Development of Notophthalmus viridescens embryos

Although the North American, adult, red-spotted newt has been an excellent model in the study of appendage regeneration at the tissue and cell levels, experiments involving embryonic forms have been essentially impossible to perform at the molecular level due to the great difficulty in breeding newts in the laboratory. Recently, we reported our methods to successfully spawn many thousands of embryos of Notophthalmus viridescens for developmental studies on a year round basis. As no detailed examination of embryogenesis of this amphibian exists, we provide a scanning electron microscopic overview of N. viridescens development. Furthermore, we chronicle the life cycle of the newt, when produced under laboratory conditions, which also undergoes two metamorphoses as newts in the wild.     

Predator-prey interactions between two amphibian species: effects of insecticide exposure

The presence of environmental contaminants may alter predator-prey interactions among aquatic species by altering activity levels of predators or prey, or by altering predator avoidance behavior. The outcome of a predatory encounter may be dependent upon whether both species are exposed to a contaminant simultaneously, or whether exposure occurs only in one of the species. In a laboratory experiment, I used the insecticide carbaryl to examine predation of southern leopard frog tadpoles (Rana sphenocephala) by adult red-spotted newts (Notophthalmus viridescens) under four conditions: both tadpoles and newts exposed, neither tadpoles nor newts exposed, and either newts or tadpoles only exposed. After one hour, exposed newts consumed half as many tadpoles as non-exposed newts. Carbaryl potentially affected newt activity enough to reduce time spent searching for prey, or may have altered the speed and coordination necessary to capture tadpoles. After six hours, non-exposed and exposed newts consumed similar numbers of tadpoles, most likely indicating recovery from exposure. After 24 h, predation rates were lowest when both newts and tadpoles were simultaneously either exposed or not exposed, and were greatest when newts and tadpoles were not exposed simultaneously. This study suggests that when tadpoles and newts are exposed to a sublethal level of a contaminant simultaneously, that predation rates do not differ from those observed under natural conditions, but exposure of either predator or prey at different times can disrupt predator-prey dynamics. This revised version was published online in August 2006 with corrections to the Cover Date.     

Differential Kin Discrimination by Red-spotted Newts (Notophthalmus viridescens) and Smooth Newts (Triturus vulgaris)

In organisms lacking parental care or when eggs of more than one female are layed together, parents preferentially tend to avoid eating their own offspring. In some species of newts (Caudata: Salamandridae), there is no parental care and cannibalism of eggs and larvae occurs when philopatric adults remain in the pond throughout the breeding season where eggs and larvae develop. Kin discrimination by both adult newts and larvae would be expected to occur during the breeding season as it would enable individuals to benefit from the nutrients obtained from eating eggs and larvae while they avoid eating close relatives. The effects of kinship on cannibalism of eggs and larvae were examined in two species of newts. In separate, but similar, paired behavioral trials adult female red-spotted newts, Notophthalmus viridescens, and smooth newts, Triturus vulgaris, were allowed to consume their own egg or an egg from another female. Female red-spotted newts, but not smooth newts, preferred to eat eggs of other females rather than their own, and overall smooth newts were ‘reluctant’ to eat conspecific eggs. To test for ontogenetic shifts in discrimination abilities, additional experiments were performed with adult female red-spotted newts paired with their own larvae and a larva from another female. In addition, the effects of kinship on the attraction or repulsion of larvae was also studied in red-spotted newts. Females showed no significant discrimination abilities during acts of cannibalism on larvae. Larvae spent similar amounts of time with siblings and non-siblings. These results suggest that the ability of adult newts to discriminate among eggs varies between species and that the elicitation of a kin-discrimination response may be context-dependent. For red-spotted newts, there was no evidence of discrimination abilities within and between other life-history stages.     

Anuran occupancy of created wetlands in the Central Appalachians

Evaluating the adequacy of created wetlands to replace the functions of lost natural wetlands is important because wetland mitigation is a major tool used to offset wetland losses. However, measurements such as vegetative cover and presence of wildlife may not provide sufficient evidence that created wetlands are functioning properly and thus examining the ecology of wetland biota such as that of amphibians may be a more useful surrogate for function. The objectives of this study were to compare the occupancy and detection of calling anurans in created wetlands relative to beaver-created wetlands. Five-, 10-min, and broadcast call surveys were performed at 24 wetlands throughout the Central Appalachians once every month from March through August of 2009 and 2010. Occupancy modeling was used to estimate the occupancy and detection of individual species, incorporating relevant environmental variables. The occupancy of anurans did not differ between human-created and beaver (Castor canadensis)-created wetlands. Detection of anurans was largely unaffected by call survey type, but several environmental covariates had a significant effect on the detectability of calling anurans. Our results suggest that the function of providing adequate chorusing habitat for adult anurans is being fulfilled by the created wetlands that we examined.     

Thermal influence on metabolic rates and a bioenergetic budget for Notophthalmus viridescens from southwestern Massachusetts

Notophthalmus viridescens has been reported to overwinter on land in southwestern Massachusetts, whereas these newts hibernate in water in southwestern Ohio. Aquatic and terrestrial metabolic rates of newts from Massachusetts were measured at different exercise speeds and acclimation temperatures in order to better understand their seasonal energetic budgets. Oxygen uptake at 25°C increased with increased swimming and walking speeds and reached a plateau at speeds of 60 and 90 cm/min, whereas at 5°C, oxygen consumption linearly increased with swimming speeds. Aerobic transport costs of the newts thus decreased with increased locomotor speeds at 25°C but remained unchanged when the newts were exercised in water at 5°C. Anaerobic metabolic rates of the newts on land were little affected by acclimation temperature but also increased linearly with walking speeds at both 5°C and 25°C. Anaerobiosis contributed most of the energy for the locomotion of the newts. These newts stored an average of 12 mg lipd/g body mass, which could apparently support their survival at 5°C for 46 days without food on land but only for 18 days in water. These calculations, based on measured metabolic rates and energy reserves, support field observations of red-spotted newts hibernating on land in Massachusetts.     

Keystone predators (eastern newts, Notophthalmus viridescens) reduce the impacts of an aquatic invasive species

Predation, competition, and their interaction are known to be important factors that influence the structure of ecological communities. In particular, in those cases where a competitive hierarchy exists among prey species, the presence of certain keystone predators can result in enhanced diversity in the prey community. However, little is known regarding the influence of keystone predator presence on invaded prey communities. Given the widespread occurrence of invasive species and substantial concern regarding their ecological impacts, studies on this topic are needed. In this study I used naturalistic replications of an experimental tadpole assemblage to assess the influence of predatory eastern newts, Notophthalmus viridescens, on the outcome of interspecific competition among native and nonindigenous tadpoles. When newts were absent, the presence of the tadpoles of one invasive species, the Cuban treefrog, Osteopilus septentrionalis, resulted in decreased survival and growth rate of the dominant native species, Bufo terrestris, and dominance of the tadpole assemblage by O. septentrionalis. However, the presence of one adult newt generally reduced or eliminated the negative impacts of O. septentrionalis tadpoles, resulting in comparable survival and performance of native species in invaded and noninvaded treatments. Differential mortality among the tadpole species suggests that newts preyed selectively on O. septentrionalis tadpoles, supporting the hypothesis that newts acted as keystone predators in the invaded assemblage. The presence of nonindigenous larval cane toads, Bufo marinus, did not significantly affect native species, and this species was not negatively affected by the presence of newts. Collectively, these results suggest that eastern newts significantly modified the competitive hierarchy of the invaded tadpole assemblage and reduced the impacts of a competitively superior invasive species. If general, these results suggest that the presence of certain species may be an essential factor regulating the ecological impacts of biological invasions.     

Stress and changes in the blood of newts,Notophthalmus viridescens,during early regeneration

Summary During the summers of 1984 and 1985, adult red-spotted newts,Notophthalmus viridescens, were maintained in the laboratory at 23°±0.5°C under natural photoperiods. From each of the experimental animals, the right forelimb was amputated just proximal to the elbow. Control newts were not manipulated surgically. Eight, 15, and 22 days after the time of amputation, equal numbers of regenerating and control animals were sacrificed, and blood smears of each individual were prepared with Wright's stain.Mean differential counts of leukocytes of the two groups of newts indicated that the relative number of neutrophils increased and the relative number of lymphocytes decreased in the regenerating animals as compared to their controls (Fig. 1 and Fig. 2). Earlier studies had shown that lymphopenia and neutrophilia occur in red-spotted newts treated with hydrocortisone or with ACTH or subjected to environmental stress (Bennett and Daigle 1983). Consequently, it is suggested that amputation and/or early regeneration may stimulate the increased production of hormones associated with stress in vertebrates, which may, in turn, influence regeneration, itself, and that the detailed study of the distribution of leukocytes inNotophthalmus viridescens may provide an assay with which to study the regulation of regeneration in this species.     

Magnetic compass orientation in the Eastern red-spotted newt (Notophthalmus viridescens)

Summary Laboratory tests were carried out to examine the orientation behavior of adult Eastern red-spotted newts (Notophthalmus viridescens) to earth-strength magnetic fields. Groups of 30 to 40 newts were housed in water-filled, all-glass aquaria with an artificial shoreline at one end. The aquaria were located in a greenhouse or outdoors adjacent to the laboratory building, and aligned on either the magnetic north-south or east-west axis. Tests were carried out in an enclosed indoor arena. Newts were tested in four horizontal alignments of the magnetic field: the ambient magnetic field (magnetic north at North) and three altered fields (magnetic north rotated to East, South or West). Data were analyzed after pooling the magnetic bearings from all four conditions in such a way as to retain the component of the newts' orientation that was a consistent response to the magnetic field. Elevation of training tank water temperature was used to increase the newts' motivation to orient in the direction of shore. Newts exposed to a training tank water temperature of 33–34 °C just prior to testing exhibited consistent unimodal magnetic compass orientation. The direction of orientation was altered predictably by changing training tank alignment and location relative to the laboratory building. The results provide the first evidence of a strong, replicable magnetic compass response in a terrestrial vertebrate under controlled laboratory conditions. Further, the present study demonstrates that the Eastern newt is able to learn a directional response relative to the earth's magnetic field.     

A comparison of created and natural wetlands in Pennsylvania,USA

Recent research suggests that created wetlands do not look, or function, like the natural systems they are intended to replace. Proper planning, construction, and the introduction of appropriate biotic material should initiate natural processes which continue indefinitely in a successful wetland creation project, with minimal human input. To determine if differences existed between created and natural wetlands, we compared soil matrix chroma, organic matter content, rock fragment content, bulk density, particle size distribution, vegetation species richness, total plant cover, and average wetland indicator status in created (n = 12) and natural (n = 14)wetlands in Pennsylvania (USA). Created wetlands ranged in age from two to 18 years. Soils in created wetlands had less organic matter content, greater bulk densities, higher matrix chroma, and more rock fragments than reference wetlands. Soils in reference wetlands had clay loam textures with high silt content, while sandy clay loam textures predominated in the created sites. Vegetation species richness and total cover were both greater in natural reference wetlands. Vegetation in created wetlands included a greater proportion of upland species than found in the reference wetlands. There were significant differences in soils and vegetation characteristics between younger and older created wetlands, though we could not say older created sites were trending towards the reference wetland condition. Updated site selection practices, more careful consideration of monitoring period lengths, and, especially, a stronger effort to recreate wetland types native to the region should result in increased similarity between created and natural wetlands.     

Macroinvertebrate community convergence between natural,rehabilitated, and created wetlands

Wetland restoration practices can include rehabilitating degraded wetlands or creating new wetlands. Empirical evidence is needed to determine if both rehabilitated and created wetlands can support the same macroinvertebrate communities as their natural counterparts. We measured long‐term macroinvertebrate community change in seasonal wetlands known as Delmarva Bays in Maryland, U.S.A. We compared a rehabilitated, a created, and a natural Delmarva Bay. We hypothesized that the created and rehabilitated wetlands would develop different macroinvertebrate communities. We also hypothesized that the community composition of the rehabilitated wetland would become more similar to that of the natural wetland than to that of the created wetland over 9 years encompassed by this study. We monitored the macroinvertebrates, including both predators and primary consumers, and environmental conditions in the three wetlands from March to August in 2005, 2006, 2007, and 2012. Cluster analysis indicated that from 2005 to 2007, the macroinvertebrate community of the rehabilitated wetland and the created wetland were more similar to each other than to the natural wetland. In 2012, the rehabilitated wetland was more similar to the natural wetland than to the created wetland. This similarity was driven principally by changes in the composition of primary consumer taxa. Our results suggest that rehabilitated Delmarva Bays are more likely to support a natural macroinvertebrate community than are created wetlands. Restoration practices that rehabilitate existing wetlands may be preferred over practices that create new wetlands when restoration project goals include developing natural macroinvertebrate communities in a short period of time.     

Replacement of a unique population of newts (<Emphasis Type="Italic">Taricha granulosa mazamae</Emphasis>) by introduced signal crayfish (<Emphasis Type="Italic">Pacifastacus leniusculus</Emphasis>) in Crater Lake,Oregon

The signal crayfish (Pacifastacus leniusculus) was introduced to Crater Lake in 1915 and now threatens the local extinction of an endemic salamander, the Mazama newt (Taricha granulosa mazamae). More than a century after their introduction, crayfish have expanded in distribution to occupy nearly 80% of the lakeshore. Although newts remain in uninvaded areas, they are almost entirely absent in crayfish occupied areas. Abundance of benthic macroinvertebrates was dramatically reduced in locations with crayfish compared with areas of the lake where crayfish were absent. Isotopic signatures of newt and crayfish tissue confirm overlap in the diets of the two species and demonstrate their similar position in the Crater Lake food web. Mesocosm experiments conducted with newts and crayfish revealed that crayfish prey directly on newts, displace newts from cover, and generally alter newt behavior. Combined, this evidence suggests that further crayfish expansion likely will cause additional declines in newt abundance and distribution, and could lead to extinction of the unique population of newts in Crater Lake.     

Repellent function of male pheromones in the red-spotted newt

Pheromones act as attractants and sexual stimulants in most vertebrates. For example, in red-spotted newts, Notophthalmus viridescens, female pheromones attract males, and male pheromones increase female receptivity. However, no studies have determined whether male vertebrates produce a pheromone that repels competing males. Through a series of olfactory mate selection tests, we found that sexually motivated male red-spotted newts produce a pheromone that functions to repel other approaching males. Our finding is the first report of a repelling function for pheromones in male vertebrates. The pheromones may act to increase both the sender's and receiver's mating success when the operational sex ratio (OSR) is male biased.     

Litter decomposition in created and reference wetlands in West Virginia,USA

Large amounts of resources have gone into wetland mitigation in recent years; however, it is still unclear whether wetland function is being replaced along with wetland area. Litter decomposition is linked to numerous wetland functions. In this study, we measured plant litter decomposition potential over 12 months in 8 created and 8 reference wetlands located in the Allegheny Mountains of West Virginia, USA. Broadleaf cattail (Typha latifolia L.) litter bags were placed in each wetland and collected at 3 month intervals. Linear decomposition rate constants and percent mass remaining were similar between wetland types (created and reference) and among Cowardin classifications (palustrine: unconsolidated bottom, aquatic bed, emergent, and scrub/shrub). Created wetland age was not correlated with decomposition potential. Our study found that created wetlands had similar litter decomposition potential as reference wetlands indicating that similar processes are likely acting upon litter decomposition within both natural and created wetlands.     

Genetic structure of natural populations of the red-spotted newt,Notophthalmus viridescens

Genetic variation is described at 15 loci in 2 neotenic and 12 nonneotenic populations of red-spotted newts. Though high levels of genetic similarity (I=0.990) were found among all populations, allele frequencies at six of the eight most polymorphic loci show significant heterogeneity across populations. Change in allele frequencies at two of these loci (Pep-2 and Ldh-1) is significantly correlated with latitude. Interspecific homologies are established for newt peptidases based on substrate specificities and lactate dehydrogenases based on tissue distribution, thermal stability, and kinetic properties. Nonneotenic populations are highly variable (H=0.157) and neotenic populations are only slightly, but significantly, less variable (H=0.120). The high levels of heterozygosity detected in nonneotenic populations may result from large effective population size and/or environmental heterogeneity. The unexpectedly high heterozygosity values obtained for the neotenic populations may indicate adult dispersal or the presence of some previously undetected red efts at these localities. In any case, a major change in life history has apparently had little effect on the genetic structure of these populations.This research was supported by grants from the Blakeslee Fund of Smith College.     

Conservation genetics of the central newt (Notophthalmus viridescens) in Iowa: the importance of a biogeographic framework

In light of global declines in amphibian populations, genetic data have become increasingly important for understanding population structure and for revealing hidden diversity. At the species level, Notophthalmus viridescens is an IUCN species of “least concern”, but the subspecies N. v. louisianensis (central newt) is listed as “threatened” in Iowa, a state on the western periphery of the species range. Genetic data were collected from 282 N. v. louisianensis from 14 sites in Iowa. Sequences from 1,054 nucleotides of mitochondrial DNA from Iowa newts revealed unexpected diversity in the form of two major haplotype groups that are not sister clades, with southern Iowa N. v. louisianensis being more closely related to N. v. piaropicola (peninsula newt) from Florida than to consubspecifics in Iowa. Sequence differentiation indicates that the two lineages of newts present in Iowa diverged near the beginning of the Pleistocene. Northern and southern Iowa haplotypes were found together at one site, indicating an opportunity for hybridization near Remington’s biogeographic suture zone 1, a hotspot for hybridization in other species. Three microsatellite loci provided additional evidence for distinctness of northern and southern Iowa newt populations. This study highlights the relevance of historical biogeography to conservation, as management strategies for N. v. louisianensis in Iowa must reflect previously unrecognized diversity in this species. Nuclear and mitochondrial data indicate genetic isolation of nearby populations on the same drainage, and field data suggest the decline of one study population, emphasizing the need for identification and protection of newt breeding sites in Iowa.     

Chemical defense of the eastern newt (Notophthalmus viridescens): variation in efficiency against different consumers and in different habitats

Amphibian secondary metabolites are well known chemically, but their ecological functions are poorly understood--even for well-studied species. For example, the eastern newt (Notophthalmus viridescens) is a well known secretor of tetrodotoxin (TTX), with this compound hypothesized to facilitate this salamander's coexistence with a variety of aquatic consumers across the eastern United States. However, this assumption of chemical defense is primarily based on observational data with low replication against only a few predator types. Therefore, we tested the hypothesis that N. viridescens is chemically defended against co-occurring fishes, invertebrates, and amphibian generalist predators and that this defense confers high survivorship when newts are transplanted into both fish-containing and fishless habitats. We found that adult eastern newts were unpalatable to predatory fishes (Micropterus salmoides, Lepomis macrochirus) and a crayfish (Procambarus clarkii), but were readily consumed by bullfrogs (Lithobates catesbeianus). The eggs and neonate larvae were also unpalatable to fish (L. macrochirus). Bioassay-guided fractionation confirmed that deterrence is chemical and that ecologically relevant concentrations of TTX would deter feeding. Despite predatory fishes rejecting eastern newts in laboratory assays, field experiments demonstrated that tethered newts suffered high rates of predation in fish-containing ponds. We suggest that this may be due to predation by amphibians (frogs) and reptiles (turtles) that co-occur with fishes rather than from fishes directly. Fishes suppress invertebrate consumers that prey on bullfrog larvae, leading to higher bullfrog densities in fish containing ponds and thus considerable consumption of newts due to bullfrog tolerance of newt chemical defenses. Amphibian chemical defenses, and consumer responses to them, may be more complex and indirect than previously appreciated.     

Ecosystem Development After Mangrove Wetland Creation: Plant–Soil Change Across a 20-Year Chronosequence

Mangrove wetland restoration and creation efforts are increasingly proposed as mechanisms to compensate for mangrove wetland losses. However, ecosystem development and functional equivalence in restored and created mangrove wetlands are poorly understood. We compared a 20-year chronosequence of created tidal wetland sites in Tampa Bay, Florida (USA) to natural reference mangrove wetlands. Across the chronosequence, our sites represent the succession from salt marsh to mangrove forest communities. Our results identify important soil and plant structural differences between the created and natural reference wetland sites; however, they also depict a positive developmental trajectory for the created wetland sites that reflects tightly coupled plant-soil development. Because upland soils and/or dredge spoils were used to create the new mangrove habitats, the soils at younger created sites and at lower depths (10–30?cm) had higher bulk densities, higher sand content, lower soil organic matter (SOM), lower total carbon (TC), and lower total nitrogen (TN) than did natural reference wetland soils. However, in the upper soil layer (0–10?cm), SOM, TC, and TN increased with created wetland site age simultaneously with mangrove forest growth. The rate of created wetland soil C accumulation was comparable to literature values for natural mangrove wetlands. Notably, the time to equivalence for the upper soil layer of created mangrove wetlands appears to be faster than for many other wetland ecosystem types. Collectively, our findings characterize the rate and trajectory of above- and below-ground changes associated with ecosystem development in created mangrove wetlands; this is valuable information for environmental managers planning to sustain existing mangrove wetlands or mitigate for mangrove wetland losses.     

Interactions of bullfrog tadpole predators and an insecticide: predation release and facilitation

The effect of a contaminant on a community may not be easily predicted, given that complex changes in food resources and predator-prey dynamics may result. The objectives of our study were to determine the interactive effects of the insecticide carbaryl and predators on body size, development, survival, and activity of tadpoles of the bullfrog (Rana catesbeiana). We conducted the study in cattle tank mesocosm ponds exposed to 0, 3.5, or 7.0 mg/l carbaryl, and no predators or two red-spotted newts (Notophthalmus viridescens), bluegill sunfish (Lepomis macrochirus), or crayfish (Orconectes sp.). Carbaryl negatively affected predator survival by eliminating crayfish from all ponds, and by eliminating bluegill sunfish from ponds exposed to the highest concentration of carbaryl; carbaryl exposure did not effect survival of red-spotted newts. Because crayfish were eliminated by carbaryl, bullfrogs were released from predation and survival was near that of predator controls at low concentrations of carbaryl exposure. High concentrations of carbaryl reduced tadpole survival regardless of whether predators survived carbaryl exposure or not. Presence of crayfish and newts reduced tadpole survival, while bluegill sunfish appeared to facilitate bullfrog tadpole survival. Presence of carbaryl stimulated bullfrog tadpole mass and development. Our study demonstrates that the presence of a contaminant stress can alter community regulation by releasing prey from predators that are vulnerable to contaminants in some exposure scenarios.Due to an error in the citation line, this revised PDF (published in December 2003) deviates from the printed version, and is the correct and authoritative version of the paper.     

Wetland type differentially affects ectoparasitic mites and their damselfly hosts

The effect of artificial habitat in altered landscapes on species interactions and their suite of enemies is largely unknown. Water mites have been associated with reduced fitness of model damselflies. Mite parasitism was variable, but higher for Ischnura verticalis damselflies from natural, than from artificial, wetlands in the same region. There were no differences in timing of sampling, temperature during sampling, or host age or sex composition of samples between wetland types. Landscape structure might constrain mite presence or abundance at wetland sites or wetland type might be a better predictor of mites, based on factors such as prey abundance. Fewer mites on damselflies from numerous artificial wetlands means that the strength of parasite-mediated selection is likely less than would be inferred if only natural wetlands were surveyed. Such effects of human changes in habitats on host species probably occur often.     

Parasite age-intensity relationships in red-spotted newts: does immune memory influence salamander disease dynamics?

Acquired immune memory in vertebrates influences transmission and persistence of infections, with consequences for parasite dynamics at both the individual and population levels. The potential impact of acquired immunity is of particular interest for salamanders, whose acquired immune systems are thought to be less effective than those of frogs and other tetrapods. One way to examine the importance of acquired immunity to parasite dynamics at the population level is by examining the relationship between host age and parasite infection intensity. Acquired immunity reduces infection rates in older animals, causing decreased parasite intensity in older age classes and leading to curvilinear age-intensity relationships for persistent parasites and convex age-intensity relationships for transient parasites. We used age-intensity relationships to look for the signature of acquired immunity for 12 parasite taxa of red-spotted newts (Notophthalmus viridescens), using data from a 2-year parasitological survey of six newt populations. We estimated ages from snout-vent length (SVL) based on the relationship between SVL and skeletochronologically-derived ages in a subset of newts. We found evidence of acquired immunity to two parasite taxa, bacterial pathogens and the protist Amphibiocystidium viridescens, whose convex age-intensity relationships could not be easily explained by alternative mechanisms. Our results suggest that the acquired immune response of newts is sufficient to influence the dynamics of at least some parasites.