Projecting future expansion of invasive species: comparing and improving methodologies for species distribution modeling

Modeling the distributions of species, especially of invasive species in non‐native ranges, involves multiple challenges. Here, we developed some novel approaches to species distribution modeling aimed at reducing the influences of such challenges and improving the realism of projections. We estimated species–environment relationships for Parthenium hysterophorus L. (Asteraceae) with four modeling methods run with multiple scenarios of (i) sources of occurrences and geographically isolated background ranges for absences, (ii) approaches to drawing background (absence) points, and (iii) alternate sets of predictor variables. We further tested various quantitative metrics of model evaluation against biological insight. Model projections were very sensitive to the choice of training dataset. Model accuracy was much improved using a global dataset for model training, rather than restricting data input to the species’ native range. AUC score was a poor metric for model evaluation and, if used alone, was not a useful criterion for assessing model performance. Projections away from the sampled space (i.e., into areas of potential future invasion) were very different depending on the modeling methods used, raising questions about the reliability of ensemble projections. Generalized linear models gave very unrealistic projections far away from the training region. Models that efficiently fit the dominant pattern, but exclude highly local patterns in the dataset and capture interactions as they appear in data (e.g., boosted regression trees), improved generalization of the models. Biological knowledge of the species and its distribution was important in refining choices about the best set of projections. A post hoc test conducted on a new Parthenium dataset from Nepal validated excellent predictive performance of our ‘best’ model. We showed that vast stretches of currently uninvaded geographic areas on multiple continents harbor highly suitable habitats for parthenium. However, discrepancies between model predictions and parthenium invasion in Australia indicate successful management for this globally significant weed.     

Biological control of Ziziphus mauritiana (Rhamnaceae): feasibility,prospective agents and research gaps

The tropical fruit tree, Ziziphus mauritiana (Rhamnaceae), a native of the Indian subcontinent, is a pasture and environmental weed in northern Australia and Fiji. In their native range, Ziziphus spp., including commercially cultivated Z. mauritiana and Z. jujuba, are subjected to a wide range of pests and diseases. The feasibility of classical biological control of this weed has not been explored to date. Effective biological control could reduce plant vigour and seed output, thereby limiting the spread of Z. mauritiana in Australia. Two Ziziphus species are native to Australia, hence, any prospective biological control agent should be specific to Z. mauritiana. Opportunistic field surveys and literature searches identified 133 species of phytophagous insects, 9 species of phytophagous mites and 12 plant pathogens on Ziziphus spp. Host records suggest the following are possibly specific to Z. mauritiana and hence are prospective biological control agents in Australia: the seed‐feeding weevil Aubeus himalayanus; the leaf‐feeding gracillariid moth Phyllonorycter iochrysis; the leaf‐mining chrysomelid beetle Platypria erinaceus; the leaf‐folding crambid moth Synclera univocalis; the leaf‐galling midge Phyllodiplosis jujubae; and the gall‐mites Aceria cernuus and Larvacarus transitans. Host range of the rust Phakopsora zizyphi‐vulgaris includes many Ziziphus species, including the native Z. oenoplia and hence would not be a suitable biological control agent in Australia. The powdery mildew Pseudoidium ziziphi, with a host range restricted to Ziziphus species, has not been reported on Z. oenoplia. All available information on the pests and diseases of Z. mauritiana are from cultivated varieties. Hence, future surveys should focus on wild Z. mauritiana in the Indian subcontinent in areas that are climatically similar to the regions of northern Australia, where it is currently most abundant.     

Biological control of parthenium (Parthenium hysterophorus): the Australian experience

ABSTRACT

Parthenium is a Weed of National Significance in Australia. Biological control of parthenium in Australia commenced in 1977 and since then nine insect species and two rust fungi have been introduced and established. Seven of them are widespread, however the time taken for field establishment varied widely between various agents, ranging from one to 15 years. Among them, the stem-galling Epiblema moth, the stem-boring Listronotus weevil, the seed-feeding Smicronyx weevil and the root-feeding Carmenta moth occur in all parthenium-infested areas at high population densities. The leaf-feeding Zygogramma beetle occurs only in central and southern Queensland, and not in northern Queensland. The parthenium summer rust occurs seasonally in central and northern Queensland, while the parthenium winter rust is more widespread in southern Queensland than in central Queensland, but does not occur in north Queensland. The sap-feeding Stobaera planthopper and the leaf-mining Bucculatrix moth established and are widespread, but their damage levels remain very low. The stem-galling Conotrachelus weevil and the stem-boring Platphalonidia moth are believed to be established, but at very low abundance. Biological control has resulted in significant reductions in the abundance and impact of parthenium in Australia. As a result, the area infested with parthenium in central Queensland has declined since the mid-1990s. Due to the absence of many of the effective agents in southern and south-eastern Queensland, agents from central Queensland are being redistributed there. Additionally, based on Australian success, many of these agents have also been introduced into other countries around the world.     

The applicability of the plant vigor and resource regulation hypotheses in explaining Epiblema gall moth–Parthenium weed interactions

A basic question in insect–plant interactions is whether the insects respond to, or regulate plant traits, or a complex mixture of the two. The relative importance of the directions of the influence in insect–plant interactions has therefore been articulated through both the plant vigor hypothesis (PVH) and the resource regulation hypothesis (RRH). This study tested the applicability of these hypotheses in explaining the interactions between Parthenium hysterophorus L. (Asteraceae) and its stem‐galling moth, Epiblema strenuana Walker (Lepidoptera: Tortricidae). Parthenium plants exposed to galling were sampled at three sites in north Queensland, Australia, over a 2‐year period, and the relationship between gall abundance and plant vigor (plant height, biomass, flowers per plant, and branches per plant) was studied. To test the predictions of PVH and RRH, the vigor of parthenium plants protected from galling using insecticides was compared to galled plants and plants that escaped from galling. The vigor of ungalled plants was less than the vigor of galled plants. The higher plant vigor in galled plants was not due to galling, as was evident from insecticide exclusion trials. The insect seemed to preferentially gall the more vigorous plants. These findings support the predictions of the PVH and are contrary to those of RRH. Since gall abundance is linked to plant vigor, galling may have only a limited impact on the vigor of parthenium. This has implications for weed biological control. If the objective of biological control is to regulate the population of a plant by a galling insect, a preference for more vigorous plants by the insect is likely to limit its ability to regulate plant populations. This may explain the paucity of successes against biocontrol of annual weeds using gall insects.     

Impact of defoliation by the biocontrol agentZygogramma bicolorataon the weed Partheniumhysterophorus in Australia

The leaf-feeding beetle Zygogrammabicolorata Pallister was introduced from Mexico intoAustralia in 1980 as a biocontrol agent for the weedParthenium hysterophorus L. (Asteraceae). Z. bicolorata became abundant in 1990, and since 1992there has been regular outbreaks resulting in thedefoliation of the weed in central Queensland. In thisstudy we evaluated the impact of defoliation by Z. bicolorata on P. hysterophorus from 1996 to1998. Z. bicolorata caused 91–100% defoliationresulting in reductions in weed density by 32–93%,plant height by 18–65%, plant biomass by 55–89%,flower production by 75–100%, soil seed-bank by13–86% and seedling emergence in the following seasonby 73–90%. At sites with continued outbreaks ofZ. bicolorata, it is expected that the existing soilseed-bank will be minimised, resulting in reduceddensity of parthenium in 6 to 7 years.     

Establishment of the biological control agent Hypocosmia pyrochroma for Dolichandra unguis-cati (Bignoniaceae) is limited by microclimate

Cat's claw creeper, Dolichandra unguis-cati (Bignoniaceae), a perennial woody vine native to tropical America, is a target for biological control in Australia and South Africa. The cat's claw creeper leaf-tying moth Hypocosmia pyrochroma (Lepidoptera: Pyralidae) from tropical South America was released as a biological control agent for cat's claw creeper in Australia from 2007 to 2010. A total of 2,277 adults, 837 pupae and 77,250 larvae were released at 40 sites in Queensland and New South Wales. Releases were made mostly in open fields (85%), and at limited sites (15%) in insect-proof cages erected over naturally occurring cat's claw creeper infestations in the field. Sampling was conducted annually in spring and autumn to monitor the establishment and dispersal of H. pyrochroma. Establishment of H. pyrochroma was first noticed in 2012 at three release sites and since then the number of established sites has increased to 80 in 2020. Establishment was evident on both ‘short-pod’ and ‘long-pod’ forms of cat's claw creeper and was more widespread in sites where releases were made within insect-proof field cages (50%) than in sites with open field releases (9%). The moth was active from late spring to late autumn with peak larval activity in late summer. To date, all field establishments have been in areas predicted by a CLIMEX model as climatically suitable but restricted mostly to riparian environment (93% of establishment), where the moth has continued to spread from 1.5 to 23 km from release sites. In contrast, there is the only limited establishment and spread in non-riparian corridors, highlighting the role of microclimate (riparian) as a limiting factor for establishment and spread. Future efforts will focus on redistribution of the agent to river/creek systems where the moth is currently not present.     

Complementing biological control with plant suppression: Implications for improved management of parthenium weed (Parthenium hysterophorus L.)

Parthenium hysterophorus L. is a weed of global significance that has become a major weed in Australia and many other parts of the world. A combined approach for the management of parthenium weed using biological control and plant suppression, was tested under field conditions over a two-year period in southern central Queensland. The six suppressive plant species, selected for their demonstrably suppressive ability in earlier glasshouse studies, worked synergistically with the biological control agents (Epiblema strenuana Walker, Zygogramma bicolorata Pallister, Listronotus setosipennis Hustache and Puccinia abrupta var. partheniicola) present in the field to reduce the growth (above ground biomass) of parthenium weed, by between 60–86% and 47–91%, in Years 1 and 2, respectively. The biomass of the suppressive plants was between 6% and 23% greater when biological control agents were present than when the biological control agents had been excluded. This shows that parthenium weed can be more effectively managed by combining the current biological control management strategy with selected sown suppressive plant species, both in Australia and elsewhere.     

Life history of Anomalococcus indicus (Hemiptera: Lecanodiaspididae), a potential biological control agent for prickly acacia Vachellia nilotica ssp. indica in Australia

Babul scale Anomalococcus indicus Ramakrishna Ayyar, a major pest of Vachellia nilotica (L.f.) P.J.H. Hurter & Mabb. on the Indian subcontinent, has been identified as a potential biocontrol agent for prickly acacia V. nilotica subsp. indica (Benth.) Kyal. & Boatwr. in Australia and was imported from southern India for detailed assessment. The life history of A. indicus under controlled glasshouse conditions was determined as a part of this assessment. Consistent with other scale species, A. indicus has a distinct sexual dimorphism which becomes apparent during the second instar. Females have three instars, developing into sexually mature nymphs after 52 days. The generation time from egg to egg was 89 days. Females are ovoviviparous, ovipositing mature eggs into a cavity underneath their body. An average of 802 ± 114 offspring were produced per female. Reproductive output was closely associated with female size; larger females produced more than 1200 offspring. Crawlers emerged from beneath the female after an indeterminate period of inactivity. They have the only life stage at which A. indicus can disperse, though the majority settle close to their parent female forming aggregative distributions. In the absence of food, most crawlers died within three days. Males took 62 days to develop through five instars. Unlike females, males underwent complete metamorphosis. Adult males were small and winged, and lived for less than a day. Parthenogenesis was not observed in females excluded from males. The life history of A. indicus allows it to complement other biological control agents already established on prickly acacia in Australia.     

Localization of feeding of Anomalococcus indicus (Hemiptera: Lecanodiaspididae) and supplementary biological notes: towards the biological management of the invasive tree Vachellia nilotica indica (Fabales: Mimosoideae) in North-Eastern Australia

Summary. Management of the invasive Vachellia nilotica indica infesting tropical grasslands of Northern Australia has remained unsuccessful to date. Presently Anomalococcus indicus is considered a potential agent in the biological management of V. n. indica. Whereas generic biological details of A. indicus have been known, their feeding activity and details of their mouthparts and the sensory structures that are associated with their feeding action are not known. This paper provides details of those gaps. Nymphal instars I and II feed on cortical-parenchyma cells of young stems of V. n. indica, whereas nymphal instars III and adult females feed on phloem elements of older shoots. Nymphal instars and adults (females) trigger stress symptoms in the feeding tissue with cells bearing enlarged and disfigured nuclei, cytoplasmic shrinkage, cytoplasmic trabeculae, abnormal protuberances and uneven cell wall thickening, unusual cell membrane proliferation, and exhausted and necrosed cells. Continuous nutrient extraction by A. indicus can cause stem death. We provide evidence that A. indicus, by virtue of its continuous feeding activity and intense population build up, can be an effective biological-management agent to regulate populations of V. n. indica in infested areas.     

Current Status of the Stem-boring Weevil Listronotus setosipennis (Coleoptera: Curculionidae) Introduced Against the Weed Parthenium hysterophorus (Asteraceae) in Australia

Effects of the stem-boring weevil Listronotus setosipennis (Coleoptera: Curculionidae) on different growth stages of the weed Parthenium hysterophorus (Asteraceae) was evaluated in a glasshouse and field cages, and its field prevalence in Queensland, Australia was assessed. In the glasshouse, L. setosipennis reduced the plant height by 51%, number of leaves by 78%, flower production by 63% and plant biomass by 54% in rosette stage plants. Damage by L. setosipennis in rosette stage plants in the field cage reduced the primary stem height by 26% and flower production by 38%, but the impact on total plant height, basal stem width, root length, number of branches, root biomass and total plant biomass was not significant. In both glasshouse and field cages, the impact of L. setosipennis on preflowering and flowering stages of parthenium was not significant. L. setosipennis was recorded in 48% of the parthenium-infested sites (n=132) sampled and 16% of the sites showed high to very high levels of incidence. L. setosipennis was more prevalent on alluvial and black soils than on clay and sandy soils. L. setosipennis is a promising biocontrol agent for regions with prolonged dry periods and erratic rainfall pattern.