Invasive American bramble is a thorn in the side of foresters, farmers and land managers across large swaths of South Africa. It chokes up grasslands, forest fringes and river banks, and is notoriously difficult to eradicate. But there is light at the end of the tunnel as a team of scientists are tracking these elusive invaders to find an effective biological control …

American bramble continues to be a major scourge to agriculture, forestry and biodiversity conservation in many of the temperate areas of KwaZulu-Natal, Mpumalanga and Limpopo provinces. The weed forms impenetrable, thorny thickets which impede the passage and access to water of livestock and other animals, replace grazing, smother young plantation trees and make the maintenance and harvesting of older trees difficult. Bramble infestations replace native vegetation, with negative consequences for natural ecosystems, particularly in temperate grasslands. They can also negatively affect specialist flower-visitors. Native bird species increase the spread and germination rates of invasive alien brambles.

There are a number of indigenous bramble species in South Africa, as well as several invasive alien ones. These all belong to the genus Rubus, which falls under the rose family Rosaceae. There are also many species, hybrids and varieties of cultivated Rubus. The most well know of these are the blackberries and raspberries, but they also include youngberries, boysenberries, cloudberries, dewberries and loganberries. There is a small berry industry in South Africa, but most of that sold in our shops is imported – Mexico, for example, is currently one of the main exporters of blackberries worldwide.

The biology of Rubus is somewhat unique, in that most or all species, although perennial, have a biennial flowering and fruiting cycle. “Primocanes” grow from the ground in the first year – long, robust stems which bear no flowers or fruit. In the second year, these become “floricanes”, which bear the flowers and fruit, and subsequently die back. The study of Rubus is also quite specialized, and comes with its own moniker – “batology” – while those who work with Rubus are known as batologists!

The genus Rubus is large and complex, and is characterized by its ability to hybridise. The genus is divided into a number of Subgenera, and within each of these, one or more ‘Sections’. It is widely distributed, with native representatives on six continents, and invasive alien species and hybrids are also widely distributed and cause great harm in certain areas. In South Africa, American bramble is the most damaging of the invasive brambles. Several introductions of various brambles into South Africa were made in the late 19th and early 20th century, chiefly with berry production in mind. By the 1930s, however, American bramble was becoming problematic: an early reference to this bramble as “Rubus cuneifolius” was by E.J. Philips and co-authors in 1939, in “Farming in South Africa”. Rubus cuneifolius is native to Florida and the southern states of the USA, with the common name “sand blackberry”. However, it was soon realized that there was more than one form of this bramble; J.P. Marais, in a 1960 report, divided it into the “Hilton Road variety” (which was shorter, more upright, and grew in more open areas) and the “Richmond variety” (taller, with more arching canes, growing more prominently in more sheltered areas with partial shade).

Jacobus Egberink carried out some of the first comprehensive studies on the weed and its control as part of his MSc in Agriculture through the University of Natal (now UKZN), completed in 1965. Dr Danie Erasmus, based at the Cedara campus of the Plant Protection Research Institute of the national Department of Agriculture (now the Agricultural Research Council’s Plant Health and Protection [ARC-PHP] institute) conducted further work in the 1980s, including on chemical control. Various other studies, on the biology and taxonomy of Rubus in South Africa, were also undertaken in the 1980s by Prof. Charles Stirton, Dr Johan Spies and Henriette du Plessis.

Worldwide, biological control of Rubus species initially achieved low success, mainly because of the complex nature of the genus, in particular its tendency to hybridise, and therefore difficulties in finding natural enemies in the region of origin that were able to develop on the introduced target weeds. With the advent of genetic techniques, success rates have increased. In South Africa, the first attempts towards Rubus biocontrol were undertaken by Dr Mike Morris and colleagues of the Agricultural Research Council’s Plant Protection Research Institute in the 1990s, using plant pathogens (rust fungi). They discovered that one of these (Kuehneola uredines), already widespread in the country, only developed on the upright form of R. cuneifolius, and was not particularly damaging. They then imported another rust fungus (Gymnoconia nitens) from Florida, USA, where it had been collected off R. cuneifolius, into their quarantine laboratory. However, this fungus only infected some specimens of the sprawling form of American bramble, as well as a commercial variety of Rubus and a native species, so it was rejected as a biocontrol agent. Because of the differences in infection patterns between upright and sprawling forms of R. cuneifolius, Dr Morris and his team believed that these might in fact be two separate species; they also realized that the upright form tended to grow at higher altitude than the sprawling form. In the early 2000s, ARC-PHP attempted to initiate genetic work in order to understand origins and identities of the forms of American bramble present in South Africa, in collaboration with Dr Lawrence (Larry) Alice of Western Kentucky University, USA, but this project did not come to fruition.

Recent efforts – from 2018 on

Given the ongoing problems caused by American bramble, interest in undertaking a feasibility study revived in 2018. A small ARC-PHP project (managed by Dr Costas Zachariades) was granted funding by the Department of Environmental Affairs (its Natural Resource Management Programmes directorate, which includes the Working for Water programme). At a similar time, the recently formed Centre for Biological Control, attached to Rhodes University, initiated a project on northern temperate weeds, under the management of Dr Grant Martin. These two units collaborated. An M.Econ. student, Brett Mason, undertook a study looking at some of the costs and benefits of Rubus in South Africa. Coincidentally, in 2017 a young dynamic researcher from the Centre of the Region Haná for Biotechnological and Agricultural Research, Olomouc, in the Czech Republic, Dr Michal Sochor, had started a study on the taxonomy and phylogeny of Rubus in South Africa, in collaboration with Dr John Manning of SANBI. Dr Sochor had previously undertaken research on the taxonomy and phylogeny of European brambles, and was thus highly experienced and knowledgeable; the European approach differs dramatically from the current American approach: European researchers tend to recognise many more species than their North American counterparts – while the latter are “lumpers”, the former are “splitters”, and will describe “microspecies”. This divergent approach has not been consistent; for example, from 1941-1945, Dr L.H. Bailey undertook the most recent comprehensive revision of the genus in North America, and recognized hundreds of species, many of which he described himself. In stark contrast, Dr Larry Alice, in a 2015 article, sank the entire Rubus section Arguti, consisting of about 110 species listed in Bailey’s monograph, under one species, R. pensilvanicus. While the assignment of variable forms to separate species or microspecies may be complex and confusing, the lumping of many variable forms under one species is not helpful for the purposes of identification of invasive forms and determination of their origins.

Dr Sochor and colleagues used several techniques in their work, including extensive field and herbarium studies across South Africa, aimed at clarifying Rubus taxonomy in the region with the help of DNA – ploidy estimation and assessment of reproductive mode. They have subsequently published their findings in two scientific papers: the first, in 2018, deals with Rubus in the Cape Floristic Region, while the second, in 2022, examines the entire country, and is thus more relevant for the purposes of American bramble. They found that the upright form of what had been previously referred to as Rubus cuneifolius is a separate species, and in a separate section of the genus, to the sprawling form. Unfortunately they were unable to put species names to these, and refer to them only as Rubus section Cuneifolii (upright) and Rubus section Arguti (sprawling). Rubus section Cuneifolii is found predominantly in KwaZulu-Natal, while Rubus section Arguti is more widespread, occurring predominantly in KZN, Mpumalanga and Limpopo. Furthermore, the Arguti plants could be divided into two commonly occurring forms. Interestingly, both Rubus section Cuneifolii and Rubus section Arguti are “facultatively apomictic” (meaning that they can reproduce asexually) – apparently this indicates that both of these invasive Rubus are in fact hybrids, not true species; in discussion with Dr Sochor, he felt it was likely that such hybridization had occurred in North America, under natural circumstances, prior to the plants being imported into South Africa. Dr Sochor and colleagues also identified two hybrids of which one parent was Arguti and the other was one of two indigenous Rubus species, but did not find any hybrids of Cuneifolii.

A year prior to the publication of the 2022 paper discussed above, Dr Bram van de Beek, a Dutch theologian who had devoted many years to the study of Rubus in South Africa, published an article focusing on the Cape, although he examined material from across the country, using only taxonomic features (i.e. no ploidy or reproductive methods). Collaborating with Dr Mark Widrlechner, an expert on Rubus taxonomy at the University of Iowa, they identified one of the two sprawling forms (Rubus section Arguti) as Rubus originalis and described the other as a new species, Rubus revealii. For the upright form, they identified a few (“stronger”) plants from KZN as Rubus pascuus but used Rubus probabilis for most plants. Dr Sochor does not feel confident in these identifications; in general we have aligned our work with Dr Sochor rather than Dr van de Beek, but we are also working with Dr Widrlechner in the USA.

In order to familiarize ourselves with the South African Rubus flora (both alien and indigenous), we joined Dr Sochor on one of his fieldtrips to South Africa, in early 2020. Despite our initial confusion as non-botanists, we soon found it quite easy to distinguish between various species based on characteristics such as leaf shape and flower colour. This trip also gave us an opportunity to look for natural enemies present on both alien and indigenous Rubus species. This proved interesting, as we found many more species of insects and pathogens on the indigenous species than the alien ones – although this is expected, it does give an indication that many natural enemies of Rubus are specialized, and secondly that were we to introduce natural enemies from North America onto these alien Rubus plants in South Africa, they have the potential to reduce the invasiveness of these plants.

Current work and the way forward

What is the relevance of the recent taxonomic and phylogenetic studies discussed above to our biocontrol project? The lack of much hybridization, together with the weediness of the plants, led us to restrict our focus to invasive North American Rubus i.e. plants previously falling under “Rubus cuneifolius” in South Africa. In order to progress, we need to firstly understand how many species, and how much genetic variability exists in these species in South Africa. We hope that it allows us to find plants growing in the USA which are close matches to at least some of these invasive Rubus, and thereby find potential biocontrol agents (insects, mites and pathogens) which are compatible with the plants. To achieve the first goal, Dr Sochor agreed to undertake genetic analysis of these species – we therefore undertook a fieldtrip across KZN in early 2023 to collect as much genetic material and herbarium specimens as possible (the latter have been lodged in the Bews Herbarium at UKZN, Pietermaritzburg). This fieldtrip confirmed previous observations that the upright form (Rubus section Cuneifolii) occurs more commonly at high altitude (KZN Drakensberg), while the sprawling form (Rubus section Arguti) occurs more commonly at lower altitude (KZN Midlands). Dr Martin also opportunistically collected some specimens in the USA while on a fieldtrip there for other purposes. Dr Sochor has conducted some analysis of our specimens, and has concluded that while the upright form is genetically quite homogeneous and consistent with a single species/hybrid, the sprawling form in KZN consists of three species/microspecies/hybrids (we still need to sample Rubus section Arguti in other provinces). Furthermore, he did not find a close match between the South African and North American specimens sampled, and the North American samples displayed a high level of variability amongst themselves.

So the identification of North American plants which are genetically close to ours remains a critical step. One way to do this is to obtain genetic material from Rubus herbarium specimens in the USA which are morphologically similar to our invasive ones. Dr Sochor has found that leaf material from herbarium specimens, even those over 100 years old, can yield good DNA. Dr Widrlechner has agreed to assist in obtaining such material, and also in re-examining Rubus specimens of species said to be similar to ours. Bearing in mind that both Rubus section Cuneifolii and Rubus section Arguti in South Africa are hybrids, we may not find a perfect match among herbarium specimens in the US, but we hope this exercise gives us some direction. If so, we can transfer our attention to the field in the US – to areas where these herbarium specimens were originally collected. Again, it would be extremely helpful if a local taxonomist such as Dr Widrlechner could assist us to identify these plants in the field. From there, there are two options, viz. (i) to survey these plants for natural enemies, and to import such natural enemies into quarantine in South Africa; (ii) a better option would be to plant out South African material among genetically similar plants in the USA on which natural enemies are present, and allow these natural enemies to colonise our South African plants on their own. In this way we will be more likely to obtain potential biocontrol agents which are compatible with our plants, and thus more likely to be successful in the field in South Africa, should they prove to have a sufficiently narrow host range (i.e. do not attack native or commercial Rubus) to be safe for release in South Africa. Whether the US biosecurity authorities would permit us to plant out South African Rubus is uncertain, but we plan to apply for permission to do so.

There is a final spoke in the works, and that is a lack of current funding. Funding from DFFE: NRMP became more erratic in 2018, and dried up completely in 2023, with no prospect of revival in the short term. CBC itself has funded some of the work since then, but its means are limited. Adequate funding would allow the US work described above to be undertaken properly, and, should natural enemies be found there that show promise as biocontrol agents, to import these into South African quarantine in order to conduct host-range testing.

Funding notwithstanding, what seemed in the 1980s and 1990s as an intractable, complex situation is now resolving itself into a more manageable research project, with some light at the end of the tunnel due to improved understanding of Rubus taxonomy and phylogeny. It is not inconceivable that within the next 10 years, an effective biocontrol agent could be released for one or more of the invasive North American brambles in South Africa, resulting in reduced vigour and competitiveness of these plants, and correspondingly, more cost-effective management using non-biocontrol methods.

Authors:-
C. Zachariades, Agricultural Research Council’s Plant Protection Research Institute
G. Martin, Centre for Biological Control

Notes from the field
Roger Poole, Member Services Co-ordinator for NCT and Agro-Chemical Liaison Officer for the Timber Industry Pesticide Working Group (TIPWG), provided some useful notes on the chemical control of American bramble …

Bramble is a tricky one due to it having two stages of growth, these being the older stems one always sees and then there is secondary (new) growth you'll find inside the thicket. Timing is critical. Best months to spray are between February through to April as the plant is building up reserves for winter so absorption of herbicide is the most efficient.

The best herbicide is metsulfuron methyl, it is slow acting and gives the best results. If there are other invasive species within the area of treatment, then one can look at glyphosate or the picloram/fluroxypyr formulation.

Glyphosate is not the best but does work on bramble that has been cut down. Depending on the size of the thickets one can apply it with a high pressure unit (bakkie sakkie) or tractor boom sprayer. Knapsacks only work on bramble that a person can walk through so you’ll have to cut the thicket with either a brush cutter or tractor-mounted slasher.

Aerial application has been done previously but water volumes need to be checked and applied as per label due to the need for penetration to ensure the mixture gets through the thicket and results in a good coverage.