Splitting of Lysimachia arvensis colour morphs

Lysimachia is a fairly large genus (few hundred species), but is relatively poorly represented in Australia, with a small handful of naturalised non-native species, and a small handful of native species (although only one, L. japonica, is widely distributed in Australia, with others such as L. fortunei or L. vulgaris var. davurica known from one or very few locations, and are possibly here via waterbird dispersal from outside Australia). One species, L. arvensis, is by far the most widely distributed in Australia and certainly the most well-known as a common weed in disturbed sites such as pastures, wasteland, weedy creeklines, etc. It's found in all Australian states and territories except the NT. Perhaps its most salient feature is the presence of two different flower colour morphs: a bright orange form and a rich blue form.

Many sources, including (until recently) Plants of the World Online - the taxonomic authority we try to adhere to for plants on iNat - give these two forms official varietal names, i.e., Lysimachia arvensis var. arvensis for the orange form, and Lysimachia arvensis var. caerulea for the blue form. Although Australia's state herbaria don't formally recognise these two variety names, they do recognise the presence of both colour morphs in Australia.

Interestingly, and perhaps one of the reasons why the two morphs have been treated as the same species for so long, is that they are not only sympatric, but in many cases directly co-occur. A perfect example is this observation just uploaded today by @russellcumming: https://www.inaturalist.org/observations/166499483

However, as some of you may have noticed, Lysimachia arvensis var. caerulea is no longer on iNat as of this morning; it has been elevated to full species, and is now L. loeflingii.

This change was based on this 2022 paper: https://academic.oup.com/botlinnean/article/199/2/557/6494517. It has also now been reflected in POWO (hence the change now in iNat).

Some relevant parts of the paper:

"Lysimachia arvensis (L.) Manns & Anderb. (synonym: Anagallis arvensis L.) and L. monelli (L.) Manns & Anderb. (synonym: Anagallis monelli L.) are two Mediterranean species that show intraspecific flower colour polymorphism (Ferguson, 1972; Pujadas, 1997). Petal colours are genetically determined (Freyre & Griesbach, 2004; Sánchez-Cabrera et al., 2021), and in both species blue- and red-flowered individuals are found due to the presence of different anthocyanins in their petals (Ishikura, 1981; Quintana et al., 2008; Sánchez-Cabrera et al., 2021). The species differ in ploidy, L. arvensis being tetraploid (revised by Pastor, 1992) and L. monelli being diploid (Kress, 1969; García Pérez et al., 1997).

...Blue- and red-flowered plants may appear in sympatric and allopatric populations, and pollinators show a preference for visiting blue-flowered plants in Mediterranean polymorphic populations (Ortiz et al., 2015) and high colour constancy patterns (Jiménez-López et al., 2020a). Hand-pollination between red and blue individuals gives rise to homogeneous F1 progeny with salmon-coloured flowers (Jiménez-López et al., 2020a), but these are infrequent in wild populations (Jiménez-López et al., 2020c). This ‘hybrid’ phenotype has been described as Anagallis × amoena Heldr. ex Halácsy (de Halácsy, 1904: 11). Nuclear microsatellite markers reconstructed two independent genetic groups for each colour morph, supporting this reproductive isolation between them (Jiménez-López et al., 2020b). All this ecological, morphological, reproductive and molecular evidence suggests that the two colour morphs of L. arvensis are independent lineages.

...Previous phylogenetic studies have scarcely explored the potential implications of corolla colour polymorphism in taxon delimitation in L. arvensis and L. monelli because this trait was considered part of the infraspecific variation. Consequently, only one colour morph per species was sampled in most of the molecular analyses (Martins, Oberprieler & Hellwig., 2003; Manns & Anderberg, 2005, 2007a; Anderberg, Manns & Källersjö., 2007; Yan et al., 2018).

...Lysimachia arvensis is a Mediterranean species currently distributed across the world as an alien species. It is annual, self-compatible (Gibbs & Talavera, 2001) and tetraploid (2x = 40; revised by Pastor, 1992).

...Our phylogenetic results based on nuclear ITS regions indicate that blue- and red-flowered individuals of L. arvensis and L. monelli are independent taxa (Fig. 2), reinforcing ecological, morphological and reproductive evidence.

...Although the presence of different ITS sequences in the red- and blue-flowered individuals of L. arvensis collected from the same populations had already been reported by Manns & Anderberg (2007b), the strong support of our ITS (100 BS, 0.999 PPS) and the species delimitation results (Table 4, Fig. 5) indicate that the two colour morphs of L. arvensis are different lineages and belong to different taxa. The ITS phylogenetic reconstruction is also congruent with recent studies on L. arvensis in which red- and blue-flowered plants were separated in different clades with nuclear microsatellite markers (Jiménez-López et al., 2020b).

...Flower colour constitutes a pivotal evolutionary force to speciation in several groups of plants (Carlson & Holsinger, 2015; Ellis & Field, 2016; Takahashi, Takakura & Kawata, 2016; Narbona et al., 2018), and it has been proposed as a ‘magic trait’, that is a trait ‘encoded by genes subjected to divergent selection that affect pleiotropically reproductive isolation’ (Servedio et al., 2011). However, according to ancestral state reconstruction, flower colour does not seem the trait promoting divergence between lineages of L. arvensis, although it does in L. monelli with posterior polyploidization events (see below). Ancestral state reconstruction of this trait invoked a blue-flowered common ancestor for this Mediterranean Lysimachia, and the transition to red-flowered plants probably occurred only once for the red-flowered common ancestor of red L. arvensis and red L. monelli. This kind of transition from blue to red flowers is quite frequent due to inactivation of a branch of the anthocyanin pathway (Rausher, 2008), and it has been found in L. arvensis lineages (Sánchez-Cabrera et al., 2021). In other plant groups, red-flowered species are usually derived from blue-flowered species (Kay et al., 2005; Wilson et al., 2007; Rausher, 2008; Wessinger & Rausher, 2012). In our study, the blue ancestor probably gave rise to two lineages, one entirely blue (which includes the blue lineage of L. arvensis) and another that subsequently separated into blue- and a red-flowered subclades (the latter including the red lineage of L. arvensis and L. monelli).

...Our results have taxonomic implications for the colour lineages of L. arvensis and L. monelli as each lineage should be defined as different taxa supported by morphological, phylogenetic and geographical data.

...Red-flowered plants of Lysimachia arvensis should maintain this name because Linnaeus in 1753 described the species from red-flowered plants.

...Blue plants of L. arvensis should be called with the specific epithet latifolia as it was the first name employed by Linnaeus in 1753 for plants with blue flowers. However, the epithet latifolia already exists in Lysimachia for a different taxon [Lysimachia latifolia (Hook.) Cholewa in Phytoneuron 28: 1–2 (2014) = Trientalis latifolia Hook., Fl. Bor. Amer. 2(9): 121 (1839), a plant described from Washington]. Therefore, we have selected the name L. loeflingii because Linnaeus in 1753 described blue-flowered plants from materials collected by Loefling in Spain."

So what all this means is that, anything we have been identifying as L. arvensis var. caerulea in Australia is now directly referrable to L. loeflingii. For iNat observations that were already ID'ed to this variety, no further work is needed, as the taxon swap moved them over automatically to the new species. However, in many cases, blue-flowered observations have been ID'ed only to species (which was fine and correct under the old scheme), so these are now incorrectly ID'ed. Similarly, observations that had one ID of L. arvensis and one ID of the blue variety will now have been bumped back to genus due to the now conflicting IDs. I'm going to go through today and correct IDs where I can.

There is one final important thing from the paper, which is the key they provide:

1 . Perennial plants, rarely annual herbs with a single stem; stem nodes with (two) three or four (five) verticillated leaves; fertile nodes with a single flower in the axil of each leaf; flowers with the corolla (14) 16–25 mm in diameter, styles 3–4 mm in length 2

  • Annual plants; nodes generally with opposite leaves, rarely with three verticillated leaves; fertile nodes with two flowers, one in each leaf axil, rarely with a single flower; flowers with corolla 3–12 (14) mm in diameter; styles 1.0–2.5 mm in length 3

2 . Flowers with orange or red corolla Lysimachia collina

  • Flowers with blue corolla Lysimachia monelli

3 . Plants generally compact; internodes generally shorter than the leaves; leaves, at least the upper ones, erect-patent, lanceolate or elliptic-lanceolate, acute; fruit pedicels generally shorter than internodes; flowers with blue corolla, with elliptical lobes, strongly serrated in the upper half of the margin, covered with hairs with (three) four (five) cells, the terminal elliptical or sub-cylindrical, about the size and shape of the adjacent cell, sometimes glabrous Lysimachia foemina

  • Plants generally graceful; internodes often longer than leaves; leaves patent, ovate or ovate-lanceolate, obtuse or subacute; fruit pedicels generally larger than internodes; flowers with blue or orange-red corolla, with lobes broadly ovate, denticulate, with the margin densely covered with hairs with three cells, the terminal globose, larger than the underlying cell 4

4 . Flowers blue 5

  • Flowers orange or red Lysimachia arvensis

5 . Small plants, generally < 10 cm in length; root neck generally covered by secondary roots; ovate leaves; flowers with blue or pale blue corolla, often surpassed by calyx; styles 1.0–1.5 mm in length; Lysimachia talaverae

  • Generally large plants, up to 60 cm in length; bare root neck without secondary roots; ovate-lanceolate leaves, at least in the upper half of the stem; blue corolla, usually longer than the calyx; styles 2.0–2.5 mm in length Lysimachia loeflingii

What is immediately noticeable here is that, when separating L. arvensis and L. loeflingii, the only differentiating character provided in the key is flower colour! This has important implications because many observations in iNat are of non-flowering plants. I'm unsure what to do with all of these; reID them to genus, or leave them as L. arvensis for now? (in my personal experience at least, arvensis is far more common, at the very least in Sydney, compared to loeflingii, so on balance of probability a lot of non-flowering observations are probably of arvensis, but of course this is not a compelling argument)

The start of the paper does mention there are other differences between the orange and blue, namely:
"...colour morphs also differ in other traits such as flowering phenology or type of herkogamy (Arista et al., 2013; Jiménez-López et al., 2020c)". They also note that "The colour morphs show different geographical distributional patterns, blue-flowered plants appearing mainly in drier Mediterranean localities and red-flowered plants being predominant in more temperate areas (Arista et al., 2013)." However, given the co-occurrence of both species in Australia in the same populations (also noted in the paper: "Blue- and red-flowered plants may appear in sympatric and allopatric populations"), this geographical aspect doesn't seem like a helpful separator.

The Arista et al. paper found differences in flowering time in Mediterranean populations. A small selection of text from that paper:

"We found significant negative associations between blue morph frequency and latitude of populations, and between similarity in blue morph frequency and geographical distance of population pairs. This means that a geographical pattern of flower colour exists in L. arvensis, and it seems to be related to climatic features, which suggests that flower colour is not a neutral trait (Mayr 1965). The correlations found between blue morph frequency and the environmental variables studied indicate that blue plants are more frequent in dryer, hotter Mediterranean localities while red plants predominate in more temperate Oceanic areas. This could reflect a differential adequacy of morphs to environmental conditions that is also supported by the fact that red plants in southern mixed populations frequently occupied the wettest or shadiest places (M. Arista & P. L. Ortiz).

...In our experimental study, germination, seedling mass, seedling survival, and flower and ovule production all showed different morph-by-environment interactions. The blue morph showed lower germination in the shade and higher seedling mass in the sun treatment, while the red morph showed lower survival in the dry–sun combination, more flowers in the sun–wet combination and more ovules at sun or wet treatments. Since some treatment effects on the components of plant performance may counteract each other, they are poor predictors of the overall effect when analysed separately. Only by considering overall fitness, instead of each trait separately, enables us to assess how each colour morph is affected by the treatments (García & Ehrlen 2002). Overall male and female fitness of blue morph was markedly higher in dry conditions, and this suggests a better tolerance to more xeric environments. However, our experimental study failed to find a clear pattern of adequacy of red morph to more mesic environments as only in wet-sun but not in wet-shade conditions was overall female fitness higher (male fitness was also higher but not significantly). In fact, the wet–shade combination seems to be the less favourable for L. arvensis as both morphs showed their lowest fitness. Thus, the Mediterranean environment seems to be more suitable for the blue morph, while the red morph seems to perform better in wet and sunny places, such as those where it usually occurs in central Europe. But, it is possible that other environmental factors not considered here could also be responsible for the geographic pattern found in our survey.

...Although most of the traits studied were affected by the experimental treatments, onset of flowering was markedly earlier in the blue morph in relation to the red morph without any morph-by-environment interactions. This difference between morphs, regardless of growing conditions, is one of our most notable results and suggests that this trait is linked to flower colour and is genetically determined. This pattern of flowering can be also observed in natural mixed populations (pers. obs.)

...Taking into account all our results, we found in L. arvensis many monomorphic populations that were spatially isolated, and some mixed populations with observational and experimental evidence of divergence in flowering times between morphs. It is obvious that long-term spatial segregation can generate reproductive isolation and trigger speciation (Mayr 1965; Coyne 1992; Doebeli & Dieckmann 2003). However, even in absence of spatial barriers, differences in flowering time between morphs could cause assortative mating, leading to a decrease in gene flow between them and eventually to allochronic speciation (Fox 2003; Weis et al. 2005; Savolainen et al. 2006; Gavrilets & Vose 2007).

...The marked difference in flowering time between colour morphs leaves open the potential for assortative mating and speciation in L. arvensis"

So maybe these differences could help differentiate in Australia, but I am unsure.

The herkogamy difference (anther–stigma separation) relevant to the Jiménez-López et al. citation above, is unhelpful here as it's a flowering trait.

For now, I'm not going to touch the existing non-flowering observations and leave them IDed as L. arvensis, even though at very least a small subset of them will almost certainly be L. loeflingii. For any non-flowering ones I upload from now on, however, I will only ID to genus. But any with flowers, I will go through now and add IDs where needed.

I will also note that, even if you don't accept this change (which of course is completely fine), L. arvensis var. caerulea is now a synonym on iNat, so it is best to use the new name L. loeflingii; they refer to the same thing on iNat. If things change again in the future, it will be much easier to change Australian observations back with minimal work.

Just for reference, here is the iNat swap: https://www.inaturalist.org/taxon_changes/125737
and here is the original flag asking for the swap: https://www.inaturalist.org/flags/613934

@nicklambert @possumpete @wcornwell @hsauquet @russellbarrett @russellcumming @alx4mtmel @scottwgavins @gregtasney @reiner @bushbandit @mattintas @mftasp @rfoster @ellurasanctuary @margl @terra_australis @jackiemiles @onetapir @gtaseski @aavankampen
Please tag anyone I've missed

Publicado el 10 de junio de 2023 por thebeachcomber thebeachcomber

Comentarios

actually just noting, in cases of non-flowering individuals, the best ID would actually be the section Anagallis, as it contains both species

Publicado por thebeachcomber hace 12 meses

Thanks Thomas. Willl you be updating the reference images for Lysimachia arvensis to exclude examples of the blue-flowering L. loeflingii?

Publicado por bushbandit hace 12 meses

yep will do, thanks for the reminder Michael

Publicado por thebeachcomber hace 12 meses

It seems rather odd to me that 2 species would be together in photos so often eg. https://inaturalist.ala.org.au/observations/137474096

Publicado por scottwgavins hace 12 meses

Oh, and thanks for putting in all that work Thomas!

Publicado por scottwgavins hace 12 meses

Why would it not be called L. caerulea, if the former taxon is var. caerulea...?

I've only seen the two sympatric once...I think the general implication here is that the co-occurrence of the two is a coincidence, and not because the same seeds can produce both forms.

Publicado por silversea_starsong hace 12 meses

no idea why they didn't use that name, perhaps it was also unavailable for some reason

I've seen a decent amount of observations now for Australia with the two co-occurring (at least 20), but out of thousands of observations that I've reviewed, have never seen a plant with both coloured flowers from the same individual

Publicado por thebeachcomber hace 12 meses

also out of interest, I have an observation of a white-flowered individual: https://www.inaturalist.org/observations/139413915
which do I ascribe this to??

Publicado por thebeachcomber hace 12 meses

Interesting essay. I'll look to see if I can see separate plants when there are the two colour morphs. If I find both colour morphs on the one plant, I'll find it hard to agree with the split. But I haven't ever looked close enough before. Thanks for the information.

Publicado por darren_fielder hace 12 meses

If there hadn't been any research into it, I would have thought this was one species that was dimorphic re the flower colour. If it is 2 species then that means each location that has both had seeds or vegetative material from both introduced. I wonder if people introducing them as garden or pot plants and intentionally with both colours could account for much of this?

Publicado por scottwgavins hace 12 meses
Publicado por thebeachcomber hace 12 meses

It mentions a salmon-coloured "hybrid" called L. x amoena. Anyone know what it looks like?

Publicado por silversea_starsong hace 12 meses

@blue_celery if you haven't seen this paper already.

Publicado por silversea_starsong hace 12 meses

I can't find any reliably labelled pics of that hybrid

Publicado por thebeachcomber hace 12 meses

ok I just finished reviewing all 2,500+ Lysimachia observations in Australia and adding IDs where necessary

I guess an important lesson here is that IDing stuff to infraspecific ranks is actually pretty useful just in case they are split off in future; if all the blue-flowered obs had been ID'ed to the variety, it all would have swapped automatically with no work needed. I was certainly guilty of a lot of the IDs to species only, so something to change in future for me too

I also found 3 Modiola caroliniana misIDed as Lysimachia during this, so that was nice to correct those

Publicado por thebeachcomber hace 12 meses

Hah, I remember defending my IDs of the blue form for this exact reference of "well what if they get split one day". But I didn't actually think this one would...

Publicado por silversea_starsong hace 12 meses

I'm thinking that the debated variety "var. lilacina" is what the hybrid "x amoena" is, given it's intermediate in colouring. I chose to taxon swap it to the hybrid name. We do not have many on iNat.

Publicado por silversea_starsong hace 12 meses

Thanks for all that work Thomas!
But if I understand correctly Lysimachia arvensis var. arvensis shouldn't exist on iNat anymore? They should now be moved/taxon swapped up to Lysimachia arvensis ?
Still does though:
https://inaturalist.ala.org.au/observations/96420826
Cheers
Brett

Publicado por ellurasanctuary hace 12 meses

that is currently under debate; see last few comments at https://www.inaturalist.org/flags/528655
(arvensis and caerulea weren't the only two varieties)

Publicado por thebeachcomber hace 12 meses

Thanks Thomas

Publicado por margl hace 12 meses

There is some fancy looking L. arvensis close to me that has white blotches on its petals. See pic 7 on http://alphitonia.com/ViewSpeciesE.cshtml?id=283

Publicado por scottwgavins hace 12 meses

looks like a disease or similar?

Publicado por thebeachcomber hace 12 meses

I was thinking that it might be a variegated cultivar from a time when they might have been sold from nurseries. I'll watch for more to see if they regularly produce flowers like this.

Publicado por scottwgavins hace 12 meses

If it still does not exist, I would recommend the creation of a complex camprising both red and the two blue species

Publicado por blue_celery hace 12 meses

Thank you for all this information!

Although functionally, your key is dichotomous, you've numbered all the lead-pairs as 1. Would you change some to 2, 3, etc., as appropriate? The key would be a lot less confusing that way.

In my area, I'll continue to label non-flowering plants as L. arvensis because the blue morph is vanishingly rare. (I've heard of one sighting and I've never seen it.) In others areas, caution could be appropriate.

Publicado por sedgequeen hace 12 meses

@sedgequeen Barbara, the blue morph is very very common in Western Australia, the orange morph is rarer here.

Publicado por margl hace 12 meses

@thebeachcomber dichotomy 2 could be better:
A: flowers with blue corolla, usually strongly serrated in the upper half of the margin, on margin with few hairs with (three) four (five) cells, the terminal elliptical or sub-cylindrical, about the size and shape of the adjacent cell, sometimes glabrous Lysimachia foemina
B: flowers with blue or orange-red corolla, denticulate, with the margin densely covered with hairs with three cells, the terminal globose, larger than the underlying cell

Characters such as habitus and leaves, in my opinion, are hard to be used with the annual species of the complex. I am afraid that someone could consider also L. foemina "graceful".
Also the shape of petals could be subjected to various intepretations. So. I think it is better to keep the quantitative characters (e.g. number of hairs cells) or easily verifiable qualitative ones (colour of petals).

Publicado por blue_celery hace 12 meses

@sedgequeen thanks, I had the numbers correct, but annoyingly the journal changed the formatting and turned them all to 1s when I published it (due to the way I had inset bullet points, it thought I had multiple lists), fixed now

@blue_celery this is the key directly from the paper

Publicado por thebeachcomber hace 11 meses

Thanks Thomas,
I was really interested to see how the ID updates many of us made over the last 24 hours effected the compare function and the auto ID suggestions. Last night (AEST) no changes to either, the only suggestions were L. arvensis but by this afternoon the blue loeflingii flowers are getting the thumbs up across WA, SA, Vic and NSW in the compare function. A good result but how many people use compare? The only related species offered in the automatic ID is still arvensis. This surely needs remedying given that this is a common plant across much of the world.

Publicado por oneanttofew hace 11 meses

L. loeflingii will be offered when the next CV model comes out, it updates every few months based on a schedule

Publicado por thebeachcomber hace 11 meses

@thebeachcomber that may be the key directly from the paper, but it never hurts to eliminate subjective characters like "graceful" where possible.

Publicado por silversea_starsong hace 11 meses

@thebeachcomber I know it is the key but I can express my point of view

Publicado por blue_celery hace 11 meses

of course, I was just clarifying that it wasn't my construction

Publicado por thebeachcomber hace 11 meses

What do you make of these plants with purple (not blue) flowers in this observation? https://www.inaturalist.org/observations/167110658

Publicado por mattintas hace 11 meses

@mattintas I suspect that could be what has been called "f. lilacina", but there's some uncertainty on how purple or pink f. lilacina ("lilac") and f. carnea ("deep peach to purple") are. I imagine one or both of these forms are what would now be referred to as the hybrid L. x amoena.

Publicado por silversea_starsong hace 11 meses

Challenge accepted...when I'm weeding now I have to search for both colours on the one plant. I would have said I've known in simply a weeding the garden (not academic) sense that the two colours were both around for 50 years or so, and grew together. I've not noticed any timing difference in the flower colours, but I wasn't looking for it either. I am in my sixties. I have lived in Melbourne as a child and gardened with my mother and had my own gardens in the Western District of Victoria for the past 40 years.

Publicado por elizabethhatfield hace 11 meses

Sorry. I meant the blue form is vanishingly rare in Oregon, U.S.A. Australia is a different place!

Publicado por sedgequeen hace 11 meses

I have tried distinguishing herbarium specimens of arvensis (orange) and loeflingii (blue). I have not been able to find any morphological differences. Both are glabrous plants, with entire, sessile leaves of the same size, calyx same size and shape; pedicels in both cases nodding and same length; corolla size same; style same length; even the seeds seem identical. This makes identification impossible for any specimen that does not bear flowers, and even those where flowers are present but the collector has not stated the colour (happens surprisingly often). It makes me wonder about the definition of a species - it seems we are heading down the road where anything that has a different gene sequence is a separate species, regardless of morphology.

Publicado por bean_ar hace 11 meses

I agree with blue_celery that we need a complex for this pair of species. The non-flowering forms cannot be distinguished and they are closely related.

Publicado por sedgequeen hace 11 meses

This taxonomic change has only just come to me attention. I have read this posting and the original paper. Very interesting.
I live in Texas and we have both Lysimachia arvensis' and *Lysimachia loeflingii growing here as well as L. foemina (very occasional).
I was wondering what, if anything, has been decided about the issue of these being impossible to distinguish from each other in the non-blooming state. I would like to embark on making sure all the observations in my area of southeast Texas (at least the ones I have contributed IDs to) are amended to reflect the new taxonomy. I have not found any sort of 'complex' designation that could be used.
Should I just use the genus designation and give a short explanation as to why I am withdrawing my previous species ID?

Publicado por sbdplantgal hace 2 meses

@sbdplantgal I am hoping that if a complex name is created, someone will let us know. It is very important for management in areas where there are likely to be a native species of Lysimachia and a weedy species like the subjects are introduced. All the information about them is obscure until flowering observations are made. In a relatively healthy area of vegetation that is also remote, observations may not be common and a complex like this can get established before it it is recognised as invasive making it more difficult to control. As a naturalised complex in Western Australia it doesn't seem to be controlled as a target except in areas where hand weeding is done, but it probably competes with many of our small annuals and small geophytes which in some vegetation, can make up a significant or even dominant proportion of a highly diverse flora. Some of our most famous tourist destinations are rich fields of annuals and one or the other of these species is most definitely in those areas. If the State Herbarium was to accept this split the data would not distinguish between the native Lysimachia and the introduced except that the naturalised taxon apparently hasn't reached the habitat of the native taxon which is tropical monsoon, so can be distinguished by location.. This region is not well collected outside of the dry season as access can be very limited in the wet and while it is drying out. Most travel in the area is in the dry winter, hopefully it is unlikely that these Lysimachia will establish there but if it did it may be difficult for data to show it has. Both naturalised taxa have been observed in an area which has rainforest in Queensland which may suggest that it is only a matter of time in the Kimberley, WA.

Publicado por margl hace 2 meses

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