxa/1363574-Dodonaea-ptarmicifolia
Dodonaea stenozyga https://www.inaturalist.org/taxa/499433-Dodonaea-stenozyga
Dodonaea viscosa https://www.inaturalist.org/taxa/122711-Dodonaea-viscosa

RESULTS: SOUTHERN AFRICA

Dodonaea viscosa 'angustifolia' https://www.inaturalist.org/taxa/122711-Dodonaea-viscosa

Erythrophysa alata https://www.inaturalist.org/taxa/183498-Erythrophysa-alata and https://pza.sanbi.org/erythrophysa-alata

Pappea capensis https://www.inaturalist.org/taxa/428559-Pappea-capensis and https://treesa.org/pappea-capensis/ and https://pza.sanbi.org/pappea-capensis

DISCUSSION

Alectryon/Pappea:

Alectryon (https://en.wikipedia.org/wiki/Alectryon_(plant)) and Pappea (https://www.inaturalist.org/posts/100103-a-comparison-of-sapindaceae-in-the-mediterranean-and-adjacent-arid-climates-of-australia-and-southern-africa#activity_comment_48b10771-dda9-4950-b8b3-291d52bbc76c) occur in the relevant semi-arid climates.

Both are

However, neither penetrates the mediterranean climate. This is partly owing to their dependence on fire-free regimes.

The Australian species, Alectryon oleifolius (https://keys.lucidcentral.org/keys/v3/scotia/key/Plants%20and%20Fungi%20of%20south%20western%20NSW/Media/Images/Alectryon_oleifolius_ssp._canescens/Alectryon_oleifolius_tree_501242_050910.jpg) differs from its approximate southern African counterpart, Pappea capensis, in

Please see

Diplopeltis:

This relatively obscure genus seems typical of the Western Australian counterpart of coastal fynbos (https://www.jstor.org/stable/2260228) under semi-arid climates. It seems to be myrmecochorous (https://www.publish.csiro.au/BT/BT9750475), and is probably adapted to wildfire.

Dodonaea:

Dodonaea has undergone an evolutionary radiation in mediterranean and adjacent arid Australia, producing various shrubby growth-forms with evergreen foliage. These include

Dodonaea is indigenous also to mediterranean and adjacent arid southern Africa. This is extremely puzzling, biogeographically and in evolutionary terms.

Dodonaea viscosa has somehow found its way across the vast oceanic barrier, probably hundreds of thousands of years ago. It is a fully integrated species in a diverse Cape Flora (https://en.wikipedia.org/wiki/Cape_Floristic_Region) that is otherwise distinct from the corresponding flora in Australia.

This sharing of D. viscosa between the landmasses is neither a 'Gondwana link' (https://en.wikipedia.org/wiki/Gondwana) nor anthropogenic. Instead, it is an unique example of a tall shrub/low tree defying major biogeographical barriers.

Furthermore, the form of D. viscosa in southern Africa seems identical to one ('angustifolia') of the seven forms recorded in the species' original range in Australia as a whole.

This raises a paradox:
Dodonaea viscosa is intraspecifically variable in Australia. However, in southern Africa it has shown no subspeciation/ecotypy/raciality/genetic drift, despite being subject to several glacial/interglacial cycles in the Pleistocene.

One species that presents a somewhat parallel biogeographical puzzle is Metrosideros angustifolia (https://www.inaturalist.org/journal/milewski/81613-metrosideros-angustifolia-myrtaceae-the-pacific-face-of-fynbos#). This anomalous member of Myrtaceae resembles D. viscosa 'angustifolia' in growth- and foliage-form, and is sympatric with it. However, it differs ecologically, by being restricted to waterlogged substrates.

Erythrophysa:

Erythrophysa alata resembles D. viscosa - which is sympatric with it in Namaqualand in Northern Cape province - in being a drought-tolerant evergreen shrub with air-filled, somewhat winged, capsular fruits. However, it differs in the following ways:

Posted on October 13, 2024 04:39 PM by milewski milewski | 16 comments | Leave a comment

October 15, 2024

A plant that seems to defy biogeographical and evolutionary rules: Dodonaea viscosa (Sapindaceae), part 2: wood density

@frontyardscientist @graysquirrel @gregtasney @alan_dandie @prof_coffee @alzym @russellcumming @tonyrebelo @jeremygilmore @arthur_chapman @botaneek @troos @thebeachcomber @mr_fab @adriaan_grobler @alastairpotts @s_k_johnsgard @joshua_tx @peakaytea @lanechaffin @kyle_campbell1 @sedgesrock

...continued from https://www.inaturalist.org/journal/milewski/99976-a-plant-that-seems-to-defy-biogeographical-and-evolutionary-rules-dodonaea-viscosa-sapindaceae-part-1#

INTRODUCTION

We have seen that Dodonaea viscosa is a fast-growing tall shrub or short tree (up to 10 m high, https://books.google.com.au/books/about/Trees_of_Southern_Africa.html?id=jtvZAAAAMAAJ&redir_esc=y), that is capable of colonising bare ground.

Based on this biological profile, D. viscosa might be expected to have light wood, economical for a rapid succesional turnover (https://www.researchgate.net/publication/381698504_Wood_Density_Functional_Trait_in_Plants).

In this Post, I point out the following:

Perhaps the most remarkable - and underappreciated - aspect of the biology of Dodonaea viscosa is the extreme density of its wood.

The exceptional nature of D. viscosa is shown, for example, by a comparison with the many other taxa of trees indigenous to New Zealand (see data at the end of this Post, with D. viscosa marked by ***).

RESULTS

Most trees in New Zealand have wood densities (specific gravity when air-dry) of about

  • 0.6 (gymnosperms), or
  • 0.7 (angiosperms).

The latter value is in line with typical 'hardwoods', e.g. English oak, which has a density of 0.72 (https://en.wikipedia.org/wiki/Quercus_robur).

In order to visualise what the range of values means:
In water, any wood with values < 1.0 floats, whereas any wood with values > 1.0 sinks.

Somewhat exceptional in the flora of New Zealand are the following trees.

Species with unusually light wood are

Species with unusually heavy wood are

Other references state the density for D. viscosa to be

The basic wood density (https://bsapubs.onlinelibrary.wiley.com/doi/10.1002/ajb2.1175#:~:text=While%20wood%20density%20has%20been,biomass%20from%20living%20tree%20volume. and https://www.sciencedirect.com/topics/agricultural-and-biological-sciences/wood-density) of D. viscosa is

For comparison, the basic wood density of English oak (Quercus robur) is 0.646.

DISCUSSION

It is noteworthy that the spp. with the heaviest wood in New Zealand

  • belong to various families (Sapindaceae, Asteraceae, Oleaceae, Ericaceae, Myrtaceae), and
  • are small-bodied enough, in several cases, to be described as shrubs rather than trees.

Furthermore, in the Myrtaceae:
Metrosideros contains both

There is a clear finding that the wood of D. viscosa - despite this species being usually shrubby and short-lived, and functioning as a pioneer in the succession of vegetation - is the heaviest of any plant indigenous to New Zealand.

The wood of D. viscosa is also remarkably consistent worldwide, regardless of climate (https://bsapubs.onlinelibrary.wiley.com/doi/10.3732/ajb.90.4.532).

These findings are all the more puzzling considering the following:
Dodonaea viscosa is the only indigenous plant in New Zealand, capable of growing into a tree, that has a cosmopolitan distribution.

What emerges is the following new perspective on the nature of D. viscosa:
This species has

In D. viscosa, the real specialisation is in:

  • unpredictable intraspecific variation - vaguely described by an 'ochlospecies' concept - that is incongruent with subspeciation, ecotypy, or raciality, and
  • extreme density of wood.

This raises two basic questions about the natural history of D. viscosa, viz.

  • What is the natural lifespan in this species? and
  • How has the extreme lignification of its stems contributed to its unique success in colonising the world during the Pleistocene?

MY COMPILATION OF THE DATA

Reference: Kennedy et al. (2023) Wood density and stiffness of New Zealand native trees and shrubs. New Zealand Journal of Forestry Science 53: 13 (https://nzjforestryscience.nz/index.php/nzjfs/citationstylelanguage/get/acm-sig-proceedings?submissionId=315&publicationId=286&issueId=8).

The values are for density (specific gravity) of air-dry wood, for species of trees indigenous to New Zealand.

Conifers:

Araucariaceae:
Agathis australis 0.468-0.630

Cupressaceae:
Libocedrus bidwillii 0.344-0.416
Libocedrus plumosa 0.380

Podocarpaceae:
Dacrycarpus dacrydioides 0.429-0.492
Dacrydium cupressinum 0.455-0.754
Halocarpus biformis 0.554
Halocarpus kirkii 0.627
Lepidothamnus intermedius 0.766
Manoao colensoi 0.515-0.788
Pectinopitys ferruginea 0.625-0.658
Podocarpus laetus 0.619
Podocarpus totara 0.435-0.559
Prumnopitys taxiflora 0.534-0.787

Phyllocladaceae:
Phyllocladus toatoa 0.675
Phyllocladus trichomanoides 0.553-0.645

Monocotyledons:

Arecaceae:
Rhopalostylis sapida 0.401

Asparagaceae:
Cordyline australis 0.522
Cordyline indivisa 0.626

Dicotyledons:

Acanthaceae
Avicennia marina 0.701

Araliaceae
Meryta sinclairii 0.462
Neopanax arboreus 0.677
colensoi 0.722
Pseudopanax crassifolius 0.757
lessonii 0.674
Raukaua edgerleyi 0.581
simplex 0.576
Schefflera digitata 0.559

Asteraceae:
Brachyglottis repanda 0.681
Olearia furfuracea 0.783
ilicifolia 0.663
lacunosa 0.748
paniculata 1.120
rani 0.850
virgata 0.776

Atherospermataceae:
Laurelia novae-zelandiae 0.382-0.465

Chloranthaceae:
Ascarina lucida 0.500

Coriariaceae:
Coriaria arborea 0.618

Corynocarpaceae:
Corynocarpus laevigatus 0.791

Cunoniaceae:
Ackama rosifolia 0.485
Pterophylla racemosa 0.572-0.639
sylvicola 0.627-0.680

Elaeocarpaceae:
Aristotelia serrata 0.575-0.593
Elaeocarpus dentatus 0.562-0.708
hookerianus 0.514-0.568

Ericaceae:
Archeria traversii 0.688
Dracophyllum latifolium 0.779
traversii 0.615
Leucopogon fasciculatus 0.901

Fabaceae:
Sophora microphylla 0.766-0.804
tetraptera 0.884-0.961

Griseliniaceae:
Griselinia littoralis 0.763
lucida 0.655

Lamiaceae:
Vitex lucens 0.959-0.980

Lauraceae:
Beilschmiedia tarairi 0.591-0.888
tawa 0.637-0.761
Litsea calicaris 0.502-0.603

Loganiaceae:
Geniostoma rupestre 0.562

Malvaceae:
Entelia arborescens 0.137-0.281
Hoheria angustifolia 0.738
lyallii 0.747
populnea 0.793
Plagianthus regius 0.506

Meliaceae:
Dysoxylum spectabile 0.491-0.678

Monimiaceae:
Hedcarya arborea 0.584

Myrtaceae:
Kunzea ericoides about 0.760 (0.731-0.943)
Leptospermum scoparium 1.076
Lophomyrtus bullata 0.694
Metrosideros excelsa 0.915
robusta 0.710-0.915
umbellata about 1.040 (0.968-1.140)
Neomyrtus pedunculata 0.741
Syzygium maire 0.637

Nothofagaceae:
Nothofagus cliffortioides 0.596-0.661
fusca about 0.600 (0.490-0.780)
menziesii 0.419-0.700
solandri 0.609-0.790
truncata 0.615-0.782

Nyctaginaceae:
Ceodes brunoniana 0.226

Oleaceae:
Nestegis cunninghamii 0.790-0.995
lanceolata 0.825
montana 1.013

Onagraceae:
Fuchsia excorticata 0.788

Pararyphiaceae:
Quintinia serrata 0.620

Pennantiaceae:
Pennantia corymbosa 0.526

Piperaceae:
Macropiper excelsum 0.519

Pittosporaceae:
Pittosporum eugenioides 0.796
tenuifolium 0.807-0.955
umbellatum 0.827

Plantaginaceae:
Veronica parviflora 0.768
salicifolia 0.822

Primulaceae:
Myrsine australis 0.805-0.991
salicina 0.844

Proteaceae:
Knightia excelsa 0.721-0.785
Toronia toru 0.836

Rhamnaceae:
Discaria toumatou 0.819

Rousseaceae:
Carpodetus serratus 0.742-0.822

Rubiaceae:
Coprosma arborea 0.654
linariifolia 0.835
lucida 0.674
repens 0.599
robusta 0.664

Rutaceae:
Melicope ternata 0.808

Santalaceae:
Mida salicifolia 0.733

Sapindaceae:
Alectryon excelsus 0.854-0.916
***Dodonaea viscosa 1.179

Scrophulariaceae:
Myoporum laetum 0.630

Strasburgeriaceae:
Ixerba brexioides 0.648-0.675

Violaceae:
Melicytus lanceolatus 0.597
ramiflorus 0.654

Winteraceae:
Pseudowintera axillaris 0.643
colorata 0.645

Posted on October 15, 2024 10:25 PM by milewski milewski | 9 comments | Leave a comment

October 17, 2024