A promising biological control agent for the invasive alien plant,Pereskia aculeata Miller (Cactaceae), in South Africa

Pereskia aculeata Miller (Cactaceae) is an invasive alien plant from Central and South America that has become a problematic environmental weed in South Africa. A potential biological control agent, the stem-wilter, Catorhintha schaffneri Brailovsky & Garcia (Coreidae), was collected in southern Brazil and imported into quarantine in South Africa. Field host range data suggested that C. schaffneri has a host range restricted to P. aculeata. No-choice nymph survival tests were then conducted on 27 test plant species in 9 families. Survival to the adult stage was only recorded on P. aculeata and the closely related Pereskia grandifolia Haw. (Cactaceae). Mortality was significantly higher on P. grandifolia with only 3% of the nymphs reaching the adult stage compared with 74% on P. aculeata indicating that P. aculeata is the primary host plant. P. grandifolia is native in South America and is of no agricultural importance in South Africa so any feeding on P. grandifolia in South Africa would have no negative environmental or economic consequences. In other tests, adult survival on P. aculeata [25.8 days (SE ± 3.74)] was significantly longer than on other test plant species [4.3 days (SE ± 0.36)] further confirming the host specificity of the species. Impact studies conducted in quarantine indicated that C. schaffneri is damaging to P. aculeata, significantly reducing the number of leaves and the shoot lengths of plants, even at relatively low insect densities. C. schaffneri is safe for release in South Africa and is likely to be a damaging and effective agent.     

Prioritisation of potential agents for the biological control of the invasive alien weed,Pereskia aculeata (Cactaceae), in South Africa

Pereskia aculeata Miller (Cactaceae) is an invasive alien species in South Africa that is native in Central and South America. In South Africa, P. aculeata outcompetes native plant species leading to a reduction in biodiversity at infested sites. Herbicidal and mechanical control of the plant is ineffective and unsustainable, so biological control is considered the only potential solution. Climatic matching and genotype matching indicated that the most appropriate regions in which to collect biological control agents were Santa Catarina and Rio de Janeiro provinces in Southern Brazil. Surveys throughout the native distribution resulted in 15 natural enemy species that were associated with the plant. Field host range data, as well as previous host plant records, were used to prioritise which of the species were most likely to be suitably host specific for release in South Africa. The mode of damage was used to determine which species were most likely to be damaging and effective if released. The most promising species prioritised for further study, including host specificity and impact studies, were the stem-wilter Catorhintha schaffneri Brailovsky & Garcia (Coreidae); the stem boring species Acanthodoxus machacalis Martins & Monné (Cerambycidae), Cryptorhynchus sp. (Curculionidae) and Maracayia chlorisalis (Walker) (Crambidae) and the fruit galler Asphondylia sp. (Cecidomyiidae). By prioritising the potential biological control agents that are most likely to be host-specific and damaging, the risk of conducting host specificity testing on unsuitable or ineffective biological control agents is reduced.     

The thermal physiology of Stenopelmus rufinasus and Neohydronomus affinis (Coleoptera: Curculionidae), two biological control agents for the invasive alien aquatic weeds,Azolla filiculoides and Pistia stratiotes in South Africa

Water lettuce, Pistia stratiotes, and red water fern, Azolla filiculoides, are floating aquatic macrophytes that have become problematic in South Africa. Two weevils, Neohydronomus affinis and Stenopelmus rufinasus, are successful biological control agents of these two species in South Africa. The aim of this study was to investigate the thermal requirements of these two species to explain their establishment patterns in the field. Laboratory results showed that both weevils are widely tolerant to cold and warm temperatures. The critical thermal minima (CT_min) of N. affinis was determined to be 5.58?±?0.31°C and the critical thermal maxima (CT_max) was 44.52?±?0.27°C, while the CT_min of S. rufinasus was 5.38?±?0.33°C and the CT_max?44.0?±?0.17°C. In addition, the lower lethal temperatures were ?9.85?±?0.06°C for N. affinis and ?6.85?±?0.13°C for S. rufinasus, and the upper lethal temperatures were 42.7?±?0.85°C for N. affinis and 41.9?±?2.52°C S. rufinasus. Using the reduced major axis regression method, the development for N. affinis was described using the formula y?=?12.976x?+?435.24, while the development of S. rufinasus was described by y?=?13.6x?+?222.45. These results showed that S. rufinasus develops twice as fast as N. affinis. Using these formulae and temperature data obtained from the South African Weather Service, N. affinis was predicted to complete between 4 and 9 generations per year in South Africa, while S. rufinasus was predicted to complete between 5 and 14 generations per year around the country. These results suggest that both species should not be limited by cold winter, nor warm summer temperatures, and should establish throughout the ranges of the weeds in South Africa.     

Using molecular methods to determine the origin of weed populations of Pereskia aculeata in South Africa and its relevance to biological control

New biological control agents are required in order to reach and sustain an adequate level of control of the declared environmental weed Pereskia aculeata Miller (Cactaceae) in South Africa. Identifying the origin of weed genotypes has been important in a number of biological control programmes and is likely to be of importance for the control of P. aculeata due to its disjunct native distribution and morphological polymorphisms between plants from different regions of the native and introduced distribution. DNA sequencing of the trnL chloroplastic intron and the phyC nuclear gene indicate that the South African weed population’s origin was in the southern region of native distribution. Inter-Simple Sequence Repeats (ISSRs) confirmed this result and added resolution to the analysis indicating that the native plants with the closest genetic distance to the South African weed population were found in Rio de Janeiro Province, Brazil. The relationship between the South African weed population and garden variety plants as well as the large genetic distance between the South African plants and the native plants suggests that the South African population may be the progeny of escaped garden variety plants that have been cultivated and possibly hybridized. The low levels of genetic variation within the South African population and the monophyly of the South African plants indicates that these plants are the progeny of a single introduction or multiple introductions from the same source. Rio de Janeiro Province in Brazil is the most appropriate region in which to survey for new biological control agents.     

Impact of the shoot-boring moth Dichrorampha odorata (Lepidoptera: Tortricidae) on growth and reproductive potential of Chromolaena odorata (Asteraceae) in the laboratory

A 9-month laboratory study was carried out to determine the impact of herbivory by a moth with shoot-boring larvae, Dichrorampha odorata Brown and Zachariades (Lepidoptera: Tortricidae) on growth and reproductive ability of its host plant, Chromolaena odorata (L.) King and Robinson (Asteraceae), a major invasive alien plant species in southern Africa. Newly hatched D. odorata larvae were inoculated onto 0 (control), 50 and 100% of the shoot tips of C. odorata in the laboratory. At all treatment levels, the basal stem diameter of C. odorata was not affected by D. odorata larval feeding. Larval feeding by D. odorata significantly reduced the height of the main shoot and flower production in C. odorata relative to the control treatment but promoted branching by increasing the number of shoot tips. However, the differences in plant height and number of flowers between the 50 and 100% inoculation levels were not statistically significant. Dichrorampha odorata is the first shoot-tip attacking agent that is being released as a biological control agent against C. odorata in South Africa. In general, the impacts of D. odorata on the weed were relatively small even though statistically significant. The findings of this study suggest that high levels of damage by the moth will modestly reduce the height, flower production, and the competitiveness of C. odorata, thereby contributing to the biological control of the weed in South Africa.     

Climate constrains the establishment and proliferation of Anthonomus santacruzi,a flower-feeding biological control agent of the invasive weed Solanum mauritianum in South Africa

The flowerbud-feeding weevil Anthonomus santacruzi Hustache (Coleoptera: Curculionidae) was released in South Africa in 2008 for the biological control of the invasive tree Solanum mauritianum Scopoli (Solanaceae). The weevil was widely deployed throughout KwaZulu-Natal province, which supports large S. mauritianum infestations, and has become well established in its warmer coastal regions. The aim of this study was to provide field evidence that climate is constraining the weevil’s distribution in South Africa. Solanum mauritianum populations were sampled at 23 sites across an altitudinal gradient in KwaZulu-Natal to determine A. santacruzi densities in relation to food availability and climatic variables. Despite significantly higher amounts of floral material on S. mauritianum at the higher altitude inland sites, A. santacruzi numbers were significantly higher at the lower altitude coastal sites. There was thus a significant negative relationship between A. santacruzi numbers and altitude and significant positive relationships between A. santacruzi numbers and both temperature and humidity. Neither rainfall nor food availability influenced A. santacruzi numbers, although lower amounts of floral material at the coastal sites may well have been caused by higher weevil densities at these sites. Anthonomus santacruzi was absent at only three sites, all at higher altitudes, further demonstrating that conditions in coastal or low-altitude regions are favourable for establishment and population proliferation. Future release efforts in KwaZulu-Natal, but also in other South African provinces, should thus be focused on coastal regions and inland regions that are below 1000 m above sea level.     

Candidates for biocontrol of Macfadyena unguis-cati in South Africa: biology, host ranges and potential impact of Carvalhotingis visenda and Carvalhotingis hollandi under quarantine conditions

Macfadyena unguis-cati (L.) Gentry (Bignoniaceae) was introduced as an ornamental in South Africa, but is fast becoming an important invasive plant in many areas. It is difficult to control the plant chemically and mechanically. The first biocontrol agent, the chrysomelid Charidotis auroguttata (Boheman), has been released. It established at some release sites, but numbers have so far remained low. Additional biocontrol agents were sought to augment C. auroguttata. The potential host ranges of two foliage feeding lace bugs, Carvalhotingis visenda (Drake and Hambleton) and C. hollandi (Drake) (Hemiptera: Tingidae) were evaluated on the basis of nymphal no-choice and adult multi-choice tests involving 23 plant species in 11 families. In no-choice tests, nymphs of both species were able to survive and complete development on M. unguis-cati only, and adults of both species only fed and oviposited on M. unguis-cati during the adult multi-choice tests. Host specificity tests thus confirm that the tingids are highly host specific biocontrol agents, and will not pose risk to any non-target plants in South Africa. A study to determine the potential impact of C. hollandi nymphal feeding on M. unguis-cati showed a significant decrease in the chlorophyll contents of leaves when compared to those of control plants. These studies indicate that, once released, the two lace bug species could contribute significantly to the biological control of M. unguis-cati in South Africa.     

Pereskiophaga brasiliensis,a natural enemy of the invasive alien cactus Pereskia aculeata,is not suitably host specific for biological control in South Africa

The stem-mining weevil, Pereskiophaga brasiliensis, was a candidate biological control agent for the invasive cactus Pereskia aculeata in South Africa. In host specificity trials, it developed on two indigenous test plant species under choice and no-choice conditions. Pereskiophaga brasiliensis is therefore not suitably host specific for release in South Africa.     

Densities of the arundo wasp,Tetramesa romana (Hymenoptera: Eurytomidae) across its native range in Mediterranean Europe and introduced ranges in North America and Africa

Tetramesa romana is a biological control agent of the giant reed, Arundo donax (Poaceae: Arundinoideae), which is an invasive weed in the riparian habitats of the Rio Grande Basin of Texas, the southwestern U.S.A. and northern Mexico. Field evaluations were conducted in the native range of T. romana in Mediterranean Europe and in the introduced ranges of Texas, California, and South Africa to compare densities of the wasp. Population densities and percentage parasitism levels for the 2017 year are compared to meteorological variables (average temperature, precipitation, and heat units). In the introduced ranges of Texas (intentional) and South Africa (adventive) T. romana population densities were 39 and 10-fold higher than in the native range, respectively. Percentage parasitism of T. romana in Texas and in the native range of Thessaloniki, Greece were 2.0% and 34.3%, respectively. Annual heat unit accumulation was 1.3–2.7-fold higher at Texas sites than at other introduced or native sites, and heat units were positively associated with exit hole counts at introduced sites. Annual precipitation was 2-fold higher at Texas and South African sites than in California and the native range sites. Favourable weather conditions and reduced parasitism in Texas along the Rio Grande, as compared to the native range, allows T. romana to reach higher population levels and cause considerable damage to A. donax.     

Invaded habitat incompatibility affects the suitability of the potential biological control agent Listronotus sordidus for Sagittaria platyphylla in South Africa

Sagittaria platyphylla (Engelmann) J.G. Smith (Alismataceae) was first recorded in South Africa in 2008 and is considered to be an emerging weed with naturalised populations occurring throughout the country. A biological control programme was initiated in Australia and surveys conducted between 2010 and 2012 yielded potential agents, including the crown feeding weevil, Listronotus sordidus Gyllenhal (Coleoptera: Curculionidae). The potential of L. sordidus as a candidate biological control agent against S. platyphylla in South Africa was examined. Although adult feeding was recorded on a number of plant species, oviposition and larval development indicated a narrow host range restricted to the Alismataceae. In South Africa, S. platyphylla populations are primarily found in inundated systems. However, laboratory studies showed that L. sordidus did not oviposit on inundated plants, potentially nullifying the impact of the insect on South African populations. It is suggested that even though L. sordidus is a damaging, specific agent, its limited impact on inundated plant populations in South Africa does not justify the inherent risk associated with the release of a biological control agent.     

Response of the invasive pompom weed,Campuloclinium macrocephalum (Asteraceae), to feeding by the foliage-deforming thrips Liothrips tractabilis (Phlaeothripidae) under outdoor conditions in South Africa

The foliage-deforming thrips Liothrips tractabilis Mound and Pereyra (Thysanoptera: Phlaeothripidae) was recently released in South Africa as the first classical biological control agent against the invasive South American herb, Campuloclinium macrocephalum (Less.) DC. (Asteraceae). The impact of the thrips on young plants and regrowth of C. macrocephalum was assessed under outdoor conditions, and the results were compared to those of earlier laboratory trials that were conducted while the agent was still under investigation in quarantine. The outdoor trials revealed that feeding by L. tractabilis reduced the growth and biomass accumulation of C. macrocephalum, particularly in young plants and to a lesser extent in regrowth, which was largely consistent with the results of the earlier laboratory trials. Thrips-infested young plants were significantly shorter, produced fewer leaves and displayed significantly lower wet root masses and lower below- and above-ground dry masses, compared to the control plants. Although thrips-infested regrowth were significantly shorter, produced fewer leaves and displayed significantly lower above-ground dry masses than the control plants, the differences in wet root masses and below-ground dry masses were not significant. Although laboratory-based impact assessments are not necessarily an accurate reflection of field impact, in this instance the results were largely validated by the outdoor assessments. The results suggest that L. tractabilis will have a negative impact on C. macrocephalum populations in the field and thus contribute towards the weed’s biological control. The establishment of L. tractabilis has recently been confirmed in South Africa, providing the opportunity to verify these predictions.     

Predicting the distribution of Eccritotarsus catarinensis, a natural enemy released on water hyacinth in South Africa

Water hyacinth [Eichhornia crassipes (Mart.) Solms (Pontederiaceae)] is the most damaging aquatic weed in South Africa, where five arthropod biological control agents have been released against it. The most recent introduction of Eccritotarsus catarinensis (Carvalho) (Heteroptera: Miridae) has failed to establish permanent populations at a number of sites in South Africa where water hyacinth is a problem. Cold winter temperatures at these sites are assumed to be the reason for these establishment failures. This assumption was tested by investigating the thermal physiology of the mirid, then incorporating these data into various predictive distribution models. Degree‐day models predict 3–14 generations per year at different localities in South Africa, and five generations at a Johannesburg site where the mirid failed to overwinter. The inability to develop sufficiently rapidly during winter months may hinder overwintering of this insect, which was predicted to develop through only one generation during the winter months of April to August in Johannesburg. A CLIMEX model also showed that cold stress limits the mirid's ability to overwinter in the interior of the country, while determination of the lower lethal limit (–3.5 °C) and critical thermal minimum (1.2 ± 1.17 °C) also indicated that extreme temperatures will limit establishment at certain sites. It is concluded that E. catarinensis is limited in its distribution in South Africa by low winter temperatures.     

Suppression of growth and reproductive capacity of the weed Lantana camara (Verbenaceae) by Ophiomyia camarae (Diptera: Agromyzidae) and Teleonemia scrupulosa (Heteroptera: Tingidae)

An 8-month study was conducted to determine the effect of herbivory by the herringbone leafminer Ophiomyia camarae Spencer (Diptera: Agromyzidae) and the lace bug Teleonemia scrupulosa Stål (Heteroptera: Tingidae) on growth and reproductive capacity of their host plant, Lantana camara L. (Verbenaceae). Ophiomyia camarae adults were released into field cages at two population densities while T. scrupulosa, a widespread and naturalised biocontrol agent in South Africa, colonised and attacked uncaged plants of similar size grown under similar field conditions. At lower initial density, O. camarae reduced stem height, stem diameter, leaf density, flower density and above-ground biomass by 5, 22, 54, 100 and 41%, respectively. At higher initial density of O. camarae, stem height, stem diameter, leaf density, flower density and above-ground biomass were substantially reduced by 19, 28, 73, 99 and 49%, respectively. Teleonemia scrupulosa also reduced stem height, stem diameter, leaf density, flower density and above-ground biomass by 37, 34, 75, 100 and 50%, respectively. The root development was less affected by both O. camarae and T. scrupulosa. Whilst higher densities of O. camarae exerted noticeable herbivore pressure, T. scrupulosa herbivory substantially hampered plant growth and reproductive capacity even at lower population levels. These findings suggest that O. camarae, released in 2001 for the control of lantana, would be effective as part of a complementary guild of biological control agents. The profound effect of herbivory by T. scrupulosa on plant growth and reproductive capacity suggests that it is making a much greater contribution to the biological control of lantana in South Africa than was previously thought.     

Phytophagous insects collected fromParkinsonia aculeata [Leguminosae: Caesalpiniaceae] in the Sonoran desert region of the southwestern United States and Mexico

Parkinsonia aculeata L. (Leguminosae: Caesalpiniaceae), a weed in northern Australia, was the target of a biological control programme from 1983–1987. Sixty-five phytophagous insect species were collected fromP. aculeata growing in the Sonoran desert region of the southwestern United States and Mexico. Six orders (Coleoptera, Diptera, Hemiptera, Lepidoptera, Orthoptera and Thysanoptera) and 30 families were represented. Of the 12 species with potential as biological control agents, only one has passed specificity testing and has been released in Australia.

---

Résumé Parkinsonia aculeata L. (Leguminosae: Caesalpinioideae), mauvaise herbe du nord de l'Australie, fut l'objet d'un programme de lutte biologique entre 1983 et 1987. Soixante-cinq espèces d'insectes phytophages furent récoltés surP. aculeata dans la région du désert Sonoran, (sud-ouest des Etats-Unis) et au Mexique. Six ordres (Coleoptera, Diptera, Hemiptera, Lepidoptera, Orthoptera et Thysanoptera) et 30 familles furent représentés. Parmi les 12 espèces qui sont potentiellement des agents de lutte biologique, une seule est acceptable selon des tests de spécificité et a été libérée en Australie.

     

Occurrence of different Lantana camara varieties across four South African provinces and their susceptibility to a biotype of the gall-forming mite Aceria lantanae

A biotype of the flower-galling mite, Aceria lantanae (Cook) (Trombidiformes: Eriophyidae) collected in Florida (U.S.A.) was released in South Africa in 2007 against Lantana camara L. (Verbenaceae) but has displayed patchy establishment. The occurrence of different L. camara varieties and their susceptibility to A. lantanae were assessed across four provinces with dense infestations. Surveys were undertaken at 113 sites during the mite’s peak infestation period (April–May) in 2013–2015. The occurrence of 13 recorded L. camara varieties differed substantially across and within these provinces. Overall, five varieties accounted for 7–45% of the sampled plants at 9–51% of the surveyed sites. The remaining eight varieties accounted for <1–4% of the plants at 2–9% of the sites. The establishment and impact of A. lantanae differed significantly between L. camara varieties. The mite established best on three varieties (163 LP, 021 WP and 015 OR), with 60–90% of plants infested. Reduced establishments were observed on seven varieties, with 18–50% of plants infested, while no establishment was recorded on three varieties. Where established, A. lantanae inflicted considerable levels of damage (i.e. 51–75% of buds infested) on the most widespread and abundant L. camara variety (163 LP) and on one less common variety (021 WP). Two uncommon varieties (015 OR, 021 P) suffered moderate levels of damage with the remainder suffering only trivial levels. The mite’s impact in South Africa could be improved by complementing the established biotype with others from Central and South America that are better matched with the poorly attacked L. camara varieties.     

Toxic effect of herbicides used for water hyacinth control on two insects released for its biological control in South Africa

The integrated control of water hyacinth, Eichhornia crassipes (Martius) Solms-Laubach (Pontederiaceae) has become necessary in South Africa, as biological control alone is perceived to be too slow in controlling the weed. In total, seven insect biological control agents have been released on water hyacinth in South Africa. At the same time, herbicides are applied by the water authorities in areas where the weed continues to be troublesome. This study investigated the assumption that the two control methods are compatible by testing the direct toxicity of a range of herbicide formulations and surfactants on two of the biological control agents released against water hyacinth, the weevil, Neochetina eichhorniae Warner (Coleoptera: Curculionidae) and the water hyacinth mirid, Eccritotarsus catarinensis (Carvalho) (Hemiptera: Miridae). A number of the formulations used resulted in significant mortality of the mirid and the weevil. Products containing 2,4-D amine and diquat as active ingredients caused higher mortality of both agents (up to 80% for the mirid) than formulations containing glyphosate. Furthermore, when surfactants were added to enhance herbicide efficiency, it resulted in increased toxicity to the insects. We recommend that glyphosate formulations should be used in integrated control programmes, and that surfactants be avoided in order to reduce the toxic nature of spray formulations to the insect biological control agents released against water hyacinth.     

Host range of Lepidelphax pistiae (Hemiptera: Delphacidae) and its potential impact on Pistia stratiotes L. (Araceae)

Pistia stratiotes L. (Araceae) is a floating aquatic plant that has become invasive in Florida. It is primarily controlled with herbicides, but two biocontrol agents have previously been released to assist in management of this species. A new potential biocontrol agent from Argentina, Lepidelphax pistiae Remes Lenicov (Hemiptera: Delphacidae), has been evaluated comprehensively for specificity after initial host range studies done in its native range indicated that it is likely specific to P. stratiotes. Host range studies indicated that this insect is specific to P. stratiotes, with no survival or reproduction occurring on any of the 42 other plant species tested. Impact studies indicated that this insect can significantly damage P. stratiotes at medium and high population densities, which were comparable to those seen in its native range. Lepidelphax pistiae is sufficiently specific enough to warrant release and has a high probability of aiding management of P. stratiotes populations in Florida.     

Phenology of Tick weed (Cleome viscosa L.) and its interaction with okra (Abelmoschus esculentus (L.) Moench) and Melodoigyne incognita

Cleome viscosa is an emerging weed with the potential of interfering with okra and influencing pests of okra. Screen house studies were conducted on the phenology of C. viscosa, its interference with okra and its interaction with root-knot nematode-infected okra. Seedlings of C. viscosa were monitored in pots for growth, yield and dry matter accumulation for 14 weeks. C. viscosa was planted with okra at densities 0, 2, 4, 6, 8 and 10 weeds per okra plant and observed for 11 weeks. Data were collected on growth, yield and dry matter of okra. Plants were inoculated with 2,500 M. incognita eggs per pot while control plants were not inoculated. C. viscosa attained 91.7 cm height and accumulated 7.8 g/plant biomass at 14 weeks after planting. The percentage reduction in okra plant height, plant dry weight and fruit yield due to interference at lowest cleome density (2 plants/pot) was 33.7, 83.6 and 82.1%, respectively. Nematode reproductive factor was significantly lower for okra alone (4.9) compared to okra with cleome (7.6). This study shows that C. viscosa is a fast-growing weed that suppressed the performance of okra even at low density, is a good host to M. incognita and increased the population of the nematode in soil.     

Risk assessment of Episimus unguiculus (Lepidoptera: Tortricidae), a biological control agent of Schinus terebinthifolia (Sapindales: Anacardiaceae) in Hawaii,USA

Schinus terebinthifolia Raddi (Anacardiaceae) is an introduced ornamental tree from South America that has become one of the most invasive weeds in Hawaii and Florida, USA. Exploratory surveys in the plant’s native range from 1950 to 2014 identified several potential biological control agents. One of these is the leaflet rolling moth Episimus unguiculus Clarke (Lepidoptera: Tortricidae), previously known as Episimus utilis Zimmerman. This biological control agent was released in Hawaii in the 1950s where high densities were occasionally observed, leading to partial control of S. terebinthifolia by the 1960s. Larvae are leaf tiers capable of completely defoliating small plants. In order to investigate the release of E. unguiculus in the continental USA, a series of laboratory no- choice, and multiple-choice tests were conducted in Florida, and a preliminary open field test with a native plant in Hawaii. Under the confined laboratory conditions imposed during the no-choice tests, E. unguiculus accepted the economically important Pistacia spp. and several other non-target plants for oviposition and development. However, in the multiple-choice tests E. unguiculus exhibited a clear preference for S. terebinthifolia relative to non-target plants accepted in the no-choice tests. Overall, the results of field observations during surveys in South America and Hawaii and host range studies completed in Hawaii and Florida showed that E. unguiculus is a narrow specialist on S. terebinthifolia, its natural host plant.     

Impact of a disease and a defoliating insect on houndstongue (Cynoglossum officinale) growth: implications for weed biological control

The noxious weed houndstongue (Cynoglossum officinale) has become a major problem on the forested rangelands in the interior of British Columbia. However, recently the fungus Phoma pomorum and the ranchman's tigermoth (Platyprepia virginalis) were identified as potential biocontrol agents of this biennial weed. Infection by the fungus Ph. pomorum resulted in the formation of large brown lesions on leaves of houndstongue. In culture, the fungus readily produced pycnidia with pycnidiospores measuring 5.7 μm × 1.7 μm. The effect of Ph. pomorum and P. virginalis on the growth of houndstongue was examined over a six week period. Leaf age strongly influenced the intra plant distribution of insect feeding and lesion formation with Ph. pomorum primarily attacking the older leaves, while P. virginalis larvae preferred feeding on young leaves. Infection of leaves by Ph. pomorum resulted in their premature death. The effect of infection by Ph. pomorum on plant weight varied among tests, but the disease usually increased the number or percentage of dead leaves and reduced root biomass. A study of the effect of plant age and disease on houndstongue showed that younger plants infected with Ph. pomorum had a slightly higher percentage of dead leaves than older plants, but that reductions in live leaf weight and root weight were similar for different age groups. Six weeks after exposure to feeding damage by P. virginalis, there was no significant effect of P. virginalis on plant weight either acting alone or in combination with Ph. pomorum.