Upward cascading effects of nutrients: shifts in a benthic microalgal community and a negative herbivore response

We evaluated the effects of nutrient addition on interactions between the benthic microalgal community and a dominant herbivorous gastropod, Cerithidea californica (California horn snail), on tidal flats in Mugu Lagoon, southern California, USA. We crossed snail and nutrient (N and P) addition treatments in enclosures on two tidal flats varying from 71 to 92% sand content in a temporally replicated experiment (summer 2000, fall 2000, spring 2001). Diatom biomass increased slightly (~30%) in response to nutrient treatments but was not affected by snails. Blooms of cyanobacteria (up to 200%) and purple sulfur bacteria (up to 400%) occurred in response to nutrient enrichment, particularly in the sandier site, but only cyanobacterial biomass decreased in response to snail grazing. Snail mortality was 2–5 times higher in response to nutrient addition, especially in the sandier site, corresponding to a relative increase in cyanobacterial biomass. Nutrient-related snail mortality occurred only in the spring and summer, when the snails were most actively feeding on the microalgal community. Inactive snails in the fall showed no response to nutrient-induced cyanobacterial growths. This study demonstrated strongly negative upward cascading effects of nutrient enrichment through the food chain. The strength of this upward cascade was closely linked to sediment type and microalgal community composition.     

The Distribution of an Introduced Mollusc and its Role in the Long-term Demise of a Native Confamilial Species

The estuarine mud snail, Batillaria attramentaria, was transported to the Pacific coast of North America with the Pacific oyster, Crassostrea gigas (C. gigas), imported from Japan in the early part of this century, and has proliferated in several bays where the oyster was introduced. Since the arrival of Batillaria there have been declines in populations of its native ecological equivalent, Cerithidea californica. This study documents the distribution of the exotic Batillaria throughout its entire introduced range, concentrating on the few bays in northern California where both snails exist sympatrically. Using dates of initial importation of C. gigas and dates of first documentation of Batillaria within a bay, I established the earliest possible date for the introduction of Batillaria in a particular area. In cases where Cerithidea also was, or had been, present within a bay, I calculated a range of time for either the continued coexistence of the two species or the time until local exclusion of the native. Density measurements of Cerithidea within these bays where the species co-occurred allowed comparison of present Cerithidea numbers to historical accounts. Results indicated that Batillaria is replacing Cerithidea in the northern marshes of California. This replacement of the northernmost Cerithidea populations is not only reducing Cerithidea's overall range, but also eliminating a race recently shown to be a genetically distinct from southern Cerithidea populations. Other studies that have demonstrated superior exploitative competitive ability by Batillaria provide a potential mechanism for this displacement pattern. Regardless, the results presented here indicate that the displacement process is slow, taking on average >50 years to complete. This study illustrates a gradual, but predictable process of exotic replacement of native species, and argues strongly against complacency toward invaders that may currently seem innocuous. This revised version was published online in July 2006 with corrections to the Cover Date.     

Release from Parasites as Natural Enemies: Increased Performance of a Globally Introduced Marine Crab

Introduced species often seem to perform better than conspecifics in their native range. This is apparent in the high densities they may achieve or the larger individual sizes they attain. A prominent hypothesis explaining the success of introduced terrestrial species is that they are typically free of or are less affected by the natural enemies (competitors, predators, and parasites) they encounter in their introduced range compared to their native range. To test this hypothesis in a marine system, we conducted a global assessment of the effect of parasitism and predation on the ecological performance of European green crab populations. In Europe, where the green crab is native, crab body size and biomass were negatively associated with the prevalence of parasitic castrators. When we compared native crab populations with those from introduced regions, limb loss (an estimator of predation) was not significantly lower in introduced regions, parasites infected introduced populations substantially less and crabs in introduced regions were larger and exhibited a greater biomass. Our results are consistent with the general prediction that introduced species suffer less from parasites compared to populations where they are native. This may partly explain why the green crab is such a successful invader and, subsequently, why it is a pest in so many places.     

Effects of an Introduced Piscivore on Native Trout: Insights from a Demographic Model

Introductions of exotic species pose a significant threat to the persistence of many native populations, including many inland fishes. In 1994, piscivorous lake trout (Salvelinus namaycush) were discovered in Yellowstone Lake, Yellowstone National Park, Wyoming, USA, one of the last strongholds of the native Yellowstone cutthroat trout (Oncorhynchus clarki bouvieri). Predation by lake trout is expected to lead to a substantial decline in the native cutthroat trout population, which may have significant negative consequences for terrestrial predators that depend on cutthroat trout for prey and for the recreational fishery of the Park. We developed a matrix demographic model for the cutthroat trout population in Yellowstone Lake to identify the life stages that are most critical for understanding population dynamics. Parameter estimates (vital rates) were manipulated to explore the possible consequences of lake trout invasion. Comparisons of our results with current estimates of population trend and age structure suggested that our model reflected current conditions of the system. Elasticity analysis of the model revealed that population growth was most sensitive to annual survival of young trout, the group that is expected to be most vulnerable to lake trout predation. Projection of our deterministic model suggested that, in addition to a decline in abundance of cutthroat trout, the effects of lake trout may be manifest as changes in age and breeding structure of the population. Simulations of a stochastic version of the model indicated that a 60% or greater decline in the cutthroat trout population could be expected within 100 years if the lake trout population were permitted to grow uncontrolled. However, an effective control strategy that prevented the establishment of a large population of lake trout substantially reduced population decline, although the reduction in the availability of adult trout to terrestrial predators and anglers may be still be substantial (20–40%). In addition to current control activities in place in the Park, we recommend a renewed emphasis on understanding and monitoring juvenile life stages of cutthroat trout. Our results demonstrate the value of existing data sets for developing models to estimate the potential impact of biological invasions on the management and conservation of native populations, especially when opportunities and resources for additional empirical studies are limited.     

Parasitism of orangestriped oakworm (Lepidoptera: Saturniidae) eggs in the urban landscape

Orangestriped oakworm, Anisota senatoria (J. E. Smith), has caused widespread defoliation of oak trees in the urban landscape of southeastern Virginia since 1985. Egg masses were collected from 1988 to 1990 to determine the impact of native egg parasites on A. senatoria populations. The most abundant egg parasite was Aprostocetus new sp. and mean egg mass parasitism was 24.6%. The eupelmid Anastatus hirtus (Ashmead), a new host record, parasitized a mean of 11.7% of A. senatoria egg masses. The encyrtid Ooencytrus sp., a new host record, had a mean egg mass parasitism of 0.09%. Inundative releases of Trichogramma minutum (Riley) in 1989 and 1990 did not increase parasitism rates and mean egg mass parasitism was 2.3%. Parasitism of first generation A. senatoria egg masses was higher compared with second generation. The four egg parasites collected in this study parasitized 30% of A. senatoria egg masses and within egg mass parasitism was 7.9%. These relatively low parasitism rates may partially explain the presence of consistently high A. senatoria populations in southeastern Virginia.     

The Asian Fish Tapeworm Bothriocephalus acheilognathi: a Potential Threat to Native Freshwater Fish Species in Mexico

Introduction of the parasite Bothriocephalus acheilognathi Yamaguti, 1934 with the herbivorous carp Ctenopharyngodon idellus, and other cyprinids, has been documented in almost every continent, except Antarctica. This study presents the current geographical distribution of this cestode in the freshwater fish of Mexico, highlighting infections in autochthonous and endemic species. It compiles existing information and presents original data. B. acheilognathi is widely dispersed among the freshwater fish of Mexico, being recorded to date in 49 fish species from 26 genera, 7 families and 5 orders. B. acheilognathi is reported from Mexico for the first time in Dionda ipni, Notropis celayensis, Yuriria alta, Gambusia vittata, Poecilia butleri, P. mexicana, Poeciliopsis baenschi, Poeciliopsis sp., Cichlasoma cyanoguttatum and C. labridens. The new hosts identified in this study bring the total number of known host species to 102 in 14 families and 7 orders of freshwater fishes around the world. Given its wide distribution among Mexican freshwater fish species, the abundance of the parasite and its high pathogenicity, parasitological data for B. acheilognathi should be considered as an important factor in native fish conservation policies. Biological changes in the freshwater habitats in Mexico caused by the introduction of exotic fish species and their parasites are virtually ubiquitous and extremely difficult to eradicate once established. As such, they should be considered as one of the most serious threats to native fish conservation.     

Social organization in a flatworm: trematode parasites form soldier and reproductive castes

In some of the most complex animal societies, individuals exhibit a cooperative division of labour to form castes. The most pronounced types of caste formation involve reproductive and non-reproductive forms that are morphologically distinct. In colonies comprising separate or mobile individuals, this type of caste formation has been recognized only among the arthropods, sea anemones and mole-rats. Here, we document physical and behavioural caste formation in a flatworm. Trematode flatworm parasites undergo repeated clonal reproduction of ‘parthenitae’ within their molluscan hosts forming colonies. We present experimental and observational data demonstrating specialization among trematode parthenitae to form distinct soldier and reproductive castes. Soldiers do not reproduce, have relatively large mouthparts, and are much smaller and thinner than reproductives. Soldiers are also more active, and are disproportionally common in areas of the host where invasions occur. Further, only soldiers readily and consistently attack heterospecifics and conspecifics from other colonies. The division of labour described here for trematodes is strongly analogous to that characterizing other social systems with a soldier caste. The parallel caste formation in these systems, despite varying reproductive mode and taxonomic affiliation, indicates the general importance of ecological factors in influencing the evolution of social behaviour. Further, the ‘recognition of self’ and the defence of the infected host body from invading parasites are comparable to aspects of immune defence. A division of labour is probably widespread among trematodes and trematode species encompass considerable taxonomic, life history and environmental diversity. Trematodes should therefore provide new, fruitful systems to investigate the ecology and evolution of sociality.     

Gringos En El Bosque: Introduced Tree Invasion in a Native Nothofagus/Austrocedrus Forest

We studied invasion into native Nothofagus/Austrocedrus forest by many introduced tree species planted between 1910 and 1940 in plantations near the center of Isla Victoria, in northern Patagonia. We located virtually all individuals of these species in 30 ha of forest in two series of transects at increasing distances from the plantations. Although these species included many reported as highly invasive elsewhere, we found little evidence for invasion on Isla Victoria, with many invasive species utterly failing to invade native forest. There was a notable decline with distance in number of introduced individuals, but wind direction appeared to be unimportant. Pseudotsuga menziesii and Juniperus communis were the only 2 species represented by many individuals, while 4 pine species plus Araucaria araucana were far less numerous and 6 other species were found fewer than 10 times each. Even those species found repeatedly were represented overwhelmingly by small individuals, and the great majority of introduced individuals were found not in native forest proper, but in somewhat open areas such as road verges, small remnant pastures, and deer trails. Invasion may be occurring, but too slowly to be clearly evidenced yet because of the longevity of the dominant native trees. Factors that may be stopping or slowing invasion include competition in gaps with native species, browsing by introduced deer, unfavorable soil, allelopathy, and natural enemies or other idiosyncratic factors for particular species. The absence of substantial invasion so far is no guarantee against future invasion, particularly if some major natural or anthropogenous disturbance were to occur.     

Effect of Two Opiine Parasitoids (Hymenoptera: Braconidae) Introduced for Fruit Fly Control on a Native Hawaiian Tephritid,Trupanea dubautiae(Diptera: Tephritidae)

Diachasmimorpha longicaudata(Ashmead) andPsyttalia fletcheri(Silvestri) are opiine parasitoids introduced into Hawaii for control of the Oriental fruit fly,Bactrocera dorsalis(Hendel) and the melon fly,Bactrocera cucurbitae(Coquillett), respectively. Both species have recently been mass-reared and released for research in augmentative biocontrol programs. Laboratory and field sleeve cage experiments were conducted to investigate the potential impact of mass-producedD. longicaudataandP. fletcherion a native Hawaiian tephritid,Trupanea dubautiae(Bryan), infesting the flowerheads of the native composite shrubDubautia raillardioidesHillebrand. Gravid females ofD. longicaudataandP. fletcheriwere confined with bloomingD. raillardioidesflowerheads infested with late instarT. dubautiae.BothD. longicaudataandP. fletcherilacked positive oviposition responses toT. dubautiaelarvae in infested flowerheads and caused neither parasitism nor mortality to the flies. However, when larvae were removed from the flowerheads and presented in screened dishes containing artificial diet of the parasitoids' normal rearing hosts (B. dorsalisandB. cucurbitae), bothD. longicaudataandP. fletcherireadily oviposited in the test larvae. Oviposition byD. longicaudatadid not significantly affect the percentage pupation ofT. dubautiae,but did reduce the emergence of adult flies. Oviposition byP. fletcherisignificantly reduced both pupation and adult fly emergence. All progeny of both parasitoid species died as eggs or first-instar larvae. Results from our experiments demonstrate that biological control programs targeted against frugivorous tephritid pests byD. longicaudataandP. fletcherihave no harmful impact on flowerhead-infestingT. dubautiae.     

Dynamics of Mycoplasmal Conjunctivitis in the Native and Introduced Range of the Host

In 1994, Mycoplasma gallisepticum, a common bacterial poultry pathogen, caused an epidemic in house finches in the eastern part of their North American range where the species had been introduced in the 1940s. Birds with mycoplasmal conjunctivitis were reported across the entire eastern United States within 3–4 years. Here we track the course of the Mycoplasma gallisepticum epidemic as it reached native, western North American populations of the house finch. In 2002, Mycoplasma gallisepticum was first observed in a native house finch population in Missoula, MT, where it gradually increased in prevalence during the next 2 years. Concurrently, house finches with conjunctivitis were reported with increasing number in the Pacific Northwest. In native populations of the host, the epidemic expanded more slowly, and reached lower levels of prevalence than in the eastern, introduced range of the species. Maximal prevalence was about half in the Missoula population than in local populations in the East. Although many factors can contribute to these differences, we argue that it is most likely the higher genetic heterogeneity in western than in eastern populations caused the lower impact of the pathogen.     

Phylogeography of the endangered tideland snail Batillaria zonalis in the Japanese and Ryukyu Islands

The phylogeography of the endangered tideland snail Batillaria zonalis in the Japanese and Ryukyu Islands was analyzed on the basis of nucleotide sequences of a mitochondrial gene for cytochrome oxidase c subunit I (COI). Extremely low genetic diversity was found in populations at both the northern and southern boundaries of the geographic distribution of this species in Japan, i.e., Sendai Bay and Iriomotejima Island, respectively, which might be attributed to the population bottleneck due to historical environmental variations and/or the recent foundation of populations in the marginal part of the inhabitable range. Most populations contained unique rare haplotypes, and significant genetic differentiation on the whole was shown, while no clear geographic genetic structure was detected between the Japanese and Ryukyu Islands or over the distribution area of B. zonalis in Japan, with the exception of significant genetic divergences in Ago Bay in the central part of Honshu and the southern part of Okinawajima Island.     

When Landscaping Goes Bad: The Incipient Invasion of Mahonia bealei in the Southeastern United States

Woodlots are forest islands embedded within an urban matrix, and often represent the only natural areas remaining in suburban areas. Woodlots represent critical conservation areas for native plants, and are important habitat for wildlife in urban areas. Invasion by non-indigenous (NIS) plants can alter ecological structure and function, and may be especially severe in remnant forests where NIS propagule pressure is high. Woody shrubs in the Family Berberidaceae have been well documented as invaders of the forest–urban matrix in North America. Mahonia bealei (Berberidaceae) is a clonal shrub native to China, and is a popular ornamental in the Southeastern United States. Mahoni bealei is listed as “present” on some local and state floras, but almost nothing is known regarding its invasion potential in the United States. We sampled 15 woodlots in Clemson, South Carolina, to assess the invasion of M. bealei and other woody non-indigenous species (NIS). M. bealei invaded 87% of the woodlots surveyed and species richness of NIS on these woodlots varied from 5 to 14. Stepwise-multiple regression indicated that less canopy cover and older M. bealei predicted greater abundance of M. bealei , and that not all subdivisions were equally invaded (P < 0.0001; r² = 0.88). The impact of M. bealei on native flora and fauna may be considerable, and it is likely to continue to spread in the Southeastern United States. M. bealei should be recognized as an aggressive invader in the Southeastern United States, with the potential for negative impacts on native flora and fauna.     

Trends in marine biological invasions at local and regional scales: the Northeast Pacific Ocean as a model system

Introduced species are an increasing agent of global change. Biogeographic comparisons of introduced biotas at regional and global scales can clarify trends in source regions, invasion pathways, sink regions, and survey effort. We identify the Northeast Pacific Ocean (NEP; northern California to British Columbia) as a model system for analyzing patterns of marine invasion success in cool temperate waters. We review literature and field surveys, documenting 123 introduced invertebrate, algal, fish, and vascular plant species in the NEP. Major invasion pathways were shipping (hull fouling, solid and water ballast; 1500s-present) and shellfish (particularly oysters) and finfish imports (commonest from the 1870s to mid-1900s). The cumulative number of successful invasions over time increased at linear, quadratic, and exponential rates for different taxa, pathways, and regions within the NEP. Regional analysis of four major NEP estuaries showed that Puget Sound and the contiguous Straits had the most introduced species, followed by Humboldt Bay, Coos Bay and Willapa Bay. Data on cumulative shipping volumes predicted smaller-scale, but not larger-scale spatial patterns in the number of shipping-mediated invasions. We identify the major challenges in scaling up from regional to global invasion analysis in cool temperate regions. Retrospective analyses for distinct biogeographic regions such as the NEP provide insight into vector dynamics and regional invasibility, and are a necessary foundation for monitoring and managing global change caused by biotic invasions.     

Bis(oxyphenylene)benzimidazoles: a novel class of anti-Plasmodium falciparum agents

A small library of 26 2,2′-[alkane-α,ω-diylbis(oxyphenylene)]bis-1H-benzimidazoles has been prepared and evaluated against Giardia intestinalis, Entamoeba histolytica, Trypanosoma brucei rhodesiense, Trypanosoma cruzi, Leishmania donovani, and Plasmodium falciparum. Among the tested compounds, eight derivatives (17, 19, 20, 24, 27, 30, 32 and 35) exhibited an anti-Plasmodium falciparum activity characterized by IC₅₀ values in the range of 180–410 nM (0.11–0.21 μg/mL) and selectivity indexes (IC₅₀ rat skeletal myoblasts L6 cells vs IC₅₀P. falciparum K1 strain) varying between 92 and more than 450. Two of the eight novel drug leads, namely compounds 19 and 32, were also active against G. intestinalis and L. donovani with selectivity indexes of 122 and >164 respectively.     

Modeling the Increase and Control of Caulerpa taxifolia, an Invasive Marine Macroalga

Population modeling based on species’ demography makes it possible to predict the pace of an invasion and evaluate the likelihood of success of different control strategies. We modeled the initial (density-independent) rate of increase of Caulerpa taxifolia (Vahl) C. Agardh (aquarium strain), a green alga that has markedly altered marine communities where it has invaded in the Mediterranean Sea. Parameter values for patch growth (from stolon extension) and reproduction (by asexual fragmentation and reattachment) were gleaned from published studies. Only the most conservative model, invoking field growth rates and low levels of fragment reattachment (2.5 m⁻² of existing patch each summer), closely matched observed increases (4–14 × annually). The most effective times for control (greatest reduction in rate of increase) were removal of established patches before summer and removal of fragments after summer. These times correspond to just before maximum growth and just after maximum reproduction, respectively. Only a combined strategy, incorporating 99% removal of all fragments and annual removal of 99% of established patches, was predicted to eliminate C. taxifolia entirely (λ < 1). This level of effort is only likely to be possible during the first few years of an invasion, arguing strongly for careful monitoring and rapid response to potential high-impact invaders.     

Competitive Growth Strategies in Intermediate Hosts: Experimental Tests of a Parasite life-History Model Using the Cestode, Schistocephalus solidus

In parasites with a complex life cycle, the fitness of an individual depends on its probability of reaching the final host and on its fecundity. Because larval growth in intermediate hosts may affect both transmission and adult size, selection should optimize growth patterns that are conditional on the presence and number of conspecific competitors. A recent model predicts that the total parasite volume per host should increase with intensity if larvae are able to vary growth depending on the number of conspecifics in the host (Life History Strategy hypothesis, i.e. LHS). Further, we would here expect growth rates to increase with intensity. By contrast, under the simplest alternative hypothesis of Resource Constraints (i.e. RC), the total parasite volume should remain constant. We experimentally infected copepods Macrocyclops albidus with the cestode Schistocephalus solidus to achieve 1, 2 or 3 parasites per host taking care that hosts had similar quality status at each infection level, and compared larval growth trajectories at the three intensity levels. The asymptotic total parasite volume was larger in double and triple infections than in single infections. Furthermore, the asymptotic total parasite volume was significantly larger in triple than in double infections but only in larger copepods that were less constrained by a host-size ceiling effect. These results, together with the fact that growth rates increased with intensity, support the LHS hypothesis: procercoids of a tapeworm may “count” their conspecific competitors in their first intermediate host to harvest its resources strategically until the next step in their complex life cycle. Co-ordinating editor: A. Biere     

Human-mediated Escalation of a Formerly Eradicable Problem: The Invasion of Caribbean Frogs in the Hawaiian Islands

Two species of neotropical frog, Eleutherodactylus coqui and E. planirostris, have been introduced into the State of Hawaii via the horticulture trade. E. coqui was introduced prior to 1988 and E. planirostris was first reported in 1994. Since these dates frog colonies have rapidly spread accidentally and intentionally and frog abundance within colonies has grown rapidly. Although these frogs were originally restricted to horticulture sites, they are now found in residential areas, resorts and hotels, and public lands. Due to the high potential biomass of introduced frogs there are realistic anthropogenic and ecological concerns associated with the spread of these frogs. Though there currently is a tool that can be used for localized control of frogs in limited circumstances, overall efforts by Federal, State, and County agencies to control the frog in Hawaii have been hampered by limited authorities and funds, disbelief in the threat, and the reluctance to act.     

Parasitism rates and sex ratios of a parasitoid wasp: effects of herbivore and plant quality

Summary We studied interactions among collards, Brassica oleracea var. acephala, the diamondback moth (DBM), Plutella xylostella (Lepidoptera: Yponomeutidae) and its parasitoid Diadegma insulare (Hymenoptera: Ichneumonidae) by manipulating plant nitrogen (N) concentrations in field and laboratory experiments. Parasitoid abundance strongly reflected DBM abundance and was related to total leaf N. Parasitism rates were high (70.7%) and density-independent. Wasp sex ratios varied markedly (3–93% female) in response to the herbivores, the plants, or both. Higher proportions of female wasps emerged from DBM larvae on plants with high leaf N than on unfertilized plants. More female wasps also emerged from larvae parasitized as larger instars. We suggest that wasps have the potential to control DBM populations through long-term numerical responses mediated by variable sex ratios.     

Dispersal Behavior, Boldness, and the Link to Invasiveness: A Comparison of Four Gambusia Species

Dispersal is a key element of a species' invasiveness. Although considerable work has addressed how dispersal influences the pattern of spatial spread of invading organisms, few studies investigate whether invasive species are in fact better dispersers than either the species they displace or less successful invaders. Recent work suggests that variation in dispersal may be due to variation in an underlying behavioral trait, boldness. Our study examined the link between dispersal, boldness, and invasiveness by comparing the dispersal characteristics and refuge use of two invasive Gambusia species to two congeners in experimental streams. The streams consisted of a series of pools (no flow) connected to a flowing channel. For each species, small groups of females were released at the middle pool, and their movement and activity were recorded over a 1-h period. We found invasive Gambusia to be more likely to disperse out of the introductory pool, to disperse sooner, to travel a greater distance in the artificial streams, and thus to exhibit greater dispersal tendencies than their close relatives. Among the invasives, Gambusia affinis had a greater dispersal tendency than G. holbrooki. We suspect this result indicates variation in the contribution of dispersal to the relative invasiveness of these species. Certain dispersal measures were correlated to time spent out of refuge, although invasive Gambusia and their relatives did not differ in the predicted manner. These results argue for the greater incorporation of experimental approaches and analyses of behavioral mechanisms in the study of invasive species.