Synergistic interactions between leaf beetle herbivory and fire enhance tamarisk (Tamarix spp.) mortality

The combined effects of herbivory and fire on plant mortality were investigated using prescribed burns of tamarisk (Tamarix ramosissima Lebed) exposed to herbivory by the saltcedar leaf beetle (Chrysomelidae: Diorhabda carinulata Desbrocher). Tamarix stands in the Humboldt Sink (NV, USA) were divided into three treatments: summer burn (August 2006), fall burn (October 2006) and control (unburned), and litter depth was manipulated to vary fire intensity within burn seasons. A gradient of existing herbivory impact was described with three plant condition metrics prior to fire: reduced proportions of green canopy, percent root crown starch sampled at the height of the growing season (August 2006), and percent root crown starch measured during dormancy (December 2006). August root crown starch concentration and proportion green canopy were strongly correlated, although the proportion green canopy predicted mortality better than August root crown starch. December root crown starch concentration was more depleted in unburned trees and in trees burned during the summer than in fall burn trees. Mortality in summer burned trees was higher than fall burned trees due to higher fire intensity, but December root crown starch available for resprouting in the spring was also lower in summer burned trees. The greatest mortality was observed in trees with the lowest December root crown starch concentration which were exposed to high fire intensity. Disproportionate changes in the slope and curvature of prediction traces as fire intensity and December starch reach reciprocal maximum and minimum levels indicate that beetle herbivory and fire intensity are synergistic.     

Competitive abilities of native grasses and non-native (Bothriochloa spp.) grasses

Old World Bluestems (OWB), introduced from Europe and Asia in the 1920s, recently have begun to raise concerns in the Great Plains. Despite suggestion in the late 1950s that OWB were weedy and negatively impacted biological diversity, they were widely introduced throughout the Great Plains for agricultural purposes. Anecdotal evidence suggests that OWB exhibit invasive characteristics that promote competitive exclusion of native species. The objective of our study was to quantify the competitive abilities of two OWB species (Caucasian bluestem; Bothriochloa bladhii (Retz.) S.T. Blake (= Bothriochloa caucasica (Trin.) C.E. Hubb.) and yellow bluestem; Bothriochloa ischaemum (L.) Keng) with three native grass species (big bluestem (Andropogon gerardii Vitman), little bluestem (Schizachyrium scoparium (Michx.) Nash), and sideoats grama (Bouteloua curtipendula (Michx.) Torr.)). A greenhouse target-neighbor study was conducted to assess both interspecific and intraspecific competition. A total of 480 pots (4.4 l) filled with native soil was used with all pair-wise combinations of species and four density treatments (six replications). Vegetative tiller height, above- and belowground biomass were measured at the end of 16 weeks. Both of the OWB significantly inhibited at least one growth parameter of the three native grass species, while most of the native species did not inhibit growth of either OWB species. Growth of B. ischaemum was enhanced when grown in association with S. scoparium. Based upon the results of our study of OWB competitive superiority and previous research, many of the characteristics possessed by OWB are found to be in common with known invasive species. Hence, we propose that two OWB are competitively superior to three common native prairie species providing them with the ability to invade and threaten the native grasslands of the Central and Southern Great Plains.     

Phreatophytes under stress: transpiration and stomatal conductance of saltcedar (Tamarix spp.) in a high-salinity environment

Background and aims

We sought to understand the environmental constraints on an arid-zone riparian phreatophtye, saltcedar (Tamarix ramosissima and related species and hybrids), growing over a brackish aquifer along the Colorado River in the western U.S. Depth to groundwater, meteorological factors, salinity and soil hydraulic properties were compared at stress and non-stressed sites that differed in salinity of the aquifer, soil properties and water use characteristics, to identify the factors depressing water use at the stress site.

Methods

Saltcedar leaf-level transpiration (E_L), LAI, and stomatal conductance (G_S) were measured over a growing season (June–September) with Granier and stem heat balance sensors and were compared to those for saltcedar at the non-stress site determined in a previous study. Transpiration on a ground-area basis (E_G) was calculated as E_L?×?LAI. Environmental factors were regressed against hourly and daily E_L and G_S at each site to determine the main factors controlling water use at each site.

Results

At the stress site, mean E_G over the summer was only 30 % of potential evapotranspiration (ET_o). G_S and E_G peaked between 8 and 9 am then decreased over the daylight hours. Daytime G_S was negatively correlated with vapor pressure deficit (VPD) (P?<?0.05). By contrast, E_G at the non-stress site tracked the daily radiation curve, was positively correlated with VPD and was nearly equal to ET_o on a daily basis. Depth to groundwater increased over the growing season at both sites and resulted in decreasing E_G but could not explain the difference between sites. Both sites had high soil moisture levels throughout the vadose zone with high calculated unsaturated conductivity. However, salinity in the aquifer and vadose zone was three times higher at the stress site than at the non-stress site and could explain differences in plant E_G and G_S.

Conclusions

Salts accumulated in the vadose zone at both sites so usable water was confined to the saturated capillary fringe above the aquifer. Existence of a saline aquifer imposes several types of constraints on phreatophyte E_G, which need to be considered in models of plant water uptake. The heterogeneous nature of saltcedar E_G over river terraces introduces potential errors into estimates of ET by wide-area methods.     

Early impacts of biological control on canopy cover and water use of the invasive saltcedar tree (Tamarix spp.) in western Nevada, USA

The success of biological control programs is rarely assessed beyond population level impacts on the target organism. The question of whether a biological control agent can either partially or completely restore ecosystem services independent of population level control is therefore still open to discussion. Using observational and experimental approaches, we investigated the ability of the saltcedar leaf beetle [Diorhabda carinulata (Brullé) (Coleoptera: Chrysomelidae)] to reduce the water use of saltcedar trees (Tamarix ramosissima Ledeb.) in two sites (Humboldt and Walker Rivers) in Nevada, USA. At these sites D. carinulata defoliated the majority of trees within 25 and 9 km, respectively, of the release location within 3 years. At the Humboldt site, D. carinulata reduced the canopy cover of trees adjacent to the release location by >90%. At a location 4 km away during the first year of defoliation, D. carinulata reduced peak (August) stem water use by 50-70% and stand transpiration (July to late September) by 75% (P = 0.052). There was, however, no reduction in stem water use and stand transpiration during the second year of defoliation due to reduced beetle abundances at that location. At the Walker site, we measured stand evapotranspiration (ET) in the center of a large saltcedar stand and found that ET was highest immediately prior to D. carinulata arrival, dropped dramatically with defoliation, and remained low through the subsequent 2 years of the study. In contrast, near the perimeter of the stand, D. carinulata did not reduce sap flow, partly because of low rates of defoliation but also because of increased water use per unit leaf area in response to defoliation. Taken together, our results provide evidence that in the early stages of population expansion D. carinulata can lead to substantial declines in saltcedar water use. The extent of these declines varies spatially and temporally and is dependent on saltcedar compensatory responses along with D. carinulata population dynamics and patterns of dispersal.     

Molecular systematics and historical biogeography of tree boas (Corallus spp.)

Inferring the evolutionary and biogeographic history of taxa occurring in a particular region is one way to determine the processes by which the biodiversity of that region originated. Tree boas of the genus Corallus are an ancient clade and occur throughout Central and South America and the Lesser Antilles, making it an excellent group for investigating Neotropical biogeography. Using sequenced portions of two mitochondrial and three nuclear loci for individuals of all recognized species of Corallus, we infer phylogenetic relationships, present the first molecular analysis of the phylogenetic placement of the enigmatic C. cropanii, develop a time-calibrated phylogeny, and explore the biogeographic history of the genus. We found that Corallus diversified within mainland South America, via over-water dispersals to the Lesser Antilles and Central America, and via the traditionally recognized Panamanian land bridge. Divergence time estimates reject the South American Caribbean-Track as a general biogeographic model for Corallus and implicate a role for events during the Oligocene and Miocene in diversification such as marine incursions and the uplift of the Andes. Our findings also suggest that recognition of the island endemic species, C. grenadensis and C. cookii, is questionable as they are nested within the widely distributed species, C. hortulanus. Our results highlight the importance of using widespread taxa when forming and testing biogeographic hypotheses in complex regions and further illustrate the difficulty of forming broadly applicable hypotheses regarding patterns of diversification in the Neotropical region.     

A general model for mortality in adult tsetse (Glossina spp.)

Tsetse exhibit a U-shaped age-mortality curve, with high losses after eclosion and a well-marked ageing process, which is particularly dramatic in males. A three-parameter (k(1) -k(3) ) model for age-dependent adult instantaneous mortality rates was constructed using mark-recapture data for the tsetse fly Glossina morsitans morsitans Westwood (Diptera: Glossinidae). Mortality changed linearly with k(1) over all ages; k(2) affected only losses in roughly the first week of adult life, and k(3) controlled the ageing rate. Mortality pooled over age was twice as sensitive to changes in k(3) as in k(1) . Population growth rate was, however, similarly affected by these two parameters, reflecting the disproportionate effect of k(3) on mortality in the oldest flies that contribute least to the growth rate. Pooled-age mortality and growth rate were insensitive to changes in k(2) . The same model also provided good fits to data for laboratory colonies of female G. m. morsitans and Glossina austeni Newstead and should be applicable to all tsetse of both sexes. The new model for tsetse mortality should be incorporated into models of tsetse and trypanosome population dynamics; it will also inform the estimation of adult female mortality from ovarian dissection data.     

Diapause in the leaf beetle Diorhabda elongata (Coleoptera: Chrysomelidae), a biological control agent for tamarisk (Tamarix spp.)

The tamarisk leaf beetle, Diorhabda elongata Brullé deserticola Chen, was collected in northwestern China and has been released in the western United States to control tamarisk (Tamarix spp.). Characteristics of diapause and reproductive development in D. elongata were examined to improve management as a biocontrol agent. Under long days, 16:8 (L:D) h, males began to emit aggregation pheromone within 2-3 d of adult emergence, mating occurred, and females oviposited within 7 d of adult emergence. Under short days, 12:12 (L:D) h, males did not emit pheromone, mating did not occur, and both males and females entered reproductive diapause marked by inconspicuous gonads and hypertrophied fat body. Ovaries of diapausing females lacked vitellogenic oocytes, and the ovarioles were clear and narrow, whereas reproductive females had enlarged ovaries with two to three yellow oocytes per ovariole. Diapausing males had thin, transparent accessory glands and ejaculatory ducts, whereas reproductive males had thick white accessory glands and white opaque ejaculatory ducts. Sensitivity to diapause-inducing photoperiods extended into the adult stage. Reproductive females ceased oviposition, resorbed oocytes, and entered diapause when switched from long to short days. Diapause-destined insects ceased feeding and entered the leaf litter 10-20 d after adult emergence, whereas reproductive insects remained on the plants and fed for at least 30 d. Reproductive insects exhibited dispersal behaviors, such as attempted flights, whereas diapause-destined insects did not show dispersal behaviors. Information gained from these studies will be used to better manage populations in the field and to improve rearing and storage in the laboratory.     

Northern tamarisk beetle (Diorhabda carinulata) and tamarisk (Tamarix spp.) interactions in the Colorado River basin

Northern tamarisk beetles (Diorhabda carinulata) were released in the Upper Colorado River Basin in the United States in 2004–2007 to defoliate introduced tamarisk shrubs (Tamarix spp.) in the region's riparian zones. The primary purpose was to control the invasive shrub and reduce evapotranspiration (ET) by tamarisk in an attempt to increase stream flows. We evaluated beetle–tamarisk interactions with MODIS and Landsat imagery on 13 river systems, with vegetation indices used as indicators of the extent of defoliation and ET. Beetles are widespread and exhibit a pattern of colonize–defoliate–emigrate, so that riparian zones contain a mosaic of completely defoliated, partially defoliated, and refoliated tamarisk stands. Based on satellite data and ET algorithms, mean ET before beetle release (2000–2006) was 416 mm/year compared to postrelease (2007–2015) ET of 355 mm/year (p < 0.05) for a net reduction of 61 mm/year. This is lower than initial literature projections that ET would be reduced by 300–460 mm/year. Reasons for the lower‐than‐expected ET reductions are because baseline ET rates are lower than initially projected, and percentage ET reduction is low because tamarisk stands tend to regrow new leaves after defoliation and other plants help maintain canopy cover. Overall reductions in tamarisk green foliage during the study are 21%. However, ET in the Upper Basin has shown a steady decline since 2007 and equilibrium has not yet been reached. Defoliation is now proceeding from the Upper Basin into the Lower Basin at a rate of 40 km/year, much faster than initially projected.     

The effects of riparian restoration following saltcedar (Tamarix spp.) biocontrol on habitat and herpetofauna along a desert stream

Amphibians and reptiles (herpetofauna) have been linked to specific microhabitat characteristics, microclimates, and water resources in riparian forests. Our objective was to relate variation in herpetofauna abundance to changes in habitat caused by a beetle used for Tamarix biocontrol (Diorhabda carinulata; Coleoptera: Chrysomelidae) and riparian restoration. During 2013 and 2014, we measured vegetation and monitored herpetofauna via trapping and visual encounter surveys (VES) at locations affected by biocontrol along the Virgin River in the Mojave Desert of the southwestern United States. Twenty‐one sites were divided into four riparian stand types based on density and percent cover of dominant trees (Tamarix, Prosopis, Populus, and Salix) and presence or absence of restoration. Restoration activities consisted of mechanically removing non‐native trees, transplanting native trees, and restoring hydrologic flows. Restored sites had three times more total lizard and eight times more yellow‐backed spiny lizard (Sceloporus uniformis) captures than other stand types. Woodhouse's toad (Anaxyrus woodhousii) captures were greatest in unrestored and restored Tam‐Pop/Sal sites. Results from VES indicated that herpetofauna abundance was greatest in the restored Tam‐Pop/Sal site compared with the adjacent unrestored Tam‐Pop/Sal site. Tam sites were characterized by having high Tamarix cover, percent canopy cover, and shade. Restored Tam‐Pop/Sal sites were most similar in habitat to Tam‐Pop/Sal sites. Two species of herpetofauna (spiny lizard and toad) were found to prefer habitat components characteristic of restored Tam‐Pop/Sal sites. Restored sites likely supported higher abundances of these species because restoration activities reduced canopy cover, increased native tree density, and restored surface water.     

Seasonal adaptations to day length in ecotypes of Diorhabda spp. (Coleoptera: Chrysomelidae) inform selection of agents against saltcedars (Tamarix spp.)

Seasonal adaptations to daylength often limit the effective range of insects used in biological control of weeds. The leaf beetle Diorhabda carinulata (Desbrochers) was introduced into North America from Fukang, China (latitude 44° N) to control saltcedars (Tamarix spp.), but failed to establish south of 38° N latitude because of a mismatched critical daylength response for diapause induction. The daylength response caused beetles to enter diapause too early in the season to survive the duration of winter at southern latitudes. Using climate chambers, we characterized the critical daylength response for diapause induction (CDL) in three ecotypes of Diorhabda beetles originating from 36, 38, and 43° N latitudes in Eurasia. In a field experiment, the timing of reproductive diapause and voltinism were compared among ecotypes by rearing the insects on plants in the field. CDL declined with latitude of origin among Diorhabda ecotypes. Moreover, CDL in southern (<39° N latitude) ecotypes was shortened by more than an hour when the insects were reared under a fluctuating 35-15°C thermoperiod than at a constant 25°C. In the northern (>42° N latitude) ecotypes, however, CDL was relatively insensitive to temperature. The southern ecotypes produced up to four generations when reared on plants in the field at sites south of 38° N, whereas northern ecotypes produced only one or two generations. The study reveals latitudinal variation in how Diorhabda ecotypes respond to daylength for diapause induction and how these responses affect insect voltinism across the introduced range.     

Patterns of parasite infection in bumble bees (Bombus spp.) of Northern Virginia

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Host specificity of different populations of the leaf beetle Diorhabda elongata (Coleoptera: Chrysomelidae), a biological control agent of saltcedar (Tamarix spp.)

The leaf beetle, Diorhabda elongata (Brullé) sensu lato, was released in 2001 for the classical biological control of exotic saltcedars, a complex of invasive Tamarix species and hybrids. It did not establish at sites south of 37°N latitude where summer daylengths are below the critical photoperiod of the northern-adapted populations of the beetle that were released. Therefore, we assessed the host specificity of four D. elongata populations collected from more southern latitudes in the Old World (Tunisia, Crete, Uzbekistan, and Turpan, China). All populations were similar to each other and the previously released populations of D. elongata in their host specificity. Larval/pupal survival for all populations was 34–100% on Tamarix test plants, 0–76% on native Frankenia plants (both in the order Tamaricales), and 0% on the remaining 28 species of plants on which all the larvae died as 1st instars. D. elongata laid high numbers of eggs on saltcedar, generally fewer eggs on athel (a moderately valued evergreen species of Tamarix) except for Uzbekistan beetles, and few to no eggs on three species of Frankenia. Few to no adults were found on Frankenia plants which also were poor maintenance hosts. The release of any of the four D. elongata populations in the southern US and northern Mexico should pose no risk to plants outside the order Tamaricales and a low risk to native, non-target Frankenia plants. Athel may be less damaged than saltcedar.     

Nitrate removal and DO levels in batch wetland mesocosms: Cattail (Typha spp.) versus bulrush (Scirpus spp.)

Nitrate removal rates and dissolved oxygen (DO) levels were evaluated in small batch-mode wetland mesocosms with two different plant species, cattail (Typha spp.) and bulrush (Scirpus spp.), and associated mineral-dominated sediment collected from a mature treatment wetland. Nitrate loss in both cattail and bulrush mesocosms was first-order in nature. First-order volumetric rate constants (k_V) were 0.30 d⁻¹ for cattail and 0.21 d⁻¹ for bulrush and rates of nitrate loss were significantly different between plant treatments (p < 0.005). On an areal basis, maximum rates of nitrate removal were around 500 mg N/(m² d) early in the experiment when nitrate levels were high (> 15 mg N/L). Areal removal rates were on average 25% higher in cattail versus bulrush mesocosms. DO in mesocosm water was significantly higher in bulrush versus cattail (p < 0.001). DO in bulrush generally ranged between 0.5 and 2 mg/L, while DO in cattail mesocosms was consistently below 0.3 mg/L. Based on cumulative frequency analysis, DO exceeded 1 mg/L around 50% of the time in bulrush, but only 2% of the time in cattail. DO in bulrush exhibited a statistically significant diel cycle with DO peaks in the late afternoon and DO minimums in the early morning hours. Difference in nitrate removal rates between wetland plant treatments may have been due to differing plant carbon quality. Cattail litter, which has been shown in other studies to exhibit superior biodegradability, may have enhanced biological denitrification by fueling heterotrophic microbial activity, which in turn may have depressed DO levels, a prerequisite for denitrification. Our results show that the cattail is more effective than bulrush for treating nitrate-dominant wastewaters.