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 Οκτώβριος 13, 2024 0439 ΜΜ by milewski milewski

Σχόλια

Αναρτήθηκε από milewski 9 ημέρες πριν

@tonyrebelo @adriaan_grobler @alastairpotts @botaneek

PAPPEA CAPENSIS

The following aspects of the biology of Pappea capensis are noteworthy.

Firstly, it has a somewhat disjunct distribution, localised in a) Namaqualand sense lato (https://www.inaturalist.org/observations/138622531), and b) the Klein Karroo and Eastern Cape, separately from a wide range in the mesic savannas of East and southern Africa. A particularly odd disjunction is the occurrence of a mainly tropical species into zones combining winter-rainfall and aridity.

Secondly, it incongruently possesses features indicating nutrient-richness, and features indicating nutrient-poverty.

Indicating nutrient-richness are its a) association with dry climates and - in mesic climates - termite mounds, b) hedged growth-form (https://www.inaturalist.org/observations/210093221 and https://www.inaturalist.org/observations/241088122 and https://www.inaturalist.org/observations/232319784 and https://www.inaturalist.org/observations/196917873 and https://www.inaturalist.org/observations/196900564), suggesting palatability to herbivores, c) restriction to fire-free vegetation, and d) complex heterophylly/heteroblasty, affecting the size and shape of leaves, which can revert to a distinctive ('juvenile' in the case of saplings) form after shoots are damaged, and also affecting the form of the shoots (compressed/elongated/epicormic)(https://www.inaturalist.org/observations/130093681 and https://www.inaturalist.org/observations/202707854 and https://www.inaturalist.org/observations/244490793).

Indicating nutrient-poverty are its a) semi-sclerophylly, in which the leaves are somewhat lignified, b) tendency to evergreenness, c) lack of stem-spinescence, d) dull green hue of the foliage (https://www.inaturalist.org/observations/241338081), e) slow growth, longevity, and dense wood, even in mesic climates, and f) affinity for lichens (https://www.inaturalist.org/observations/211682038).

Thirdly, its endozoochorous dispersal involves an unusual form of fleshy fruit (https://www.inaturalist.org/observations/169211993 and https://www.inaturalist.org/observations/168305680). This combines a persistent capsular pericarp (https://www.inaturalist.org/observations/150511973), which starts to dehiscence when still green (https://www.inaturalist.org/observations/104095528 and https://www.inaturalist.org/observations/123370364 and https://www.inaturalist.org/observations/224853478), filled with a succulent food-body (arillode, https://www.inaturalist.org/observations/193818101). For its part, the arillode has an odd origin in the micropyle of the ovule.

The ecological nature of P. capensis can perhaps be summarised by the adjectives 'arid/eutrophic', 'bird-dispersed', 'long-lived', and 'megaherbivore-adapted'.

The incongruence in the quality of the foliage of P. capensis is indicated by the following quote from van Wyk (1974, page 360): "In the eastern Cape province the species is regarded as an exceptionally good fodder plant. Wild animals [in Kruger National Park] are not particularly fond of it, however, and up to the present only elephants, giraffe and impala have been observed eating the young leaves and twigs."

Αναρτήθηκε από milewski 8 ημέρες πριν

Further thoughts on Pappea capensis, stimulated by https://www.inaturalist.org/observations/10324739:

Pappea capensis is ecologically comparable with Sideroxylon inerme, Euclea undulata (https://www.inaturalist.org/taxa/533457-Euclea-undulata), and Schotia afra.

Like those tall shrubs/low trees, it is seldom recorded as seedlings/saplings, despite the obvious mechanisms of dispersal and sowing.

However, the clonal vegetative reproduction in Sideroxylon and Euclea help to explain the persistence of these plants. Is P. capensis known likewise to be clonal, in the sense of suckering (https://www.britannica.com/science/suckering)?

Unlike the other genera mentioned here, Pappea seems to have an induced response in its anti-herbivore strategy. I refer to the heterophylly/heteroblasty, in which damage to shoots results in the replacements having 'juvenile' (serrated) leaves.

Please note that S. afra (https://www.inaturalist.org/taxa/138594-Schotia-afra) resembles P. capensis in its geographical disjunction between a) the lower Orange River, b) the Klein Karroo, and c) Eastern Cape.

Αναρτήθηκε από milewski 7 ημέρες πριν

@tonyrebelo @adriaan_grobler @alastairpotts

An approximate confamilial counterpart for Pappea capensis in Australia is Alectryon diversifolius (https://www.inaturalist.org/observations/213040839 and https://www.inaturalist.org/observations/232835280 and https://www.inaturalist.org/taxa/824439-Alectryon-diversifolius and https://plantnet.rbgsyd.nsw.gov.au/cgi-bin/NSWfl.pl?page=nswfl&lvl=sp&name=Alectryon~diversifolius and https://apps.lucidcentral.org/rainforest/text/entities/alectryon_diversifolius.htm).

This Australian species of Sapindaceae tends to occur in 'vine thickets' (https://en.wikipedia.org/wiki/Rainforests_and_vine_thickets) on relatively nutrient-rich soils, which are exempt from wildfire.

Its fruit (https://www.inaturalist.org/observations/74116500 and https://www.inaturalist.org/observations/74116870 and https://www.inaturalist.org/observations/166922471 and https://www.inaturalist.org/observations/117555052 and https://www.inaturalist.org/observations/112024525) is similar to that of P. capensis, albeit smaller (https://www.inaturalist.org/observations/82277061), but not known as edible for humans.

However, the Australian species is restricted to the subtropics, where rain falls in summer. Unlike P. capensis, it does not extend to the mediterranean and adjacent arid climate. Here it is replaced by a congener, Alectryon oleifolius (https://www.inaturalist.org/taxa/783975-Alectryon-oleifolius), which less resembles P. capensis.

Like P. capensis, A. diversifolius shows remarkable heterophylly/heteroblasty (in size as well as shape) in its foliage. Some leaves have entire, smooth margins (https://www.inaturalist.org/observations/242410461 and https://www.inaturalist.org/observations/236863961), whereas others have toothed margins (https://www.inaturalist.org/observations/239570190).

However, a difference is in degree of sclerophylly (= foliar lignification). The 'juvenile' leaves of A. diversifolius a) qualify as foliar-spinescent, whereas those of P. capensis do not, and b) are more variable in shape (https://www.inaturalist.org/observations/222592401) than those of P. capensis.

Αναρτήθηκε από milewski 7 ημέρες πριν

Dodonaea viscosa extends to the Richtersveld (https://www.inaturalist.org/observations/183183322 and https://www.inaturalist.org/observations/10934687), whereas Erythrophysa alata apparently does not.

Αναρτήθηκε από milewski 7 ημέρες πριν
Αναρτήθηκε από milewski 7 ημέρες πριν

Wow that's one detailed journal post! Thanks for the tag even though I'm only familiar with the species of Sapindaceae native and introduced to Canada

Αναρτήθηκε από frontyardscientist 7 ημέρες πριν

@tonyrebelo @alastairpotts @adriaan_grobler

ALECTRYON OLEIFOLIUS

Reference: pages 117-120 in Burnside D et al. (1995, https://www.publish.csiro.au/RJ/pdf/RJ9950243 and https://library.dpird.wa.gov.au/pubns/2/)

Also see https://www.inaturalist.org/posts/68618-the-puzzle-of-hakea-as-an-indigenous-woody-weed-in-currantbush-mixed-shrubland-in-western-australia#.

The western Gascoyne region (https://en.wikipedia.org/wiki/Gascoyne and https://www.agric.wa.gov.au/rangelands/rangelands-western-australia) of Western Australia is relevant to this Post.

This is despite the subtropical location, and because rain (mean annual 226 mm) falls mainly in winter (https://www.climate.top/australia/carnarvon/graphs/).

The climate here differs from the same latitude in southern Africa (https://en.wikipedia.org/wiki/Maltah%C3%B6he), where the even sparser rain falls in summer instead (https://en.climate-data.org/africa/namibia/hardap-region/maltahoehe-169737/).

Alectryon oleifolius is one of the three most common 'low trees' in 'Currant Bush Mixed Shrub', a vegetation type occurring on the 'less sandy', duplex soils over more than 4000 square kilometres.

This vegetation type "occupies the land systems flanking the erodible river floodplains in the Carnarvon area" (https://en.wikipedia.org/wiki/Carnarvon,_Western_Australia).

The other two of the most common 'low trees' are Acacia xiphophylla (https://www.inaturalist.org/taxa/964520-Acacia-xiphophylla) and Santalum spicatum (https://www.inaturalist.org/taxa/793376-Santalum-spicatum).

"While [Scaevola spinescens, https://www.inaturalist.org/taxa/421592-Scaevola-spinescens] is the most distinctive of the palatable species and is locally dominant within stands in good condition, the general vegetation comprises a rich mixture of both low and tall shrub species".

My commentary:

'Currant Bush Mixed Shrub' vegetation is particularly relevant to this intercontinental comparison. This is because this community, which contains an approximate counterpart for the southern African species Pappea capensis, resembles a southern African rather than an Australian ecological syndrome, in several ways.

I refer to a tendency to a) relative nutrient-richness (alluvial), b) freedom from intense wildfire, c) stem-spinescence (mainly in S. spinosa), d) palatability to herbivores, e) drought-deciduousness in some plants or fleshy foliar texture in others, and f) endozoochorory (with fleshy fruits/'arillate' diaspores in Alectryon, Scaevola, Stylobasium, Solanum, Acacia, Chenopodium, Enchylaena, Atriplex, Rhagodia, Santalum, and Eremophila).

Αναρτήθηκε από milewski 6 ημέρες πριν

Excerpts from the description of Pappea capensis on page 199 in Palmer (1977, https://books.google.com.au/books/about/A_Field_Guide_to_the_Trees_of_Southern_A.html?id=BI0LAQAAIAAJ&redir_esc=y):

"In moist eastern and north eastern areas [of southern Africa] medium-sized, somewhat drooping, luxuriantly leafy, often mistaken for a wild fig, in arid scrub up to 4-5 m, one to several-stemmed, crown somewhat flat...Leaves...up to 8 cm long (arid forms about 3 cm)...often stiff, margins wavy, entire or shortly toothed...Fruit: a 1-3-lobed capsule, usually round, 8-18 mm wide, 'shell' hard, brittle, splitting to show red shiny flesh, shells later brown, persistent"

Αναρτήθηκε από milewski 5 ημέρες πριν

INTRASPECIFIC VARIATION IN PAPPEA CAPENSIS

Reference: Van Wyk (1974, https://www.abebooks.com/9780360001589/Trees-Kruger-National-Park-Volume-0360001580/plp and https://catalogue.nla.gov.au/catalog/1052708 and https://books.google.com.au/books/about/Trees_of_the_Kruger_National_Park.html?id=2DctAQAAIAAJ&redir_esc=y)

Kruger National Park (https://en.wikipedia.org/wiki/Kruger_National_Park) has a summer-rainfall climate, Here, two distinct forms of P. capensis have been called 'lowland' and 'mountain'.

The lowland form, which resembles that common in e.g. Eastern Cape, is restricted in Kruger National Park to the region south of the Tropic of Capricorn.

The mountain form occurs on rocky/sloping terrain over the full latitudinal span of the Park. It differs from the lowland form in being shrubby, multistemmed, and deciduous, with simple, thin branches; the foliage is sparsely distributed on the branches, and consists of exceptionally large (usually 9 cm and up to 17 cm long) leaves (https://www.inaturalist.org/observations/204134534 and https://www.inaturalist.org/observations/199680523).

The lowland form flowers in Sept.-Nov., fruiting in summer; by contrast, the mountain form flowers in Feb.-March and fruits in late winter and early spring.

Αναρτήθηκε από milewski 4 ημέρες πριν

Density of wood (air-dry) is greater in Dodonaea viscosa (1.2-1.25) than in Pappea capensis (0.86). This is despite the fact that the former grows rapidly and acts as a pioneer, whereas the latter grows slowly and does not act as a pioneer.

Αναρτήθηκε από milewski 4 ημέρες πριν
Αναρτήθηκε από milewski 3 ημέρες πριν

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