Photosynthesis and water‐use efficiency: A comparison between invasive (exotic) and non‐invasive (native) species

Abstract Invasive species have been hypothesized to out‐compete natives though either a Jack‐of‐all‐trades strategy, where they are able to utilize resources effectively in unfavourable environments, a master‐of‐some, where resource utilization is greater than its competitors in favourable environments, or a combination of the two (Jack‐and‐master). We examined the invasive strategy of Berberis darwinii in New Zealand compared with four co‐occurring native species by examining germination, seedling survival, photosynthetic characteristics and water‐use efficiency of adult plants, in sun and shade environments. Berberis darwinii seeds germinated more in shady sites than the other natives, but survival was low. In contrast, while germination of B. darwinii was the same as the native species in sunny sites, seedling survival after 18 months was nearly twice that of the all native species. The maximum photosynthetic rate of B. darwinii was nearly double that of all native species in the sun, but was similar among all species in the shade. Other photosynthetic traits (quantum yield and stomatal conductance) did not generally differ between B. darwinii and the native species, regardless of light environment. Berberis darwinii had more positive values of δ¹³C than the four native species, suggesting that it gains more carbon per unit water transpired than the competing native species. These results suggest that the invasion success of B. darwinii may be partially explained by combination of a Jack‐of‐all‐trades scenario of widespread germination with a master‐of‐some scenario through its ability to photosynthesize at higher rates in the sun and, hence, gain a rapid height and biomass advantage over native species in favourable environments.     

Photosynthesis, nitrogen allocation and specific leaf area in invasive Eupatorium adenophorum and native Eupatorium japonicum grown at different irradiances

The mechanisms underlying biological invasions are still not well elucidated. In this study, ecophysiological traits of invasive Eupatorium adenophorum and native E.   japonicum were compared at 10 irradiances in field. I hypothesized that the invader may allocate a higher fraction of leaf nitrogen (N) to photosynthesis and have higher light-saturated photosynthetic rate ( P _max) and specific leaf area (SLA) than E.   japonicum . The invader had a significantly higher ability to acclimate to high irradiance than E.   japonicum , while it showed a similar shade-tolerant ability. The invader indeed allocated a higher fraction of leaf N to photosynthesis than E.   japonicum , which, with its high leaf N content ( N _A), resulted in a higher N content in photosynthesis ( N _P), contributing to its higher biochemical capacity for photosynthesis and P _max. However, the invader had a significantly lower SLA than E.   japonicum , contributing to its higher P _max but increasing its area-based leaf construction cost. The abilities to acclimate to a wider range of irradiance and to allocate a higher fraction of leaf N to photosynthesis, and the higher P _max, N _A, N _P and leaf area ratio may contribute to the invasion of the invader. High SLA is not always necessary for invasive species.     

氮对苹果幼树水分利用效率的影响

以２年生盆栽新红星／平邑甜茶苹果树为试材,初步探讨了土壤不同水分状况下氮肥对植株水分利用效率（ＷＵＥ）及有关参数的影响,结果表明,充足供水时,随施Ｎ量的增加,植株ＷＵＥ降低,施Ｎ导致气孔导率（Ｇｓ）增大,对蒸腾（Ｔｒ）的提高幅度大于光合（Ｐｎ）;供水不足时,施Ｎ植株的ＷＵＥ明显高于对照,不同施Ｎ水平的ＷＵＥ表现为：高Ｎ〉中Ｎ〉低Ｎ,ＷＵＥ改善是由于叶肉羧化能力提高,导致光合增强。     

Invasive feral pigs impact native tree ferns and woody seedlings in Hawaiian forest

Invasive mammals can fundamentally alter native plant communities, especially on isolated islands where plants evolved without them. The globally invasive feral pig (Sus scrofa) can be particularly destructive to native plant communities. Tree ferns are an important understory component in many forests facilitating the establishment of a variety of species. However, the extent and effects of feral pig damage to tree ferns, and associated impacts on plant community regeneration, are largely unknown. We quantified the effect that feral pig damage has on tree fern growth, survival, and epiphytic woody seedling abundance over 1 year on 438 randomly selected tree ferns of three endemic species (Cibotium chamissoi, Cibotium glaucum, and Cibotium menziesii) in a Hawaiian montane wet forest with high tree fern and feral pig densities. Across all tree fern species, feral pigs damaged 13 % of individuals over 1 year. Compared with undamaged tree ferns, moderately- to heavily-damaged individuals had decreases of 4 to 27 % in trunk length increment and lost tenfold more fronds. Tree fern angle (standing, leaning, prone, or semi-prone) and woody seedling abundance co-varied with feral pig damage. Specifically, damaged tree ferns were more often prone or semi-prone and supported more seedlings, but also had annual mortality up to 34 % higher than undamaged tree ferns. Overall, feral pig damage had substantial negative effects on tree ferns by reducing growth and survival. Given the importance of tree ferns as regeneration sites for a variety of native plants, feral pig damage to tree ferns will likely alter future forest composition and structure. Specifically, feral pig damage to tree ferns reduces potential establishment sites for species that either regenerate preferentially as epiphytes or are currently restricted to epiphytic establishment due to ground rooting by feral pigs.     

Responses to nitrogen pulses and growth under low nitrogen availability in invasive and native tree species with differing successional status

Invasive species are frequently found in recently disturbed sites. To examine how these disturbance-dependent invasive species exploit resource pulses resulting from disturbance, twelve physiological and morphological traits, including age-dependent responsiveness in leaf traits to nitrogen pulse, were compared between Bischofia javanica, an invasive tree species in Ogasawara islands, and three native Ogasawara species, each having a different successional status. When exposed to a nitrogen pulse, invasive B. javanica showed higher increases in photosynthetic capacity, leaf area, epidermal cell number and cell size in leaves of broad age classes, and root nitrogen absorption ability than two native mid-/late or late-successional species, but showed no particular superiority to a native pioneer species in these responses. Under low nitrogen, however, it showed the largest relative growth rate among the four species, while the native pioneer showed the lowest growth. From these results, we concluded that the combination of moderately high responsiveness to resource pulses and the ability to maintain steady growth under resource limitations may give B. javanica a competitive advantage over a series of native species with different successional status from early to late-successional stages.     

Nitrogen allocation,partitioning and use efficiency in three invasive plant species in comparison with their native congeners

In this study, we hypothesized that invasive species may allocate a higher fraction of leaf nitrogen (N) to photosynthesis than phylogenetically related native species. To test this hypothesis, we determined N allocation and other ecophysiological traits of three invasive species in comparison with their respective native congeners by measuring response curves of photosynthesis to intercellular CO₂ concentration. The invasive species of Peperomia and Piper indeed allocated a higher fraction of leaf N to photosynthesis and were more efficient in photosynthetic N (N _P) partitioning than their native congeners. The two invasive species partitioned a higher fraction of N _P to carboxylation and showed a higher use efficiency of N _P, while their native congeners partitioned a higher fraction of N _P to light-harvesting components. The higher N allocation to photosynthesis and the higher N _P partitioning to carboxylation in the two invaders were associated with their higher specific leaf area. Nitrogen allocation and partitioning were the most important factors in explaining the differences in light-saturated photosynthetic rate and photosynthetic N use efficiency (PNUE) between the two invasive species and their native congeners. The differences in N allocation-related variables between the invasive and native species of Amaranthus could not be evaluated in this study due to the method. Except PNUE, resource capture- and use-related traits were not always higher in all three invasive species compared to their native congeners, indicating that different invasive species may have different syndrome of traits associated with its invasiveness.     

Water use and water-use efficiency of the invasive Centaurea maculosa and three native grasses

The Eurasian herb Centaurea maculosa Lam. has invaded millions of hectares of semi-arid grasslands in western North America. Its success may reflect that it may be more competitive than native species, it is not grazed by large herbivores, it was introduced without its native enemies, it may interfere with native species via allelopathy, or most likely some combination of these factors. Greater competitive ability could include greater use of limiting soil resources, such as water, or more efficient use of soil water, thereby inhibiting establishment, survival, and reproduction of native species. We measured water use and water-use efficiency of Centaurea and three native grasses, Pseudoroegneria spicata [Scribn. and Smith] A. Love, Pascopyrum smithii [Rybd.] A. Love, and Festuca idahoensis Elmer, in a glasshouse. Water-use efficiency was determined by the traditional measure of biomass produced per mass of water used, and by carbon-isotope discrimination (). Centaurea did not use the most water, or use water more efficiently (based on biomass (g)/ water (kg) and carbon-isotope discrimination) than all three native grasses. We also determined carbon-isotope discrimination of Centaurea and dominant native grasses during the 1999 and 2000 growing seasons at three field sites. Centaurea rosettes had the lowest water-use efficiency (greatest carbon-isotope discrimination), followed by mature plants of Centaurea, and then native grasses. Water-use efficiency of mature Centaurea plants and native grasses was greater in late summer than early summer. Centaurea's success as an invasive species in North America cannot be attributed to greater use of soil water or greater water-use efficiency than native grasses.     

An invasive tree fern alters soil and plant nutrient dynamics in Hawaii

Invasive species that alter ecosystems are often successful competitors due to their effects on nutrient cycling. Sphaeropteris cooperi (Cyatheaceae; Australian tree fern) has been invading intact Hawaiian rainforests for decades and displacing the dominant native tree fern Cibotium glaucum (Cibotiaceae). S. cooperi produces more leaves that grow faster, contain more N and P, and decompose faster than C. glaucum leaves. Our experiment tested the effects of additions of leaf litter from native and non-native tree ferns on the growth and nutrient content of four native angiosperm species in forest (N-rich) and landslide (P-rich) soils. Both litter treatments inhibited growth initially in all species, but subsequent responses were species-specific. Compared to control treatments, the increase in biomass was highest in the fast-growing Carex wahuensis and Hibiscus arnottianus with S. cooperi litter in landslide soil. Leaf N in C. wahuensis was higher with S. cooperi litter and in forest soil, as expected, but other leaf nutrient responses showed some evidence of nutrient immobilization from litter addition. Several growth measures were higher with S. cooperi than C. glaucum litter and in forest than landslide soil, suggesting that N availability is the strongest driver of growth. Our results show that S. cooperi can alter nutrient cycling in Hawaiian plants, sometimes with positive effects on growth. However, under natural conditions, native plants must compete for these additional nutrients with S. cooperi and other invasive species. This study contributes to invasion biology as the first to examine the impact of leaf litter of an invasive fern on native species.     

Photosynthesis and photoinhibition after night chilling in seedlings of two tropical tree species grown under three irradiances

We investigated the physiological effect of night chilling (CN) on potted seedlings of two tropical tree species, Calophyllum polyanthum and Linociera insignis, in Xishuangbanna, southwest China. Seedlings grown under 8, 25, and 50 % daylight for five months were moved to a 4–6 °C cold storage house for three consecutive nights, and returned to the original shaded sites during the day. CN resulted in strong suppression of photosynthesis and stomatal conductance for L. insignis, and reduced photorespiration rates, carboxylation efficiency, and maximum photochemical efficiency of photosystem 2 (PS2) at dawn and midday for both species. CN increased dawn and midday rates of non-photochemical quenching, and the contents of malondialdehyde and H₂O₂ for both species. CN also induced inactivation or destruction of PS2 reaction centres. The impacts of CN on tropical seedlings increased with the number of CN. Shading could significantly mitigate the adverse effects of CN for both species. After 3-d-recovery, gas exchange and fluorescence parameters for both species returned to pre-treatment levels in most cases. Thus CN induced mainly stomatal limitation of photosynthesis for L. insignis, and non-stomatal limitation for C. polyanthum. C. polyanthum was more susceptible to CN than L. insignis. Fog, which often occurs in Xishuangbanna, could be beneficial to chilling sensitive tropical seedlings in this area through alleviating photoinhibition or photodamage by reducing sunlight.     

Shade mitigates photoinhibition and enhances water use efficiency in kiwifruit under drought

We tested the hypotheses that a reduction of incident light of 50 % over sun-acclimated leaves of water stressed kiwifruit (Actinidia deliciosa var. deliciosa) would (1) reduce stomatal limitations to carbon supply and (2) mitigate the inactivation of the primary photochemistry associated with photosystem (PS) II, thereby this increases carbon gain and water-use efficiency (WUE). Groups of field-grown vines were either shaded or left naturally exposed and subjected to progressive water stress in order to study moderately and severely droughted vines, while other groups were well irrigated. Daily variations in leaf gas exchange and midday chlorophyll (Chl) a fluorescence were determined once plants had −0.6 MPa (moderate stress) and −1.0 MPa pre-down leaf water potential (severe stress). Variations in Chl pigment content and specific leaf area (SLA) are also discussed. Results reveal that 50 % shade application maintained efficiency of PSII close to 0.8 even under severe drought so that to prevent its large decline (0.65) recorded in sunlit leaves. Under moderate stress level stomata behaviour dominated upon metabolic impairments of PSII. Reduction of irradiance increased WUE (15–20 %) in droughted vines, representing a valuable intervention to save photosynthetic apparatus and improve WUE in vines experiencing typical Mediterranean summer stresses.     

Photosynthesis,photoinhibition and low temperature acclimation in cold tolerant plants

Cold acclimation requires adjustment to a combination of light and low temperature, conditions which are potentially photoinhibitory. The photosynthetic response of plants to low temperature is dependent upon time of exposure and the developmental history of the leaves. Exposure of fully expanded leaves of winter cereals to short-term, low temperature shiftsinhibits whereas low temperature growthstimulates electron transport capacity and carbon assimilation. However, the photosynthetic response to low temperature is clearly species and cultivar dependent. Winter annuals and algae which actively grow and develop at low temperature and moderate irradiance acquire a resistance to irradiance 5- to 6-fold higher than their growth irradiance. Resistance to short-term photoinhibition (hours) in winter cereals is a reflection of the increased capacity to keep Q_A oxidized under high light conditions and low temperature. This is due to an increased capacity for photosynthesis. These characteristics reflect photosynthetic acclimation to low growth temperature and can be used to predict the freezing tolerance of cereals. It is proposed that the enhanced photosynthetic capacity reflects an increased flux of fixed carbon through to sucrose in source tissue as a consequence of the combined effects of increased storage of carbohydrate as fructans in the vacuole of leaf mesophyll cells and an enhanced export to the crown due to its increased sink activity. Long-term exposure (months) of cereals to low temperature photoinhibition indicates that this reduction of photochemical efficiency of PS II represents a stable, long-term down regulation of PS II to match the energy requirements for CO₂ fixation. Thus, photoinhibition in vivo should be viewed as the capacity of plants to adjust photosynthetically to the prevailing environmental conditions rather than a process which necessarily results in damage or injury to plants. Not all cold tolerant, herbaceous annuals use the same mechanism to acquire resistance to photoinhibition. In contrast to annuals and algae, overwintering evergreens become dormant during the cold hardening period and generally remain susceptible to photoinhibition. It is concluded that the photosynthetic response to low temperatures and susceptibility to photoinhibition are consequences of the overwintering strategy of the plant species.     

A decision tree approach to modelling nitrogen fertiliser use efficiency in New Zealand pastures

Quantifying nitrogen (N) fertiliser use efficiency (NFUE) in pastoral systems has important implications for fertiliser management from both economic and environmental points of view. The potential of a decision tree approach for modelling NFUE in New Zealand pastures was investigated. The decision tree model suggested that the time of applying N fertiliser was the most important factor influencing NFUE, with August or September (early spring in New Zealand) being the best time of application. The interaction of rainfall and temperature, rainfall, phosphorus (P) fertiliser history, soil Olsen P and slope were other important factors influencing NFUE. The model was validated for 11 of the 16 trials tested with a predictive accuracy of 69%. The mechanisms by which these factors influenced NFUE and the uncertainty associated with the model prediction were discussed. It was concluded that this type of modelling approach can be used to predict NFUE and thereby to assist decisions on when and where to apply N fertiliser in pastures for increasing productivity while reducing the environmental impact.     

Leaf construction cost, nutrient concentration, and net CO2 assimilation of native and invasive species in Hawaii

To examine the predictability of leaf physiology and biochemistry from light gradients within canopies, we measured photosynthetic light-response curves, leaf mass per area (LMA) and concentrations of nitrogen, phosphorus and chlorophyll at 15–20 positions within canopies of three conifer species with increasing shade tolerance, ponderosa pine [Pinus ponderosa (Laws.)], Douglas fir [Pseudotsuga menziesii (Mirb.) Franco], and western hemlock [Tsuga heterophylla (Raf.) Sarg.]. Adjacent to each sampling position, we continuously monitored photosynthetically active photon flux density (PPFD) over a 5-week period using quantum sensors. From these measurements we calculated FPAR: integrated PPFD at each sampling point as a fraction of full sun. From the shadiest to the brightest canopy positions, LMA increased by about 50% in ponderosa pine and 100% in western hemlock; Douglas fir was intermediate. Canopy-average LMA increased with decreasing shade tolerance. Most foliage properties showed more variability within and between canopies when expressed on a leaf area basis than on a leaf mass basis, although the reverse was true for chlorophyll. Where foliage biochemistry or physiology was correlated with FPAR, the relationships were non-linear, tending to reach a plateau at about 50% of full sunlight. Slopes of response functions relating physiology and biochemistry to ln(FPAR) were not significantly different among species except for the light compensation point, which did not vary in response to light in ponderosa pine, but did in the other two species. We used the physiological measurements for Douglas fir in a model to simulate canopy photosynthetic potential (daily net carbon gain limited only by PPFD) and tested the hypothesis that allocation of carbon and nitrogen is optimized relative to PPFD gradients. Simulated photosynthetic potential for the whole canopy was slightly higher (<10%) using the measured allocation of C and N within the canopy compared with no stratification (i.e., all foliage identical). However, there was no evidence that the actual allocation pattern was optimized on the basis of PPFD gradients alone; simulated net carbon assimilation increased still further when even more N and C were allocated to high-light environments at the canopy top. Received: 12 August 1998 / Accepted: 25 March 1999     

氮肥运筹对小麦产量、氮素利用效率和光能利用率的影响

连续2年在西南冬麦区的重庆、仁寿、广汉、西昌4个地点,开展3种施氮水平(每公顷纯氮0、120、180 kg,简写为N₀、N₁₂₀、N₁₈₀)和3种氮肥分配模式(N_A:底肥100%;N_B:底肥70%+苗肥30%;N_C:底肥60%+拔节肥40%)的田间试验,监测小麦花后冠层叶片SPAD值、群体光合速率(CAP)、光能利用等生理参数和籽粒产量,计算氮素利用效率、光能利用率等.结果表明: 随施氮水平增加,小麦上三叶SPAD值、CAP、光合有效辐射(PAR)截获率和产量均呈增加趋势,而氮肥农学利用效率、生产效率、吸收效率和利用效率呈降低趋势.氮肥后移的增效作用因施氮水平而异,SPAD于N₁₈₀增效明显,而CAP于N₁₂₀增效明显,不同氮肥管理模式的光能利用率因地点而异.氮肥后移能明显提高小麦氮肥农学效率、生产效率、吸收效率和氮素表观回收率,但氮肥利用效率则略有减少.氮肥后移效果N_C总体优于N_B处理.不同地点比较,广汉的SPAD值、CAP、PAR截获率、氮肥利用参数较高,其产量也相应最高;西昌的产量、SPAD值及氮素利用效率较高,但其光能利用率和CAP较低;重庆和仁寿的SPAD值、光能利用率及氮素利用效率均较低,其产量也最低.小麦生物产量与各地点的籽粒产量、CAP、SPAD值和PAR累积截获量均呈显著或极显著的正相关关系.表明不同生态区域增施氮肥都能促进小麦增产,氮肥后移可进一步优化产量结构、改善氮肥和光能利用效率,但存在年份和地点差异,各地需要制定有针对性的氮肥管理模式.     

Differences in nitrogen use strategies between native and exotic tree species: predicting impacts on invaded ecosystems

Background and aim

Exotic plant species can alter the nitrogen cycle in invaded ecosystems. We assess the differences in nitrogen use strategies and litter production and dynamics among three native riparian trees (Fraxinus angustifolia, Populus alba and Ulmus minor) and three co-occurring exotics (Ailanthus altissima, Robinia pseudoacacia and Ulmus pumila), currently spreading throughout river banks in inner Spain. We aim to predict the ecological consequences of a replacement of the natives by the exotics.

Methods

We compared the leaf lifespan, nitrogen resorption efficiency in leaves, nitrogen mean residence time, amount and timing of litter production and amount of nitrogen returned to soils between these native and exotic species.

Results

We found differences among species in all the variables measured, but not between native or exotic origins. Species were ranked from the most to the least conservative nitrogen use strategy as follows: U. pumila was the most conservative species, followed by the three natives (with an intermediate strategy), A. altissima and finally by the nitrogen-fixer R. pseudoacacia. The studied exotic species would produce contrasting impacts on the nitrogen cycle upon invasion.

Conclusions

On the basis of our results, we predict little impacts on the nitrogen cycle if U. pumila dominates the landscape. Despite being nitrogen-fixer R. pseudocacia would not increase soil nitrogen availability in the study area due to its low litter production and litter decomposition rates. In contrast, we predict an increase in nitrogen availability of soils upon A. altissima invasion, as this species produces a high amount of nitrogen rich and labile litter. This study offers a striking example of the contingencies involved in predicting the ecosystem impacts of exotic plant invasion.     

Photosynthesis and photoinhibition in two xerophytic shrubs during drought

Seasonal changes in water relations, net photosynthetic rate (P _N), and fluorescence of chlorophyll (Chl) a of two perennial C₃ deciduous shrubs, Ipomoea carnea and Jatropha gossypifolia, growing in a thorn scrub in Venezuela were studied in order to establish the possible occurrence of photoinhibition during dry season and determine whether changes in photochemical activity of photosystem 2 (PS2) may explain variations of P _N in these species. Leaf water potential () decreased from –0.2 to –2.1 MPa during drought in both species. The P _N decreased with in I. carnea and J. gossypifolia by 64 and 74 %, respectively. Carboxylation efficiency (CE) decreased by more than 50 and 70 % in I. carnea and J. gossypifolia, respectively. In I. carnea, relative stomatal limitation (L_s) increased by 17 % and mesophyll limitation (L_m) by 65 % during drought, while in J. gossypifolia L_s decreased by 27 % and L_m increased by 51 %. Drought caused a reduction in quantum yield of PS2 (_PS2) in both species. Drought affected the capacity of energy dissipation of leaves, judging from the changes in the photochemical (q_P) and non-photochemical quenching (NPQ) coefficients. Photoinhibition during drought in I. carnea and J. gossypifolia was evidenced in the field by a drop in the maximum quantum yield of PS2 (F_v/F_m) below 0.8 and also by non-coordinated changes in _PS2 and quantum yield of non-photochemical excitation quenching (Y_n). Total soluble protein content on an area basis increased with but the ribulose-1,5-bisphosphate carboxylase/oxygenase content remained unchanged. A reduction of total Chl content with drought was observed. Hence in the species studied photoinhibition occurred, which imposed an important limitation on carbon assimilation during drought.     

Congeneric invasive versus native plants utilize similar inorganic nitrogen forms but have disparate use efficiencies

入侵植物与同属本地植物利用相似的无机氮但具有不同的利用效率 土壤无机氮对外来植物入侵具有重要作用。虽然沿完整无机氮梯度研究入侵植物与本地植物的氮利用策略对理解入侵成功很有必要,但相关研究依然相对不足。为此,我们选择6对入侵植物和同属本地植物(即12种植物)开展盆栽实验,设置11种硝铵比(每种比例包含低水平和高水平)形成完整的氮梯度(一端完全是硝态氮而另一端完全是铵态氮)。本实验中每种植物-氮组合重复8次,共栽培2112株植物。我们测定生物量及其分配、生长优势、叶绿素含量和低氮耐受性。结果显示,入侵植物和本地植物在任何一种硝铵比条件下均能正常生长,两者的生长、分配和耐受性反应随硝铵比梯度接近平行。入侵植物比本地植物长得更大,具有更高的叶绿素含量、根生物量分配和低氮耐受性。这些结果表明,入侵植物与同属本地植物利用相似的无机氮形态但具有不同的利用效率。这些氮利用对策在一定程度上有助于外来植物入侵。     

Stoichiometry of ferns in Hawaii: implications for nutrient cycling

We asked if element concentrations in ferns differ systematically from those in woody dicots in ways that could influence ecosystem properties and processes. Phylogenetically, ferns are deeply separated from angiosperms; for our analyses we additionally separated leptosporangiate ferns into polypod ferns, a monophyletic clade of ferns which radiated after the rise of angiosperms, and all other leptosporangiate (non-polypod) ferns. We sampled both non-polypod and polypod ferns on a natural fertility gradient and within fertilized and unfertilized plots in Hawaii, and compared our data with shrub and tree samples collected previously in the same plots. Non-polypod ferns in particular had low Ca concentrations under all conditions and less plasticity in their N and P stoichiometry than did polypod ferns or dicots. Polypod ferns were particularly rich in N and P, with low N:P ratios, and their stoichiometry varied substantially in response to differences in nutrient availability. Distinguishing between these two groups has the potential to be useful both in and out of Hawaii, as they have distinct properties which can affect ecosystem function. These differences could contribute to the widespread abundance of polypod ferns in an angiosperm-dominated world, and to patterns of nutrient cycling and limitation in sites where ferns are abundant.     

Complex impacts of an invasive omnivore and native consumers on stream communities in California and Hawaii

The effects of invasive species on native communities often depend on the characteristics of the recipient community and on the food habits of the invasive species, becoming complicated when the invader is omnivorous. In field enclosure experiments, we assessed the direct and interactive effects of an invasive omnivorous crayfish (Procambarus clarkii) and either native herbivorous snails (Physella gyrina) or shrimp (Atyoida bisulcata) on stream communities in California and Hawaii, respectively. Based on literature data and the characteristics of each study site, we predicted that crayfish would affect primarily algal-based trophic linkages in an open California stream but detritus-based trophic linkages in a shaded Hawaiian stream, with trophic cascades mediated through crayfish effects on primary consumers being observed in both systems. As predicted, crayfish in California directly reduced periphyton, filamentous algae, sediment, and snail levels, but generated a cascade by decreasing snail densities and increasing periphyton biomass. Contrary to prediction, crayfish did not reduce total invertebrate biomass. As predicted, crayfish in Hawaii reduced leaf litter, filamentous algae, and benthic invertebrate biomass. Contrary to our predictions, however, a trophic cascade was not observed because shrimp did not affect periphyton levels, crayfish did not reduce shrimp abundance, and crayfish had greater negative impacts on filamentous algae than did shrimp. Our findings highlight that the same invasive species can generate different effects on disparate systems, probably as mediated through the availability of different food types, flexibility in the invasive species’ food habits, and complex pathways of trophic interaction.