Insect biology and host range: risk assessment in biological control J-R Baars BioControl Research Unit School of Biology and Environmental Science University College Dublin Natural Enemies Species: Common name: Order: Family: Pyrrhalta nymphaeae Waterlily leaf beetle Coleoptera Chrysomelidae Life History: Egg batches 6-20 3 Larval instars 10 eggs/day 13-24 days High leaf turnover 5-17% net primary production Larva-Adult Damage Plant Hosts Spatterdock, Nuphar advena (L.) Sibth. & Sm. (Nymphaeaceae) Waterlilies, Nymphaea spp. (Nymphaeaceae) Smartweed, Polygonum hydropiperoides Michx. (Polygonaceae) Smartweed, Polygonum amphibium L. (Polygonaceae) Bog myrtle, Myrica gale L. (Myricaceae) Water shield, Brasenia schreberi J.F. Gmel. (Cabombaceae) Arrowheads, Sagittaria sp. (Alismataceae) Willows, Salix sp. (Salicaceae) Water chestnut, Trapa natans L. (Trapaceae) Top down control Framework for Weed BC Target weed ecology Exploration for potential control agents Evaluation of biological control potential Host specificity testing Agent release and redistribution Agent evaluation Briese DT 2000 Classical Biological Control. In: Australian Weed Management Systems (ed. B Sindel) pp. 161-192.l Target Weed Ecology Lagarosiphon major Submerged aquatic weed Vegetative growth Spread by fragmentation & layering Top 75cm of submerged shoots (50cm x 50cm) = 160m2 Leaf surface area Complex ecology dependant on location Exploration – Origin Identifying the native range of the weed Locate evolutionary centres of origin – locate phytophage diversity Searching areas best ecoclimatically matched Characterise agent & weed populations using molecular markers Exploration – Origin Lagarosiphon is native to sub-Saharan Africa & Madagascar 9 species are described (Symoens & Triest, 1983) L. major native to southern Africa Herbarium specimens held by SANBI Altitude range: 750 - >2000m a.s.l Exploration – Origin Shoot-tip borer Leaf-miner Leaf defoliator Leaf Pathogen Stem-borer Agent selection criteria Host specificity Effectiveness Two philosophies: • Process and release a large number of candidates to find successful agents • Prioritise by understanding the ecology of the weedherbivore system, targeting specific parts of the weed’s life cycle Life cycle: Hydrellia lagarosiphon Adult Widespread Damaging Eggs Short life cycle Indications that it is host specific Other similar ephydrids used as biocontrol agents Pupa Larva Ephydridae in Ireland ~29 species in the genus Hydrellia ~12 recorded in Ireland Largest genus in the Hydrellinae Hydrellia lagarosiphon on L. major Little ecology known about the species Exception include species of economic importance e.g. Hydrellia griseola Ephydrids as pests Hydrellia griseola, Smaller Rice Leaf Miner Pest on wheat, barley, rice, maize, and timothy Leaf mines causes reduction of plant photosynthesis intensity and of crop yield 14-16% damage to leaf surface, rice yields decrease by 6-9% Populations regulated by parasitic wasps Ephydrids as BC agents 2 species released in US ~ Hydrellia pakistanae ~ H. balciunasi Target species Hydrilla verticillata Originate from Asia and Australasia Released in Southern US after host specificity testing Balciunas et al 2002 Ephydrids as BC agents Leaf impact Released 1987 In early 2000s high populations were recorded Low levels of leaf damage ~20% reduce photosynthesis Ephydrids as BC agents Populations in US Taking ~18 yrs to build up Deliberately released in 30 sites Spread to sites 300-400km from release sites Grodiwitz et al 2004 Ephydrids as BC agents Before Fly impact damages Hydrilla infestations in Lake Seminole US Monocultures replaced by mixture of species After Grodiwitz et al 2004 Evaluation of BC potential 300 Hydrellia lagarosiphon 250 1-11 larva in shoot tips in the country of origin Leaf damage increases with larval density 150 100 Number of leaves damaged ±50 leaves damaged/larva 200 Carrying capacity suggests 3-4 larvae can be maintained per shoot tip 50 0 -1 0 1 2 3 4 5 6 No. Pupae 10 9 1 3 5 7 9 8 7 6 5 4 Number of Larvae 3 2 1 0 0 2 4 6 8 10 12 14 16 Days 18 20 22 24 26 28 30 Evaluation of BC potential Damage stimulates growth Side shoots increase with herbivory Shoot tip viability dependant on size and levels of larval damage 5 4 Shoot viability decreases with increasing larval density 3 2 1 Number of side shoots 0 1 3 Larval densities 5 Host specificity testing Aims to predict the damage to nontarget species following release Colonisation of nontarget species & temporary spill-over The process has evolved over time as our understanding of host-plant interactions improve An analytical process is followed to assess the potential risks Ecology, behaviour and phylogeny Determine the fundamental and realised host range Host specificity testing Fundamental host range Absolute limits of a species host range Independent of ecological setting Realised host range Variation in host acceptance following release Spatial and temporal overlap of species Risk assessment tools Test plant lists Extensive list of plants selected using ‘Centrifugal phylogenetic method’ (Wapshere 1975) ~ a sequence of plants from those most closely related to the target weed to progressively more distantly related Experimental tests • No-choice tests • Choice tests • Field tests Test plant list Family Butomaceae Species Butomus umbellatus Eupatorium cannabinum Alismataceae Sagittaria sagittifolia Sagittaria latifolia Sagittaria rigida Sagittaria subulata Baldellia ranunculoides Luronium natans Alisma plantago-aquatica Alisma lanceolatum Alisma gramineum Damasonuim alisma Hydrocaritaceae Hydrocharis morsus-ranae Elodea canadensis Hydrilla verticillata Elodea nuttallii Stratiotes aloides Egeria densa Elodea callitrichoides Vallisneria spiralis Najadanceae Najas flexilis Najas marina Najas graminea Scheuchzeriaceae Scheuchzeria palustris Aponogetonaceae Aponogeton distachyos Juncaginaceae Triglodin palustre Triglodin maritimun Potamogetonaceae Potamogeton natans Potamogeton polygonifolius Potamogeton nodosus Potamogeton coloratus Potamogeton lucens Potamogeton gramineus Potamogeton lucens x P. perfoliatus (P. x salicifolius) Potamogeton gramineus x P. lucens (P. x zizii) Potamogeton alpinus Potamogeton gramineus x P. perfoliatus (P. x nitens) Potamogeton praelongus Potamogeton epihydrus Potamogeton perfoliatus Potamogeton friesii Potamogeton pusillus Potamogeton rutilus Potamogeton obtusifolius Potamogeton trichoides Potamogeton berchtoldii Potamogeton compressus Potamogeton crispus Potamogeton acutifolius Potamogeton filiformis Potamogeton pectinatus Groenlandia densa Zannichelliaceae Zannichellia palustris Ruppiaceae Ruppia maritima Ruppia cirrhosa Tanaka et al 1997. The phylogeny of the family Hydrocharitaceae inferred Tanaka et al 1997 Phylogeny Classification proposed by Les et al 2006 Les et al 2006. A reappraisal of phylogenetic relationships in the monocotyledon family Hydrocharitaceae (Alismatidae) Aliso 22: 211-230. Proposed Classification No native species in the subfamily Anacharidoideae 3 genera need consideration ~Hydrocharis ~Stratiotes ~Najas Other alien species include Hydrilla & Vallisneria Les et al 2006 Les et al 2006. A reappraisal of phylogenetic relationships in the monocotyledon family Hydrocharitaceae (Alismatidae) Aliso 22: 211-230. Related plants 3 genera need consideration ~Hydrocharis ~Stratiotes ~Najas Host screening Adult Host range determined by larval stage Eggs Pupa Larva Host screening Behavioural constraints False +ves False -ves (Heard 2000) Heard, T.A., 2000. Concepts in insect host-plant selection behaviour and their application to host specificity testing. Host screening 35 20 oC 30 25 Fecundity of adults dependant on temperature 16 oC 20 15 13 oC 10 Accumulative number of eggs Egg viability reduces through the life time of adult fly 10 oC 5 0 2d 4d 6d 8d 10d 12d 14d 16d 18d 20d 22d 24d 26d Days 100 20 oC 16 oC 13 oC 10 oC 80 60 40 % Egg viability 20 0 2d 4d 6d 8d 10d 12d Days 14d 16d 18d 20d 22d Risk Assessment The use of host specificity testing and field hostuse studies to make pre-release relativity-based predictions of likelihood that the agents threaten nontarget plants Test conditions designed for candidate agent evaluated Host tests can predict the likelihood of nontarget attack Retrospective assessments in the USA and Australia indicate that host testing procedures can predict field host host Acknowledgements Support by the following are kindly acknowledged Hydrellia lagarosiphon • First discovered in 2008, IFI funded survey (Baars et al. 2010 - Hydrobiologia) • New species to science • Described by John Deeming (Wales Natural History Museum) (Deeming, 2011 –African Entomology) • Two additional populations maintained from collection trip in SA (May 2010) • Variation in the male genitalia • DNA analysis (barcoding) Life+ project CAISIE Survey Objectives • Collection trip to import known candidates (i.e. Bagous spp.) • Survey to establish the presence of additional candidates • Collection trip conducted in April-May 2010 • Over 50 sites were visited, 18 with L. major Shoot tip midge (Chironomidae) • Shoot-tip mining midge (cf. Polypedilum sp.) • Similar species found on other Hydrocharitaceae • Host-specificity in question • Taxonomy in question • Not easily reared under laboratory conditions Leaf feeding moth (Lepidoptera) • Leaf feeding larvae (Nymphulinae, Paraponyx spp. & Synclita spp.) • Host specificity in question • USA call to consider Lepidopteran species to be considered for Hydrilla verticillata Climate match native vs exotic • Different species persisting in different climatic areas, better pre-adapted? • Biotypes of species Thermal tolerance • One of the main contributory factors to failure in weed biocontrol