Thai K. Van

Herbivory alters resource allocation and compensation in the invasive tree Melaleuca quinquenervia

Abstract.  1. Plants may compensate for the effects of herbivory, especially under favourable growing conditions, limited competition, and minimal top‐down regulation. These conditions characterise many disturbed wetlands dominated by introduced plants, implying that exotic, invasive weeds in these systems should exhibit strong compensatory responses.

Alteration of water hyacinth (Eichhornia crassipes (Mart.) Solms) leaf dynamics and phytochemistry by insect damage and plant density

Abstract Water hyacinths (Eichhornia crassipes (Mart.) Solms) grown outdoors were subjected to 0, 25, or 250 water hyacinth weevils (Neochetina eichhorniae Warner) per culture tank. Plant densities were then reduced by 75% in half of the tanks within each weevil stocking rate. Biweekly leaf production, mortality, and loss were monitored for 22 weeks. After 12 weeks, leaf lamina tissue was analyzed for N, P, K, phenolics, carbohydrates, Na, Ca, Mg, Mn, Fe, and Zn concentrations. High weevil populations increased leaf production in low plant densities (0.170 leaves rosette−1 day−1). Otherwise weevil infestations had little overall effect on production rates (average 0.144 leaves rosette−1 day−1), but infested plants produced smaller leaves. Leaf mortality was 50% higher and the plants lost leaves 72% faster than the controls when subjected to high weevil populations. Weevil infestations had no effect on tissue N levels when plant densities were but levels were lower with weevils (2.8%) than without them (3.5%) when plant densities were high. Likewise, N was higher when plant densities were low (3.4%) than when they were high (2.8%) if weevil populations were high. Tissue P, K, Fe and Zn concentrations increased when either weevil level or plant density increased. Tissue Mg inversely decreased with increasing weevil levels. It is concluded that weevil infestations disrupt water hyacinth leaf dynamics by increasing leaf mortality rates. This effect is most severe in high plant densities but in low densities plants may partially compensate through increased production of smaller leaves. Both insect damage and plant density alter tissue nutrient concentrations, but the significance of changes is confounded by plasticity in leaf size. Weevil infestations effectively reduce total quantities of nutrients accumulated within whole leaf laminas but this is manifested more by reduced leaf size than by reduced tissue loads.

Geographic Distribution and Dispersal Rate of Oxyops vitiosa (Coleoptera: Curculionidae), a Biological Control Agent of the Invasive Tree Melaleuca quinquenervia in South Florida

Abstract We assess the geographic distribution and rate of spread of Oxyops vitiosa (Pascoe), a classical biological control agent of the invasive Australian tree Melaleuca quinquenervia (Cav.) S.T. Blake. This weevil has been released at 135 locations in south Florida, where it now occurs in 9 of 19 infested counties. When averaging dispersal distances among four representative sites, O. vitiosa spread at a rate of 0.99 (±0.28) km/yr, ranging from 0.10 to 2.78 km/yr. The rate of spread by O. vitiosa across melaleuca-dominated habitats was influenced by both ecological- and human-mediated parameters, including M. quinquenervia stand fragmentation (spatial separation among host plants), the number of weevils released, and time since release. The rate of spread was positively correlated with stand fragmentation level: high = 2.04, medium = 1.07, and low = 0.30 km/yr. By incorporating the dispersal rate from the highest fragmentation level into a simulation model we predicted that 138 months (June 2008) would be required for 50% of the habitat currently invaded by melaleuca to become infested at an economic weevil density (0.5 individuals per branch tip). At medium and low fragmentations, the model predicts 182 (February 2012) and 191 (November 2012) months, respectively. After examining the output from this basic model, we identified 16 possible redistribution sites that may accelerate the spread of the weevil.

Regeneration Potential of the Canopy-Held Seeds of Melaleuca quinquenervia in South Florida

Melaleuca quinquenervia produces and maintains extensive seed reservoirs in the forest canopy. We collected capsules from different infructescences (clusters; Cluster I is the youngest, located at the most distal position, and Cluster VII is the oldest, located at the most proximal position) on branches from dry, seasonally flooded, and permanently flooded habitats. Extracted seeds were soaked for 10 d in sterile deionized water and/or 2,3,5,-triphenol tetrazolium chloride (TTC) stain to assess viability. Microscopic inspections revealed that only 15% of the seeds were embryonic (filled), 50% of embryonic seeds were viable (stained red with TTC), and 73% of viable seeds were germinable after 10 d. The remaining 27% of viable seeds may have been dormant or, possibly, required special conditions for germination. A higher percentage of seeds were embryonic on trees at permanently flooded habitats (18%) when compared to dry (14%) or seasonally flooded (14%) habitats. Overall seed viabilities and germinabilities were comparable among the three habitat types. Proportions of filled seeds were constant among infructescence positions within each habitat. Both viability and germinability of seeds varied with infructescence age, both being highest in Clusters II-V and lowest in Cluster VII.

Initial Impacts and Field Validation of Host Range for Boreioglycaspis melaleucae Moore (Hemiptera: Psyllidae), a Biological Control Agent of the Invasive Tree Melaleuca quinquenervia (Cav.) Blake (Myrtales: Myrtaceae: Leptospermoideae)

Abstract Invasion of south Florida wetlands by the Australian paperbark tree, Melaleuca quinquenervia (Cav.) S.T. Blake (melaleuca), has caused adverse economic and environmental impacts. The tree’s biological attributes and favorable ambient biophysical conditions combine to complicate efforts to restore and maintain south Florida ecosystems. Management requires an integrated strategy that deploys multiple biological control agents to forestall reinvasion and to supplement other control methods, thereby lessening recruitment and regeneration after removal of existing trees. This biological control program began during 1997 when an Australian weevil, Oxyops vitiosa (Pascoe), was released. A second Australian insect, the melaleuca psyllid (Boreioglycaspis melaleucae Moore), first introduced during 2002, has also widely established. After inoculation of the psyllid in a field study, only 40% of seedlings survived herbivory treatments compared with 95% survival in controls. The resultant defoliation also reduced growth of the surviving seedlings. A weevil-induced decline at a site comprised mainly of coppicing stumps had slowed after a 70% reduction. Psyllids colonized the site, and 37% of the remaining coppices succumbed within 10 mo. The realized ecological host range of B. melaleucae was restricted to M. quinquenervia; 18 other nontarget plant species predicted to be suboptimal or nonhosts during laboratory host range testing were unaffected when interspersed with psyllid-infested melaleuca trees in a common garden study. Evaluations are ongoing, but B. melaleucae is clearly reducing seedling recruitment and stump regrowth without adversely impacting other plant species. Manifestation of impacts on mature trees will require more time, but initial indications suggest that the psyllid will be an effective supplement to the weevil.

Decline in exotic tree density facilitates increased plant diversity: the experience from Melaleuca quinquenervia invaded wetlands

The Australian tree Melaleuca quinquenervia (melaleuca) formed dense monocultural forests several decades after invading parts of Florida and the Caribbean islands. These dominant forests have displaced native vegetation in sensitive wetland systems. We hypothesized that native plant diversity would increase following recent reductions in density of mature melaleuca stands in south Florida. We therefore examined data on changes in melaleuca densities and plant species diversity derived from permanent plots that were monitored from 1997 to 2005. These plots were located within mature melaleuca stands in nonflooded and seasonally-flooded habitats. Two host-specific biological control agents of melaleuca, Oxyops vitiosa and Boreioglycaspis melaleucae, were introduced during 1997 and 2002, respectively. Also, an adventive rust fungus Puccinia psidii and lobate-lac scale Paratachardinapesudolobata became abundant during the latter part of the study period. Overall melaleuca density declines in current study coincided with two to four fold increases in plant species diversity. The greatest declines in melaleuca density as well as the greatest increases in family importance values and species diversity indices occurred in nonflooded as compared to seasonally-flooded habitats. Most pioneer plant species in study sites belonged to Asteraceae, Cyperaceae, Poaceae, and Ulmaceae. The rapid reduction in melaleuca density and canopy cover during the study period may be attributed to self-thinning accelerated by the negative impact of natural enemies. Densities of other woody plants, particularly Myrica and Myrsine, which were sparsely represented in the understory by a few suppressed individuals also declined during the same period, possibly due to infestation by the generalist lac-scale. These findings indicate that natural-enemy accelerated self-thinning of melaleuca densities is positively influencing the native plant diversity and facilitating the partial rehabilitation of degraded habitats.

Melaleuca quinquenervia dominated forests in Florida: analyses of natural-enemy impacts on stand dynamics

Melaleuca quinquenervia (melaleuca) is a native of Australia but has become an invasive plant in Florida, USA. We conducted a long-term demographic study of melaleuca in three sections (central, transitional, and peripheral) of monoculture stands located in Florida, and quantified absolute density, diameter at breast height and basal area of trees by section at three sites. Additionally, we monitored the impacts of natural enemy (insects and fungi) on melaleuca populations which became apparent after 2001. Both absolute density and basal area, from before (1997–2001) and after noticeable natural-enemy impact (2002–2005), were compared. Prior to the natural-enemy impact, absolute density of melaleuca trees declined primarily due to self-thinning and associated losses of small trees, but diameter at breast height increased, as did the basal area. Later during the period when natural enemies prevailed, absolute density declined at a significantly greater rate across all sections but was highest at the periphery. The decrease in mean absolute density and basal area/ha of melaleuca during the natural-enemy impacted period coincided with the increased incidence of the populations of plant-feeding insects and fungi. The mean diameter at breast height continued to increase in all sections of the stands throughout the study period. An increasing trend in basal area prior to natural-enemy impact was reversed after increase in natural-enemy abundance and noticeable impact in all three sections of the stands. These findings lend support to a growing body of literature that implicates natural enemies as increasingly important density-independent regulators of M. quinquenervia populations.

Molecular and morphological examination of Cyrtobagous sp. collected from Argentina, Paraguay, Brazil, Australia, and Florida

Two members of the floating fern genus Salvinia (Salviniaceae), S. minima Baker and S. molesta Mitchell, have established in the United States. Cyrtobagous salviniae Calder and Sands (Coleoptera: Curculionidae), long established on Florida S. minima, was released in Texas and Louisiana as a biocontrol agent for both species. Subsequently, sequence analysis of the 28S rRNA D2 expansion domain suggested that the Florida and Brazilian populations (used worldwide for biocontrol) of C. salviniae might constitute two cryptic species. In response, the Brazilian weevil was imported from Australia and released instead onto S. molesta. We sampled C. singularis Hustache and C. salviniae from their native ranges in Brazil, Argentina, and Paraguay and sequenced them (D2) along with Australian and Florida samples. The genetic distance between C. singularis and C. salviniae samples is much greater (almost 5×) than the distance between either the Florida and Brazilian samples or the Brazilian and Argentinean/ Paraguayan C. salviniae samples. Since C. singularis and C. salviniae are cryptic species, the Florida and Brazilian populations (or for that matter Brazilian and Argentinean/Paraguayan) could reasonably be described as demes or ecotypes. Occurrence data indicates that, in parts of their ranges, C. salviniae and C. singularis are not only sympatric but also feed on the same plant species at the same site. While host adaptation (species preferences) likely occurs within local demes, both species seem capable of adapting to the available resource (Salvinia species). Finally, a polymerase chain reaction (PCR) primer was developed to distinguish the Florida and Brazilian/Australian types.

A genetic survey of Salvinia minima in the southern United States

The genetic relationships among 68 samples of Salvinia minima (Salviniaceae) were investigated using RAPD analysis. Neighbor joining, principle components, and AMOVA analyses were used to detect differences among geographically referenced samples within and outside of Florida. Genetic distances (Nei and Li) range up to 0.48, although most are under 0.30, still relatively high levels for an introduced, clonally reproducing plant. Despite the diversity AMOVA analysis yielded no indication that the Florida plants, as a group, were significantly different from the plants sampled elsewhere in its adventive, North American range. A single, genetically dissimilar population probably exists in the recent (1998) horticultural introduction to Mississippi. When the samples were grouped into 10 regional (but artificial) units and analyzed using AMOVA the between region variance was only 7.7%. Genetic similarity among these regions may indicate introduction and dispersal from common sources. The reduced aggressiveness of Florida populations (compared to other states) may be due to herbivory. The weevil Cyrtobagous salviniae, a selective feeder, is found in Florida but not other states. The genetic similarity also suggests that there are no obvious genetic obstacles to the establishment or efficacy of C. salviniae as a biological control agent on S. minima outside of Florida.