Genetic diversity of introduced weeds

These notes are over two decades old, from a time when I was working on the declaration of weeds under legislation, but they may still be worth a glance. There has been a tension in biosecurity policies between the demand for minimal restraints on trade and a precautionary principle that warns against allowing the entry of any new material of species that have already demonstrated weedy behaviour. The free trade lobby argues that if a weed is already so abundant in Australia that eradication or containment programs have ceased, then restrictions on its entry to the continent (particularly as a contaminant in goods) can no longer be justified. Their position is based on a naïve, pre-Darwinian concept of species as homogeneous entities.

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Introduced populations of a plant may be either more or less uniform than those in its native range.  This can be attributed partly to high or low phenotypic plasticity of the species and the broad or narrow range of habitats in which it is naturalised. Species which can produce a wide range of phenotypes, in adaptation to the range of local environments, are pre-adapted to become weeds.  Naturalised Oxalis pes-caprae is quite uniform in genotype because it has depended on asexual reproduction for its spread in Australia, but it adapts widely in size according to microhabitat and seasonal conditions.

But genetic variation may also increase or decrease with naturalisation, depending on the biology and breeding system of the plant and the type of selection to which it is exposed.

We might expect naturalised plants to show less genetic variation than the same species in its native range, because the whole naturalised population has descended from the few individuals that were its founders.  There has been a “bottleneck” in genetic diversity at the time when the species was transported to its new habitat.

This appears to be demonstrated by Reseda lutea in South Australia. The disjunct distribution of the first records around the coast suggest that there were several successive introductions from overseas. In its native range in Europe and the Mediterranean two subspecies and five varieties are recognised, distinguished by such things as the presence or absence of an aril on the seed, seed size, number and arrangement of ovules, leaf shape and division, and the dissection of the petals. However, collections of this species from throughout its South Australian range show no significant variation in these or other morphological characters (e.g. flower colour, vestiture) which vary in European populations. It is a very uniform species here, suggesting that, even if it was introduced more than once, all the founders of the population came from a similar source.  Reseda lutea was introduced in ships’ ballast around the turn of the century, and our material is a good match for collections from close to the shoreline in Britain and western Europe.

The bottleneck may be beneficial to the introduced species in the longer term, helping it become better adapted after an initial lag phase. Due to the phenomenon of dominance complementation, a period of inbreeding could purge deleterious recessive alleles from a population since these will have an increased chance of appearing in homozygotes than can be eliminated by selection (Barrett & Charlesworth, 1991).

On the other hand, the genetic variance of a plant may increase suddenly as it spreads in a new habitat during the logistic growth phase that follows the lag. Fisher (1930) predicted that mutant alleles, even if they did not increase fitness, were more likely to become established in an expanding population than in a stable one.  Populations of an r-strategist weed invading a new habitat are an extreme case, expanding even more rapidly than the populations considered by Fisher. The pressure of natural selection against mutant alleles is proportionately lower in that situation and genetic variation might be expected to rise.  And such was the case with many of our common annual weeds of broadacre agriculture, which dispersed as quickly as farming land was opened up in the early 19th century and built up large populations in a new vacant niche.

Naturalised populations of salvation Jane (Echium plantagineum) appear to have a high level of genetic variability (Brown & Burdon, 1983), and at all 16 isozyme loci they examined were as diverse as the European source populations (Burdon & Brown, 1986). This may be due to the amount of variation that was present in the original parent plants which may have come from more than one region, or to mutations that had occurred in the 140-odd generations since the species was introduced.

Even when the population has reached a stable size, if the plant has occupied a range of habitats natural selection can be expected to preserve that portion of the new genetic variability which increases fitness in these habitats; this has also been found in salvation Jane which shows clinal variation across south-eastern Australia (Wood & Degabriele, 1985) in morphology as shown by measurements of leaves and flower parts and also in its physiology – in its response to different temperature and moisture regimes.

One reason for the difference in behaviour of Echium and Reseda may be that the former is an annual but Reseda is a perennial and therefore has passed through fewer generations since arriving in Australia with proportionately less opportunity for the establishment of new alleles.

The difference in breeding systems of the two species would tend to reinforce this effect.  Echium is an outbreeder but Reseda is an inbreeder whose flowers self-fertilise if not pollinated by insects, therefore there has been less opportunity for recombination in Reseda. Additionally, Echium exists in populations many orders of magnitude larger than those of Reseda, especially if we count the clonal patches of the latter as unit individuals. Again, Fisher predicted that greater levels of genetic variance are produced and maintained in abundant species than in rare ones.

Another naturalised perennial, Myrsiphyllum asparagoides, also shows much less variation in Australia than in its native southern Africa (Cooke & Robertson, 1990). The stapeliad Orbea variegata is extremely variable in its floral and vegetative morphology across its wide native range (White & Sloane, 1937) but the material naturalised in South Australia is quite uniform in morphology, implying that it was introduced from only a small part of its range, most likely close to Cape Town.

On the other hand, the annual Rumex hypogaeus also introduced from South Africa shows little variation in isozymes across its introduced range in Australia. This may simply reflect the similarly low level of variation in these enzymes its native range (Panetta, 1990).

Weeds and other invasives are species released from fitness constraints and fortuitously adapted to their new home. This adaptation may come about only after hybridisation and recombination between successive introductions from different source populations (Ellstrand & Schierenbeck, 2000). Consequently, it is important that the introduction of additional genotypes of the weed species that are already established here be prevented. Any new addition of germplasm may turn out to be one that can significantly improve the adaptation of the species to local habitats, and so increases its invasiveness and/or impact as a weed.

The same general principles could be applied to other pest organisms of biosecurity concern.

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References

Barrett, S.C.H. & Charlesworth, D. (1991) Effects of a change in the level of inbreeding on genetic load. Nature 352: 522-524.

Brown, A.H.D. & Burdon, J.J. (1983) Multilocus diversity in an outbreeding weed, Echium             plantagineum L. Aust. J. Biol. Sci. 36: 503-510.

Burdon, J.J. & Brown, A.H.D. (1986) Population genetics of Echium plantagineum L. – target weed for biological control. Aust. J. Biol. Sci. 39: 369-378.

Cooke, D.A. & Robertson, M. (1990) Bridal creeper, Myrsiphyllum asparagoides, in South Australia. Proc. 9th Aust. Weeds Conf. Adelaide 113-115.

Ellstrand, N.C. & Schierenbeck, K.A. (2000) Hybridization as a stimulus for the evolution of invasiveness in plants? Proc. Nat. Acad. Sci. 13: 7043-7050.

Fisher, R.A. (1930) The Genetical Theory of Natural Selection. (Oxford University Press).

Panetta, D. (1990) Isozyme variation in Australian and South African populations of Emex australis Steinh. Aust. J. Bot. 38: 161-167.

White, A. & Sloane, B.L. (1937). The Stapelieae. edn 2 (Abbey San Encino: Pasadena).

Wood, H. & Degabriele, R. (1985) Genetic variation and phenotypic plasticity in populations of Paterson’s curse (Echium plantagineum L.) in south-eastern Australia. Aust. J. Bot. 33: 677-686.


Inflorescence cormlets in Watsonia meriana

All corms and cormlets produced by a Watsonia are axillary. Sympodial branching is a diagnostic character of the genus, with the main corm being exhausted by producing an annual flowering stem and one or several new lateral underground corms each subtended by a basal leaf. Additional smaller corms, variously called cormlets, cormils or bulbils are typically produced in the axils of lower cauline leaves in many species including W. aletroides (Burm.f.)Ker Gawl., W. humilis Miller and W. meriana (L.)Miller.

The Watsonia inflorescence is a spike with each solitary, sessile flower subtended by an outer bract and a less robust inner bract. The weedy variety W. meriana var. bulbillifera (J.Mathews & L.Bolus)D.A.Cooke is distinguished by having the lower flowers in its inflorescence replaced by clusters of cormlets. This variety is usually a sterile triploid depending on above-ground cormlets for dispersal, and is known as bulbil watsonia.

However, some genotypes of diploid W. meriana var. meriana can produce small solitary cormlets or cormlet clusters in the inflorescence, at least in cultivation. The extent of cormlet development is variable from one year to another; they are sometimes absent and are never as large or numerous as in var. bulbillifera.

Accession 183. At the right of the picture is a typical solitary cormlet 3mm wide produced in the axil of the reduced leaf subtending an inflorescence branch – but on the outside of that branch. This has been observed on all branches in six out of eight years. To the left is a cluster of six cormlets replacing the lowermost flower on the main axis; this has only been observed once.

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Accession 187. A cluster of three cormlets replacing the lowermost flower of the inflorescence. This has only been observed once in eight years.

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I used to wonder if the cormlet clusters of var. bulbillifera might be homologous to the twelve organs (six tepals, three stamens, three carpels) in a flower. This hypothesis was attractive because, although the number of cormlets in each closely packed cluster is variable, twelve is a typical number. The observations reported here make the hypothesis much less likely. There is a gradient transition from stem cormlets through inflorescence cormlets to clusters. The position of the cormlets in accession 183, axillary to the phyllome subtending an inflorescence branch but abaxial to that branch as if an extra axillary growing point, is rather surprising.

The consistent development of inflorescence cormlet clusters is still the morphological character separating var. bulbillifera from var. meriana. However, the difference should be understood as one of degree rather than a clear dichotomy.

Accession 183 was a gift from Mr Graeme Dallimore who collected it on Mornington Peninsula, Victoria. Accession 187 is said to be Cronin material discarded from Melbourne Botanic Garden in 1995. Both readily produce seed, implying that they are diploids.

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References

Conran, J.G., Wilson, P.A. & Houben, A. (2004). Pollination and ploidy changes in South Australian populations of bulbil watsonia, Watsonia meriana (L.)Miller var. bulbillifera (J.Mathews & L.Bolus)D.A.Cooke (Iridaceae). Herbertia 57: 57-70.

Cooke, D.A. (1998) Bulbil watsonia is a variety of Watsonia meriana (L.)Miller (Iridaceae). J. Adelaide Bot. Gard. 18: 5-7.

Goldblatt, P. & Manning, J.C. (2020) Iridaceae of southern Africa. Strelitzia 42. (South African National Biodiversity Institute: Pretoria).


Herbertia lahue in Australia

Now that Deble (2021) has clarified the nomenclature of this group of related species, it’s clear that the Herbertia commonly grown in Australian gardens is H. lahue (Molina)Goldblatt, which is remote from H. pulchella Sweet.

Its closest relatives include H. amoena Grisebach and H. caerulea (Herbert)Herbert. These three are sympatric in South America, but H. lahue is the commonest in central Chile, and so may be better adapted to grow autumn to winter in a Mediterranean climate where it continues flowering through spring until dryness forces it to go dormant for the summer. The epithet lahue comes from a Chilean common name.

In my experience of growing H. lahue in Adelaide, it is self-compatible or possibly apomictic, with every flower producing a full capsule of seed after opening for one day. Seed germinates readily after sowing in autumn and plants may flower in their second year. Although perennial, the bulbs do not multiply rapidly and may die after a few years. Like many other r-strategist irids, it depends on seed to maintain a population.

The true Herbertia lahue is recognised by the combination of characters:
– a single flower per spathe (instead of two)
– outer tepals 10–18 mm wide
– anthers straight, 5–7 mm long
– apices of the 3 style branches only shortly bifid and straight (not recurved)

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Reference

Deble, L.P. (2021) Herbertia lahue (Iridaceae) and its allies. Balduinia 17: 2-13.


An Iris resembling Iris aphylla

Many rhizomatous bearded irises are loosely called “Iris germanica” in Australia, and I have to admit I was guilty of this in the Flora of Australia (1984) and the Flora of South Australia (1986). Even the complex garden hybrids known as Tall Bearded Irises are sometimes lumped under this binomial, although the real Iris germanica L. is highly unlikely to have contributed to any of them, being difficult to use in a breeding program.

Among them are several distinctive cultivated clones; you may find them in your garden, or your neighbours’ gardens, or growing feral on waste ground from plantings several (human) generations ago. It seems approriate to call them clones as they represent a small number of quite uniform phenotypes with no intermediate forms, and rarely if ever produce seed. One of these irises that is frequent around Adelaide resembles I. aphylla L. at first glance, but can only be included in the broadest concept of this species.

It’s clear that the epithet “aphylla” used by Linnaeus in Sp. Pl. 38 (1753) did not mean that the plant is leafless at flowering time, although that may be the case in the colder parts of its historical range, which extends from the Balkans north to Germany and east to Russia. He actually wrote “scapo nudo longitudine foliorum” – with a naked scape the same length as the leaves – implying that he saw leaves and flowers together, as shown in the habit photo at Plants of the World Online. By “scapo nudo” he surely meant that the scape did not bear any leaves apart from the bracts.

Rhizomatous perennial herb to 70 cm high. Leaves in a basal fan of 5-12, ensiform, glaucous, to 37 mm wide, 50 cm long, apices acute and straight or gently incurved, present throughout the year. Scape elliptic in cross-section, to 50 cm tall, naked apart from a bract subtending each of the 0-2 short branches. Bracts slightly inflated, cymbiform, herbaceous but densely purple-streaked. Flowers solitary or paired within the bracts. Perianth tube to 3 cm long. Perianth lobes obovate, to 6 cm long, indigo to dark violet, the outer series (falls) patterned with a dark network on paler ground in the lower part, with a beard of white hairs, those near the haft yellow-tipped. Anthers 15 mm long, white, on 15 mm dark blue filaments. Style branches indigo, shading to almost white at base. Ovary fusiform, 6-angled, to 15 mm long. Fruit and seeds not seen. Flowers in August in Adelaide, long before I. germanica and the Tall Bearded Irises.

Thus it differs from I. aphylla sens. strict. in the leaves, which are seasonal, fewer, broader, and strongly falcate in that species (Dabrowska et al., 2019).

There are several irises in southern Europe close to I. aphylla and included in it by Service (1997) that have also been given names at species level. They may be of hybrid origin (Colosante & Mathew, 2008) but none of them matches the one described above.

Many of the supposed species in Iris Section Iris could be ancient garden hybrids that have been given binomials as if they were actual species that consist of wild populations. It’s a similar story in many horticultural genera such as Citrus, where our familiar oranges and lemons are ancient hybrids bred from less familiar Asian species. Long before the well-documented hybridisation of irises over the past two centuries (Darlington, 1973) there may have been earlier cycles of hybridization, selection, and the survival of some forms as feral populations that botanists in recent times have interpreted as species. My father used to say that history is a lot longer than most people think, and this is surely true of the history of plant domestication.

I. aphylla sens. strict. is a tetraploid with 4n = 48 chromosomes, while I. albicans Lange and I. germanica are reduced tetraploids with 44. All three behave as obligate outbreeders in cultivation; seed can only be produced by hand pollination with compatible pollen. When documented hybridisation began in the 19th century the first Tall Bearded Iris cultivars were diploids produced from I. variegata L. and I. pallida Lam. with 2n = 24, and the contemporary tetraploids and hexaploids are derived from them.

For comparison, here is a plant matching Iris germanica L. in morphology and pigmentation – although I haven’t seen its chromosomes.

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References

Colasante, M. & Mathew, B. (2008). Species of natural hybrid origin and misinformation in the Irises: A reappraisal of the presence of I. aphylla L. in Italy. Plant Biosystems 142: 172-178.

Dabrowska, A., Smigala, M. Denisow, B. & Winiarczyk, K.(2019) Biology of flowering and insect visitors of Iris aphylla L. (Iridaceae). Turk. J. Bot. 43: 798-808.

Darlington, C.D. (1973) Chromosome Botany and the Origins of Cultivated Plants. 3rd edn (Allen & Unwin: London).

Service, N. (1997) Section Iris. In The Species Group of the British Iris Society A Guide to Species Irises. 17-56. (Cambridge University Press: Cambridge).


Watsonia fulgens

Watsonia fulgens (Andrews)Pers. based on Antholyza fulgens Andrews was regarded as a nomen confusum by Goldblatt (1989) because the type illustration could not be matched to any wild population. Andrews’ description of this plant whch had been introduced to England in 1792 was little more than a diagnosis differentiating it from Antholyza ringens (= Babiana ringens): it had much longer glabrous leaves that remained green until new growth appeared, and bright scarlet, curved trumpet shaped flowers with large spreading lobes.

Ker Gawler (1802) treated it as a distinctive variety of Watsonia iridifolia (Jacq.)Ker Gawl., which is another name of uncertain application. An illustration by Planchon (1856) under W. iridifolia var. fulgens matches a clone that is still widely grown in Melbourne although apparently not commercially available. Planchon noted that it flowered in autumn with a scape to 1-2 metres long, far exceeding the leaves, simple or sometimes branched in vigorous specimens. Plants of this name were being sold in England by 1820 (Loddiges, 1820). In New South Wales, Macarthur (1843) had a plant he called Watsonia iridiflora fulgens and presented material to the Sydney Botanic Gardens in 1831.

The following description is based on accession 180 in my collection:

Evergreen, proliferating, to 150 cm tall. Basal leaves about 4, to 60 cm long, 35 mm wide, bright green with faint glaucous striations and thin green margins. Stem leaves 2, bract-like, slightly inflated. Flowers 24-28 (to 4 open at once) on a brown axis plus 0-2 short branches. Bract acute, to 19 mm long, exceeding the internode, brown-herbaceous. Bracteole subequal, obtuse or notched at apex. Perianth intense orange-red, with a paler star inside throat, alternating red and pale stripes inside tube. Tube to 49 mm long; basal part to 23 mm long; distal part cylindric, curved, to 26 mm long, 8 mm wide at mouth. Ridges absent. Lobes semi-flared with flat margins; outer acute, oblanceolate, to 27 mm long, 11 mm wide; inner elliptic, obtuse, to 28 mm long, 14 mm wide. Stamens closely arcuate with style, anthers 11 mm long, purple with purple pollen. Style branches far exceeding anthers, red with paler stigmas. Capsule cylindric, truncate, to 25 mm long, brown. Seeds with two short wings, 8-10 mm long, dark brown.

Unlike Watsonia tabularis and W. fourcadei, this plant is undamaged by full summer sun in Adelaide as long as it gets enough water. New shoots appear in January while the previous year’s leaves are still green. Flowering is irregular any time from April to September.

There is a superficial resemblance to photos of wild W. zeyheri in colouring: orange-red flowers on a dark axis. But accession 180 is clearly separated from this species by its size, truncate capsules, autumn-spring flowering season, non-thickened leaf margins and the rather characteristic pale star marking in the flowers.

One possible origin could be a garden selection from random hybrids between W. tabularis and W. zeyheri or W. angusta, with strong, hardy growth in cultivation due to F1 vigour. An irregular flowering season is common in Watsonia hybrids between parents with differing phenologies. It also resembles my hybrids of typical W. tabularis pollinated by W. fourcadei in such features as size, flower colour and capsule shape. The four species mentioned in this paragraph are closely related and were treated as the Subsection Angustae in Goldblatt’s revision.

Below is Planchon’s illustration. The prominent leaf venation may be the artist’s interpretation of the striated glaucous bloom emphasising the longitudinal veins.

And the type illustration from Andrews. Assuming it is the same plant as Planchon illustrated, this is less informative. Perhaps it was grown in shaded or otherwise unfavourable conditions, as he described it as only 3 feet tall.

The plant known as Watsonia fulgens has been a “thing” for over 200 years. If it does not match any wild population, perhaps it should be treated as a cultivar. Unfortunately the name has been loosely applied in horticultural literature, for example to W. angusta by Campbell (1986). Watsonia fulgens sensu Montague (1930) was probably a hybrid cultivar; it was described as having pale-rose flowers appearing early in spring. It was distributed by Law Somner (1933) and may have been identical to the Watsonia fulgens described as a deep pink in Brunning’s 1905 and 1918 catalogues.

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References

Andrews, H.C. (1801) Botanist’s Repository 3: t.192.

Brunning, F.H. (1905) Manual of Seeds, Bulbs, Horticultural Sundries. (F.H. Brunning Pty Ltd: Melbourne).

Brunning, F.H. (1918) Winter Flowers, Bulbs, Spring Flowering Sweet Peas. (F.H. Brunning Pty Ltd: Melbourne).

Campbell, E. (1989) Watsonia. In Walters et al. (eds) The European Garden Flora 1: 385-386. (Cambridge University Press: Cambridge).

Goldblatt, P. (1989) The genus Watsonia. (National Botanic Gardens: Kirstenbosch).

Ker Gawler, J.B. (1802) The Botanical Magazine 17: t.600.

Law Somner Pty Ltd (1933) Law Somner Catalogue 1933-34. (Law Somner Pty Ltd: Melbourne).

Loddiges, C.L., Loddiges, G. & Loddiges W. (1820) Catalogue of Plants which are sold by Conrad Loddiges and Sons, nurserymen, at Hackney, near London. (Loddiges: London).

Macarthur, W. (1843) Catalogue of Plants Cultivated at Camden.

Montague, P. (1930) The new watsonias should be freely grown. The Australian Garden Lover 6: 33.

Persoon, C.H. (1805) Synopsis Plantarum 1: 42.

Planchon, J.E. (1856) Flore des Serres et des Jardins de l’Europe. 11: 1.


Almost a belladonna, but not quite

The belladonna lily, Amaryllis belladonna L., is familiar in Australian gardens and also out on roadsides and other public places where it had been planted – or dumped – decades ago. The umbels of large pink flowers appear on unbranched leafless stems at the beginning of autumn.

It has been hybridised with Brunsvigia josephinae (Redouté) Ker Gawl. to produce F1 hybrids that are almost as large and spectacular as the Brunsvigia and are grouped under the name Amarygia tubergenii. They have cartwheel-shaped umbels of many relatively small flowers on longer pedicels.

There are also much more common hybrids, known as Amarygia parkeri (W.Watson)H.E.Moore. Roger Spencer suggested in the Horticultural Flora of South-Eastern Australia that these are actually hybrids with another South African amaryllid, Cybistetes longifolia (L.) Milne-Redh. & Schweick, but was reluctant to complicate the nomenclature further by adopting a new hybrid genus.

An average example of A. parkeri.

It is not easy to distinguish the original belladonna lily from this latter hybrid. The ‘pure’ belladonnas tend to have fewer (less than 13) and larger flowers, which are on even shorter pedicels than in A. parkeri. The various forms of Amarygia parkeri tend to have more flowers in a more symmetrical umbel, and like the Cybistetes they have a conspicuous yellow carotenoid pigment inside the perianth tube.

The acyanic cultivar A. parkeri ‘Hathor’ showing the yellow pigment.


A partial checklist of named Watsonia cultivars

Watsonia is a genus of the Iridaceae with about 53 species in southern Africa. They are perennial herbs growing from corms and producing spikes of showy flowers adapted to pollination by birds or insects. The species are generally interfertile, all being outbreeders with the same diploid chromosome number. Their wide range in size, phenology and flower colour, along with the ease of working with their large simple flowers, make them attractive subjects for collectors and amateur hybridists.

During the early 20th century there was interest in commercial production of named cultivars for home gardens and cut flowers, but the genus has been rather neglected since then. The following checklist is a ‘first pass’ through the referenced publications, with a bias toward those cultivars that have been released in Australia. It is not certain if every cultivar on this list is still extant.

You can download the list as a 210Kb pdf file from this link.

Watsonia ‘Leng’

DOI: 10.13140/RG.2.2.16217.39520


Watsonia coccinea

Watsonia coccinea Herbert ex Baker is native to the South-western Cape in seasonally wet sites on sandy flats.

It has been cultivated in Australia since the 1840s, sometimes under the misapplied name of W. humilis. The earliest recorded importation from the Cape was by Alexander Macleay of Elizabeth Bay via Captain Farquard Campbell in 1838. The specimen illustrated here was purchased from Tesselaars nursery in Victoria in 2002.

coccinea

In cultivation it grows to 40 cm tall, exceptionally to 1 metre but never with more than 12 flowers. The bright scarlet perianth has an arched, narrow cylindrical tube 4-5 cm long marked internally with six darker red lines, and hooded lobes 24 to 28 mm long.

W. coccinea flowers later than many of the winter-growing watsonias, in late October and consequently the flowers are vulnerable to damage by thrips. It is less useful in the garden than the small forms of W. meriana for this reason, and because it is a “shy bloomer” with some full-sized corms producing only foliage if planted too densely or given less than full sunlight during the winter. It has apparently contributed its flower shape and warm colouration to a few of the cultivars bred by Cowlishaw and Cronin in the early 20th century, crossing with larger plants derived from W. borbonica.

References

Goldblatt, P. (1989) The genus Watsonia. 148 pp. (National Botanic Gardens: Kirstenbosch) ISBN 062012517

Macleay, A. (1843) Plants received at Elizabeth Bay. (Ms in Mitchell Library, Sydney, 2009/115).


Watsonia tabularis

Watsonia tabularis J.Mathews & L.Bolus has been in cultivation in Australia since the 19th century but does not appear to have contributed to the pedigrees of any hybrid cultivars bred in this country. It is endemic to the Cape Peninsula of South Africa and may be closest to the more widespread and variable Watsonia fourcadei J.Mathews & L.Bolus.

Plants grown from seed recently imported from South Africa have flowers of pale pink with a darker tube as shown in the photo. These represent the high altitude form; plants from lower altitudes differ in having bright orange flowers.

tabularis

W. tabularis is evergreen, making most growth during autumn and spring then flowering in November to January.

Reference

Goldblatt, P. (1989) The genus Watsonia. 148 pp. (National Botanic Gardens: Kirstenbosch) ISBN 062012517


Two advantages of being inedible

A tweet from Patrick Moore to the effect that most of the pesticides present in the food we eat are produced by the crops themselves set me thinking about the “arms race” between plants to avoid being eaten and at the same time encourage herbivores to eat other competing species.

Eating or being eaten can be viewed as a very simple game. In order to survive, any plant or animal, any living organism, must eat. That is, take in from outside the substances that it needs to grow its own body and to provide energy to run its internal processes. It also must not be eaten if it is going to survive for long. Winning this leg of the game consists of convincing any predator that it cannot be eaten, to use the terminology of  Stephens (1993).

Plants have developed the ‘must eat’ game virtually to its limit by now. Housing symbiotic chloroplasts that fix carbon by photosynthesis, absorbing other nutrient elements, and the metabolic pathways that produce the whole plant have been established since the Palaeozoic. Even the later innovations of CAM and C4 photosynthesis have been around for millions of years.

Dennis Stephens (1994) further suggested that the main game among plants is ‘must not be eaten’ because they have not yet evolved as  far as they can go in that department. They are still in an arms race with herbivores, with pathogens and with each other. Plants may develop spines or other physically deterring outgrowths that convince hungry herbivores that they are not edible. They may use nectar to encourage ants to wander over their surface and clean up feeding insects. But most importantly, they may produce any of a vast range of chemicals (secondary metabolites) that make them bad-tasting or toxic to the particular herbivores that threaten them. These are all physical manifestations of the strategy of being inedible.

As an aside, it’s interesting that the development of toxic secondary metabolites is a speciality of the angiosperms or flowering plants that have dominated land vegetation since the Cretaceous age. Ferns can be inedible too, but they are much less rich in this kind of chemistry. And the notably toxic members of the gymnosperms are not the really ancient ones, but those that have diversified since the Cretaceous in competition with angiosperms – the cycads, Ephedra, Taxus and some related conifers.

So there is an evolutionary pressure on plants to not be eaten by convincing herbivores that they are inedible. The most obvious benefit from this – when it succeeds –  is that they do not lose biomass or get killed outright by herbivory.

But there can be a second advantage for the uneatable. Consider a three-way game between two plants and a herbivore, such as sheep grazing on a pasture of grass containing thistles. A thistle’s spines make it unpalatable; either totally inedible, or so hard for the sheep to eat that it will not be nibbled as long as any edible, palatable grass remains around it.

So the harder the sheep graze, the less the grass will compete with the thistles for light, water and ground space. The combination of grazing pressure and their spiny defence against being eaten has given them a powerful strategy in their own competitive game with the grass.

It’s also interesting to consider the strategy of the grass. It might actually derive some benefit from being grazed along with broadleaf weeds that lack the thistle’s defence, since it is better equipped to regrow after grazing than they are. But that’s another story.

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References

Stephens, D.H. (1993) Expanding on Level 5, Sex. Letter tape of 6 May 1993.

Stephens, D.H. (1994) Postulates, Self and the Obsessive IP. Letter tape of August 1994.