Invasive <Emphasis Type="Italic">Tamarix</Emphasis> (Tamaricaceae) in South Africa: current research and the potential for biological control

Most species of Tamarix originate in Eurasia and at least five species have become invasive around the world, including South Africa. However, T. usneoides is indigenous to southern Africa, where the potential for biological control of the invasive species is being investigated. Recent research on the invasive species is reviewed here with particular reference to these South African biocontrol efforts. The successful biological control programme against invasive Tamarix in the USA, using several species of “Tamarisk beetle”, is being used as a guide for the South African research. The South African programme is complicated by firstly, the presence of the indigenous T. usneoides which raises the precision of host-specificity required, and secondly, the introduced and indigenous Tamarix have a high intrinsic value for phytoremediation of mine tailings dams in South Africa. The phylogenetic proximity of these Tamarix species to each other has contributed to this challenge, which has nevertheless been successfully addressed by molecular techniques used to separate the species. In addition, classical morphological techniques have been used to separate the Tamarisk beetles, so that now they can generally be matched to Tamarix tree species. Overall, it is concluded that given the broad knowledge now available on the ecology and identity of both the trees and their biocontrol agents, the prospects for successful biological control of Tamarix in South Africa are good.     

A synoptic review of <Emphasis Type="Italic">Tamarix</Emphasis> biocontrol in North America: tracking success in the midst of controversy

Woody shrubs in the genus Tamarix L. (Tamaricaceae) were introduced into western North America in the nineteenth century and have invaded riparian areas, acting as drivers of ecosystem change by altering fire cycles, soil chemistry, hydrology and native plant composition. The scope and severity of the invasions provided impetus for a classical weed biological control program using Diorhabda spp. (Coleoptera: Chrysomelidae). Since the first releases in 2001 Diorhabda spp. have moved into many of the areas dominated by Tamarix resulting in defoliations, canopy dieback, and in some locations substantial Tamarix mortality. Success of the program has been overshadowed by concern that Tamarix is used by a federally-listed bird sub-species, the southwestern willow flycatcher. The controversy has led to lawsuits, cancelled biological control research and release permits and to a negative perception of Tamarix biocontrol by some. Long term success is likely, but only with continued monitoring and riparian restoration will the program reach its full potential.     

<Emphasis Type="Italic">Tamarix</Emphasis> (Tamaricaceae) hybrids: the dominant invasive genotype in southern Africa

Hybridization is regarded as a rapid mechanism for increasing genetic variation that can potentially enhance invasiveness. Tamarix hybrids appear to be the dominant genotypes in their invasions. Exotic Tamarix are declared invasive in South Africa and the exotic T. chinensis and T. ramosissima are known to hybridize between themselves, and with the indigenous T. usneoides. However, until now, it was not known which species or hybrid is the most prevalent in the invasion. With a biocontrol programme being considered as a way of suppressing the alien Tamarix populations, it is important to document the population genetic dynamics of all species in the region. This investigation sought to identify Tamarix species in southern Africa and their hybrids, describe their population structure, and reveal the geographic origin of the invasive species. To achieve this, nuclear Internal Transcribed Spacer (ITS) sequence data and the multilocus Amplified Fragment Length Polymorphisms (AFLPs) markers were used. Phylogenetic analysis and population genetic structure confirmed the presence of three species in South Africa (T. chinensis, T. ramosissima and T. usneoides) with their hybrids. The indigenous T. usneoides is clearly genetically distant from the alien species T. chinensis and T. ramosissima. Interestingly, the Tamarix infestation in South Africa is dominated (64.7 %) by hybrids between T. chinensis and T. ramosissima. The exotic species match their counterparts from their places of origin in Eurasia, as well as those forming part of the invasion in the US.     

Unresolved native range taxonomy complicates inferences in invasion ecology: <Emphasis Type="Italic">Acacia dealbata</Emphasis> Link as an example

Elaborate and expensive endeavours are underway worldwide to understand and manage biological invasions. However, the success of such efforts can be jeopardised due to taxonomic uncertainty. We highlight how unresolved native range taxonomy can complicate inferences in invasion ecology using the invasive tree Acacia dealbata in South Africa as an example. Acacia dealbata is thought to comprise two subspecies based on morphological characteristics and environmental requirements within its native range in Australia: ssp. dealbata and spp. subalpina. Biological control is the most promising option for managing invasive A. dealbata populations in South Africa, but it remains unknown which genetic/taxonomic entities are present in the country. Resolving this question is crucial for selecting appropriate biological control agents and for identifying areas with the highest invasion risk. We used species distribution models (SDMs) and phylogeographic approaches to address this issue. The ability of subspecies-specific and overall species SDMs to predict occurrences in South Africa was also explored. Furthermore, as non-overlapping bioclimatic niches between the two taxonomic entities may translate into evolutionary distinctiveness, we also tested genetic distances between the entities using DNA sequencing data and network analysis. Both approaches were unable to differentiate the two putative subspecies of A. dealbata. However, the SDM approach revealed a potential niche shift in the non-native range, and DNA sequencing results suggested repeated introductions of different native provenances into South Africa. Our findings provide important information for ongoing biological control attempts and highlight the importance of resolving taxonomic uncertainties in invasion ecology.     

Lack of human-assisted dispersal means <Emphasis Type="Italic">Pueraria montana</Emphasis><Emphasis Type="Italic">var. lobata</Emphasis> (kudzu vine) could still be eradicated from South Africa

The legume, Pueraria montana var. lobata (kudzu vine) is one of the worst plant invaders globally. Here we present the first study of P. montana in South Africa. We found only seven P. montana populations covering an estimated condensed area of 74 hectares during the height of the growing season. Based on a species distribution model, it appears that large parts of the globe are suitable, including parts of the eastern escarpment of South Africa (where most populations occur). South African populations of P. montana appear to have a similar ecology to populations in the USA: high growth rates, low seed germination, no natural long-distance dispersal, little herbivory and vigorous post-fire resprouting. In contrast to the USA, most South African populations do flower and flowers are capable of producing seed in the absence of pollinators. However, P. montana appears to have never been widely planted in South Africa, and the incursion was for many years restricted to a single introduction site. The comparison between the invasions of P. montana in the USA and South Africa highlights the often overriding importance of human-assisted dispersal and cultivation in creating widespread invasions, and should serve as a warning to people who have proposed to utilize the species in Africa.     

Spatial modeling improves understanding patterns of invasive species defoliation by a biocontrol herbivore

Spatial modeling has proven to be useful in understanding the drivers of plant populations in the field of ecology, but has yet to be applied to understanding variation in biocontrol impact. In this study, we employ multi-scale analysis (Moran’s Eigenvector Maps) to better understand the variation in tree canopy exposed to defoliation by a biocontrol beetle (Diorhabda spp.). The control of the exotic tree Tamarix in riparian areas has long been a priority for land managers and ecologists in the American southwest. Diorhabda spp. was introduced as a bio-control agent beginning in 2001 and has since become an inseparable part of Tamarix-dominated river systems in the southwest. Between 2013 and 2016 tamarisk dieback was assessed at 79 sites across Grand County, Utah, arguably the epicenter of Diorhabda impact in the U.S. Canopy cover of Tamarix was between 73 and 81% at these sites, with the percent that was live cover fluctuating by year with a minimum of 42%. Using a traditional general linear model, we found that readily and commonly measured environmental factors could explain only up to 26% of the variation in Tamarix live canopy each year. The number of defoliations was correlated with an increase rather than a decrease in percent live canopy, suggesting compensatory growth. Spatial structure alone explained 22–40% of variation. We found fine scale spatial structure at less than 10 km and broad scale spatial structure from 10 to 30 km. Combining both traditional and novel spatial statistical methods we increased that percentage to 43–63%, depending on year. These results suggest that scientists and land managers must look beyond commonly measured environmental variables to explain non-random biocontrol impact in this system. In particular, this study points to the potential for biotic interactions and variation in flood cycles for further exploration of the identified spatial structure.     

Evaluating the efficacy of <Emphasis Type="Italic">Hypogeococcus</Emphasis> sp. as a biological control agent of the cactaceous weed <Emphasis Type="Italic">Cereus jamacaru</Emphasis> in South Africa

We evaluated the efficacy of Hypogeococcus sp. (Hemiptera: Pseudococcidae) as a biological control agent of the cactaceous weed Cereus jamacaru De Candolle (Queen of the Night cactus) in South Africa. This weed has been described as being under complete biological control due to the action of Hypogeococcus sp., although no formal post-release evaluation had been conducted prior to this study. Biological control was associated with significant reductions in fruiting, plant survival and plant densities, while plant population age structures were negatively affected. Weed populations infected by Hypogeococcus sp. were typified by low or non-existent recruitment and are expected to diminish with time. Populations where Hypogeococcus sp. was absent displayed extensive recruitment, and are predicted to expand or self-replace, if left unchecked. These data indicate that Hypogeococcus sp. has a significant negative effect on C. jamacaru at the individual plant and population level, and given sufficient time provides complete biological control over this weed in South Africa.     

Gram-Positive Bacteria with Probiotic Potential for the <Emphasis Type="Italic">Apis mellifera</Emphasis> L. Honey Bee: The Experience in the Northwest of Argentina

Apis mellifera L. is one of the most important natural pollinators of significant crops and flowers around the world. It can be affected by different types of illnesses: american foulbrood, nosemosis, varroasis, viruses, among others. Such infections mainly cause a reduction in honey production and in extreme situations, the death of the colony. Argentina is the world’s second largest honey exporter and the third largest honey producer, after China and Turkey. Given both the prominence of the honey bee in nature and the economic importance of apiculture in Argentina and the world, it is crucial to develop efficient and sustainable strategies to control honey bee diseases and to improve bee colony health. Gram-positive bacteria, such as lactic acid bacteria, mainly Lactobacillus, and Bacillus spp. are promising options. In the Northwest of Argentina, several Lactobacillus and Bacillus strains from the honey bee gut and honey were isolated by our research group and characterized by using in vitro tests. Two strains were selected because of their potential probiotic properties: Lactobacillus johnsonii CRL1647 and Bacillus subtilis subsp. subtilis Mori2. Under independent trials with both experimental and commercial hives, it was determined that each strain was able to elicit probiotic effects on bee colonies reared in the northwestern region of Argentina. One result was the increase in egg-laying by the queen which therefore produced an increase in bee number and, consequently, a higher honey yield. Moreover, the beneficial bacteria reduced the incidence of two important bee diseases: nosemosis and varroosis. These results are promising and extend the horizon of probiotic bacteria to the insect world, serving beekeepers worldwide as a natural tool that they can administer as is, or combine with other disease-controlling methods.     

Impact of multiple natural enemies on immature <Emphasis Type="Italic">Drosophila suzukii</Emphasis> in strawberries and blueberries

Drosophila suzukii (Matsumura) (Diptera: Drosophilidae) oviposits in ripening fruit, larvae render crops unmarketable, and significant economic losses can occur. Biological control research has focused on individual natural enemy species against immature D. suzukii. Here we combine two predators and an entomopathogenic nematode, expecting species complementarity and increased control of D. suzukii. In strawberries, Orius insidiosus (Say) (Hemiptera: Anthocoridae) plus Heterorhabditis bacteriophora Poinar (Rhabditida: Heterorhabditidae) resulted in fewest D. suzukii (81% reduction), and in blueberries, results were similar (60% reduction), although H. bacteriophora was not as effective as in strawberries, which was likely due to drier substrate conditions. There was neither strong complementarity nor interference between predators, O. insidiosus and Dalotia coriaria Kraatz (Coleoptera: Staphylinidae). Inclusion of O. insidiosus resulted in 50% fewer D. suzukii than combinations without O. insidiosus. Control of D. suzukii can be improved with multiple natural enemies, and combinations of O. insidiosus with other agents (parasitoids, fungal entomopathogens) should be tested.     

Basidiospores of <Emphasis Type="Italic">Puccinia striiformis</Emphasis> f. sp. <Emphasis Type="Italic">tritici</Emphasis> succeed to infect barberry,while Urediniospores are blocked by non-host resistance

Stripe rust (Yellow rust) caused by Puccinia striiformis f. sp. tritici (Pst) is a major disease of wheat worldwide. The use of resistant cultivars to control Pst has been very effective, low-cost, and ecologically sound. However, virulence patterns of Pst can quickly change, which may render resistant cultivars susceptible. The discovery of infection of Berberis spp. by basidiospores of Pst in 2010 raised important concerns about the evolution of new virulent races of the pathogen. Little is known about the infection process of Berberis spp. by basidiospores of Pst and the interaction between Berberis spp. and asexual urediniospores. In this study, the interaction between Pst urediniospores and Berberis spp. was investigated at histological and cytological levels. Our results indicate that Berberis spp. expresses a continuum of layered defenses comprised of structural and chemical changes in the cell wall as well as post-haustorial hypersensitive responses to urediniospore infection. Our study also re-examines in detail the infection process of Pst basidiospores on Berberis spp. and provides useful information for further research on the molecular mechanisms governing the interaction between Berberis spp. and Pst.     

<Emphasis Type="Italic">Aeromonas</Emphasis> and <Emphasis Type="Italic">Plesiomonas</Emphasis> species from scarlet ibis (<Emphasis Type="Italic">Eudocimus ruber</Emphasis>) and their environment: monitoring antimicrobial susceptibility and virulence

The present study aimed at evaluating the role of captive scarlet ibises (Eudocimus ruber) and their environment as reservoirs of Aeromonas spp. and Plesiomonas spp., and analyzing the in vitro antimicrobial susceptibility and virulence of the recovered bacterial isolates. Thus, non-lactose and weak-lactose fermenting, oxidase positive Gram-negative bacilli were recovered from cloacal samples (n = 30) of scarlet ibises kept in a conservational facility and from water samples (n = 30) from their environment. Then, the antimicrobial susceptibility, hemolytic activity and biofilm production of the recovered Aeromonas spp. and Plesiomonas shigelloides strains were assessed. In addition, the virulence-associated genes of Aeromonas spp. were detected. Ten Aeromonas veronii bv. sobria, 2 Aeromonas hydrophila complex and 10 P. shigelloides were recovered. Intermediate susceptibility to piperacillin-tazobactam and cefepime was observed in 2 Aeromonas spp. and 1 P. shigelloides, respectively, and resistance to gentamicin was observed in 4 P. shigelloides. The automated susceptibility analysis revealed resistance to piperacillin-tazobactam and meropenem among Aeromonas spp. and intermediate susceptibility to gentamicin among P. shigelloides. All Aeromonas isolates presented hemolytic activity, while 3 P. shigelloides were non-hemolytic. All Aeromonas spp. and 3/10 P. shigelloides were biofilm-producers, at 28 °C, while 10 Aeromonas spp. and 6/10 P. shigelloides produced biofilms, at 37 °C. The most prevalent virulence genes of Aeromonas spp. were asa1 and ascV. Scarlet ibises and their environment harbour potentially pathogenic bacteria, thus requiring monitoring and measures to prevent contamination of humans and other animals.     

Reproduction and potential range expansion of walnut twig beetle across the Juglandaceae

Biological invasions by insects that vector plant pathogens have altered the composition of natural and urban forests. Thousand cankers disease is a new, recent example and is caused by the complex of walnut twig beetle, Pityophthorus juglandis, and the fungus, Geosmithia morbida, on susceptible hosts, notably some Juglans spp. and Pterocarya spp. Host colonization by P. juglandis may be particularly important for disease development, but the beetle’s host range is not known. In the United States and Italy, this insect has expanded its geographic range by colonizing naïve hosts. The objective of this study was to characterize limits to, and variation within, the host range of P. juglandis and infer the extent to which hosts might constrain the geographic distribution of the insect. We examined colonization and reproduction by P. juglandis in no-choice laboratory experiments with 11 Juglans spp., one Pterocarya sp., and two Carya spp. over 2 years and found that all but the Carya spp. were hosts. Reproduction was generally greater on Juglans californica, J. hindsii, and J. nigra, than on J. ailantifolia, J. cathayensis, J. cinerea, J. major, J. mandshurica, J. microcarpa, or J. regia. Escape of an insect vector into populations of evolutionary-naïve hosts can facilitate rapid range expansion by the pest and massive mortality to hosts. Multi-continental plantings of suitable species may facilitate geographic range expansion of P. juglandis and place other, native Juglans spp. at risk.     

Control and consequences of <Emphasis Type="Italic">Spartina</Emphasis> spp. invasions with focus upon San Francisco Bay

Maritime Spartina spp. are powerful ecosystem engineers that accrete sediment, define shorelines, create habitat, and generate prodigious primary productivity both where they are native and where they have been introduced. Invasive Spartina spp. can compete vigorously with native species, diminish biota, change hydrology, and confound human uses of estuaries. Herbicides have been effective in controlling several Spartina spp. invasions. One of the most recent successes is a 15-year campaign that has virtually eliminated S. alterniflora from the large, century-old invasion in Willapa Bay, WA, USA. Hybridization between native and introduced Spartina spp. has created new species and hybrid swarms. In San Francisco Bay, CA, USA (SF Bay) a complicated situation continues to play out from the purposeful introduction of S. alterniflora, which hybridized with native California cordgrass, S. foliosa. The hybrids spread rapidly and led to a long list of environmental problems, which led to an herbicide program that was successful in greatly diminishing the hybrid and saving the open mud habitat of migratory shorebirds. However, it was belatedly realized that the non-migratory, endangered Ridgeway’s rail uses the tall, dense hybrid Spartina as a surrogate for habitat that was lost during the twentieth century to urbanization and agricultural transformation of marshes around SF Bay. This realization has made difficult the simultaneous management of hybrid Spartina, wildlife conservation, and marsh restoration in San Francisco Bay. Restoration of native vegetation could satisfy the multiple goals of preserving open mud and conserving Ridgeway’s rail.     

Taxonomy and pathogenicity of <Emphasis Type="Italic">Leptographium</Emphasis> species associated with <Emphasis Type="Italic">Ips subelongatus</Emphasis> infestations of <Emphasis Type="Italic">Larix</Emphasis> spp. in northern China,including two new species

The larch bark beetle (Ips subelongatus), which occurs in larch plantations over a vast area of eastern Asia, infects both dying and fallen trees. When its population reaches a high density, the beetle may also infect healthy trees, resulting in tree decline and, eventually, death. Leptographium spp., in both their sexual and asexual states, are mainly associated with conifer-infesting bark beetles; some species are important tree pathogens. The aims of this study were to identify the Leptographium spp. associated with I. subelongatus infestations of Larix spp. in northern China and to examine their pathogenicity towards the tree. Morphological studies and phylogenetic approaches based on multilocus DNA sequence data (ITS2- partial r28S, partial β-tubulin, and EF-1α gene regions) showed that three Leptographium species occur in association with I. subelongatus in the areas investigated: Leptographium taigense, which is recorded in China for the first time, and two new species, namely L. innermongolicum sp. nov. and L. zhangii sp. nov. Leptographium innermongolicum is closely related to L. taigense, whereas L. zhangii belongs to the Grosmannia piceaperda species complex. The pathogenicity of these Leptographium species towards mature Larix spp. was tested by stem inoculation in forests. All inoculations only resulted in small lesions on the inner bark; therefore, the three Leptographium species were not considered to be pathogenic.     

Identification and genetic diversity of two invasive <Emphasis Type="Italic">Pissodes</Emphasis> spp. Germar (Coleoptera: Curculionidae) in their introduced range in the southern hemisphere

During the first half of the twentieth century, two accidental cases of introduction of Pissodes weevils were recorded from the southern hemisphere. The weevils in South Africa were identified as the deodar weevil (Pissodes nemorensis) and those in South America as the small banded pine weevil (Pissodes castaneus). Wide distribution of the two species in their invasive range, general difficulty in identifying some Pissodes spp., and the varying feeding and breeding behaviours of the species in South Africa has necessitated better evidence of species identity and genetic diversity of both species and population structure of the species in South Africa. Barcoding and the Jerry-to-Pat region of the COI gene were investigated. Morphometric data of the South African species was analysed. Our results confirmed the introduction of only one Pissodes species of North American origin to South Africa. However, this species is not P. nemorensis, but an unrecognized species of the P. strobi complex or a hybrid between P. strobi and P. nemorensis. Only P. castaneus, of European origin, was identified from South America. We identified ten mitochondrial DNA haplotypes from South Africa with evidence of moderate genetic structure among geographic populations. Terminal leader and bole-feeding weevils did not differ at the COI locus. A single haplotype was identified from populations of P. castaneus in South America. Results of the present study will have implications on quarantine, research and management of these insect species.     

Thermostable phycocyanin from the red microalga <Emphasis Type="Italic">Cyanidioschyzon merolae</Emphasis>, a new natural blue food colorant

The demand for natural food colorants is growing as consumers question the use of artificial colorants more and more. The phycobiliprotein C-phycocyanin of Arthospira platensis is used as a natural blue colorant in certain food products. The thermoacidophilic red microalga Cyanidioschyzon merolae might provide an alternative source of phycocyanin. Cyanidioschyzon merolae belongs to the order Cyanidiophyceae of the phylum Rhodophyta. Its natural habitat are sulfuric hot springs and geysers found near volcanic areas in, e.g., Yellowstone National Park in the USA and in Java, Indonesia. It grows optimally at a pH between 0.5 and 3.0 and at temperatures up to 56 °C. The low pH at which C. merolae grows minimizes the risk of microbial contamination and could limit production loss. As C. merolae lacks a cell wall, phycocyanin with a high purity number of 9.9 could be extracted by an osmotic shock using a simple ultrapure water extraction followed by centrifugation. The denaturation midpoint at pH 5 was 83 °C, being considerably higher than the A. platensis phycocyanin (65 °C). The C. merolae phycocyanin was relatively stable at pH 4 and 5 up to 80 °C. The high thermostability at slightly acidic pH makes the C. merolae phycocyanin an interesting alternative to A. platensis phycocyanin as a natural blue food colorant.     

Primary Cutaneous Coccidioidomycosis in an Italian Nun Working in South America and Review of Published Literature

Coccidioidomycosis is a systemic disease caused by the dimorphic fungus Coccidioides, endemic in parts of the Southwestern USA and Central and South America. Two species, Coccidioides immitis and Coccidioides posadasii, were differentiated. Primary cutaneous coccidioidomycosis (PCC) has been reported rarely. An unusual case of PCC characterized by a persistent solitary lesion diagnosed in Italy in an immunocompetent Italian nun living in Argentina is described. The isolate was identified by sequence analysis as C. posadasii. Antibody screening was negative. A total of 39 cases of PCC have been reported in the literature. Infections occurred as a consequence of traumatic implantation in a natural setting in endemic areas or of accidental inoculation in laboratory workers. Importance of accurate investigation of travel history and of occupational hazards to laboratory workers is outlined.     

Dispersal of <Emphasis Type="Italic">Fusarium</Emphasis> spp. by rainwater and pathogenicity on four plant species

Research focused on the occurrence of Fusarium spp. in atmospheric dust or rainwater is not common. Preliminary studies with four sampling dates in 2007 revealed that several species of Fusarium may also be conveyed by rainwater. In order to determine the regular presence of Fusarium spp. in rainfall water, samples were systematically collected for a year (from October 2009 to October 2010) in three points on the Mediterranean coast of the province of Granada (Spain) 10-km distance between them. Throughout the year of sampling, a total of 179 rainwater samples were collected during every significant rainfall event. Eight different Fusarium species were isolated from the rainwater samples: F. oxysporum (32 %), F. proliferatum (26 %) and F. equiseti (20 %) coincide with previous studies, while F. dimerum (3 %), F. semitectum (4.7 %), F. solani (8 %), F. avenaceum (0.5 %) and F. chlamydosporum (3.7 %) were isolated for the first time from rainwater. Results were consistent with previous surveys conducted 100 km away from the sampling sites. Inoculation of 39 different isolates from five different Fusarium species showed pathogenicity on plants. Disease severity differed depending on the inoculated plant species, which means that rain water can be an effective vector to transport new pathogens into new cultivated areas. This work reveals some epidemiological aspects of Fusarium genus in natural environments. Some of the isolated Fusarium spp. are potential mycotoxin producers, such as zearalenone, fumonisin, moniliformin or nivalenol.