Is Hybridization Responsible for Invasive Growth of Non-indigenous Water-milfoils?

Heterosis, or hybrid vigor, has recently been proposed as a factor promoting invasive growth of some non-indigenous aquatic plant species, particularly those capable of spreading rapidly within and among lakes through clonal reproduction. We tested this hypothesis for variable-leaf water milfoil (Myriophyllum heterophyllum), a non-indigenous aquatic plant that has become a major management and conservation concern in New England. Using nuclear ribosomal DNA, we looked for F₁ hybrid populations of invasive M. heterophyllum in 25 New Hampshire (NH) lakes. In contrast to a previous study that found F₁ hybrid lineages of invasive M. heterophyllum in Connecticut, we did not find hybrids in our study lakes. This result has two implications: (1) pure lineages of M. heterophyllum are also capable of invasive growth, and (2) the distribution of invasive M. heterophyllum lineages (hybrid vs. pure) may be spatially structured across New England. We stress the importance of more detailed distributional and ecological studies for understanding the invasive potential of this species.     

Comparative studies on scanning electron microscopy of trophi of the genus Filinia Bory De St. Vincent (Rotifera)

Trophi of Filinia species from 16 South Island lakes and three North Island lakes of New Zealand were examined and compared with specimens from Australia, Austria, Belgium, and Turkey. Five species of Filinia (brachiata, longiseta, cf. pejleri, novaezealandiae, and terminalis) were positively identified from the New Zealand samples. Numbers of unci teeth are considered to be the most reliable features for identification within the genus. Numbers obtained from SEM for other species of Filinia (australiensis, grandis, and hofmanni) are also listed for the first time.     

RELATIONSHIPS OF SEDIMENTED DIATOM SPECIES (BACILLARIOPHYCEAE) TO ENVIRONMENTAL GRADIENTS IN DILUTE NORTHERN NEW ENGLAND LAKES1

Limnological gradients of small, oligotrophic, and low conductance lakes in northern New England were defined by principal components analysis; relationships of sedimented diatom species to the gradients were investigated by correlation analysis. Diatom distributions were most strongly related to the gradient of pH and alkalinity and the covarying variables, conductance, Mg, Ca, total Al, and exchangeable Al. Weaker relationships to lake morphology, dissolved organic carbon and water color, altitude and marine aerosol inputs, and the distinctive water chemistry of some New Hampshire lakes were also present. Results for 16 taxa of importance in our studies of lake acidity are given in detail and are compared to results from other regions of eastern North America. Planktonic taxa were absent below pH 5.5, with the exception of the long form of Asterionella ralfsii var. americana Korn. The two forms of this taxon differed ecologically: the long form (>45μm) had an abundance weighted mean (AWM) pH 4.90 and occurred mostly in lakes that were deep relative to transparency; the short form (<45μm)had an AWM pH and occurred on lakes that were shallow relative to transparency. The ecological advantage of a “splitter” approach to diatom taxonomy was demonstrated by examination of other taxa as well, including Tabellaria flocculosa (Roth) Kütz. These results have important implications for paleolimnological interpretations.     

Predictions of establishment risk highlight biosurveillance priorities for invasive fish in New Zealand lakes

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REPRODUCTION AND SYSTEMATICS OF THE MARINE ALGA DERBESIA (CHLOROPHYCEAE) IN NEW ENGLAND1,2

The morphology, gross cytology, reproduction, and habitat ecology are described for Derbesia marina based on observations of New England field populations and laboratory cultures of this plant. These data, and additional observations on cultures of several other species of Derbesia from elsewhere and on type and other important collections, have been used to evaluate the systematic relationship of New England Derbesia with other species of this genus. The single Derbesia species in New England is referred to D. marina. The systematic criteria previously used to distinguish species within Derbesia are reviewed and interpreted within the context of the present investigation. In systematic studies, we stress the importance of the use of sporangial and chloroplast morphology, the presence or absence of pyrenoids, and reproductive history. Two types of life history are reported for D. marina in New England: (1) A form of D. marina collected at 20 m reproduces directly with stephanokontous zoospores growing into sporophytic plants of Derbesia morphologically identical to their parent thalli. (2) On the other hand, at least some New England, populations of D. marina have retained the genetic potential for producing a sexual generation (Halicystis ovalis), even though the latter is unknown for the coast of northeastern North America.³ Gametophytes (H. ovalis) were produced directly from enlargement and subsequent differentiation of uncleaved lateral sporangia in 2 cultured populations of New England D. marina. A single female and numerous male vesicles formed in this manner produced gametes, but neither fertilization nor parthenogenesis occurred; thus the entire life history was not completed in culture. The occurrence of the directly reproducing deep water form of D. marina is presented as evidence for speciation of a sporophyte (Derbesia) independent of its alternate gametophyte (Halicystis). An hypothesis is advanced to explain the source and means for expression of genetic variability necessary for speciation in a population of nonsexually reproducing Derbesia.     

Relationships between zooplankton abundance and trophic state in seven New Zealand lakes

The zooplankton communities of seven Rotorua, New Zealand, lakes of different trophic status were studied in 1977–78. They were generally dominated by the calanoid copepod, Calamoecia lucasi. Bosmina meridionalis occurred in all the lakes and Ceriodaphnia dubia in most. Only small numbers of Macrocyclops albidus ever occurred. Rotifers were not studied in detail. Community composition was similar to that in other northern New Zealand lakes. No well defined patterns of seasonal change in abundance were found and the timing of changes, which were of low magnitude, was different in each lake. Clutch sizes in all species were small. Calamoecia population parameters were analysed using multivariate methods and shown to be related to lake trophic level. Population densities were higher in more productive lakes whereas breeding levels were inversely related to indices of trophic status and population abundance. It is suggested that the populations, as in other northern New Zealand lakes, are food-limited, probably as a consequence of a lack of marked climatic seasonality and the absence of major predation pressures. Groupings of the lakes based on the Calamoecia data are in general agreement with those derived from parallel studies of water chemistry, phytoplankton and macrobenthos.     

Post-glacial changes in spatial patterns of vegetation across southern New England

Aim We analysed lake‐sediment pollen records from eight sites in southern New England to address: (1) regional variation in ecological responses to post‐glacial climatic changes, (2) landscape‐scale vegetational heterogeneity at different times in the past, and (3) environmental and ecological controls on spatial patterns of vegetation. Location The eight study sites are located in southern New England in the states of Massachusetts and Connecticut. The sites span a climatic and vegetational gradient from the lowland areas of eastern Massachusetts and Connecticut to the uplands of north‐central and western Massachusetts. Tsuga canadensis and Fagus grandifolia are abundant in the upland area, while Quercus, Carya and Pinus species have higher abundances in the lowlands. Methods We collected sediment cores from three lakes in eastern and north‐central Massachusetts (Berry East, Blood and Little Royalston Ponds). Pollen records from those sites were compared with previously published pollen data from five other sites. Multivariate data analysis (non‐metric multi‐dimensional scaling) was used to compare the pollen spectra of these sites through time. Results Our analyses revealed a sequence of vegetational responses to climate changes occurring across southern New England during the past 14,000 calibrated radiocarbon years before present (cal yr bp ). Pollen assemblages at all sites were dominated by Picea and Pinus banksiana between 14,000 and 11,500 cal yr bp ; by Pinus strobus from 11,500 to 10,500 cal yr bp ; and by P. strobus and Tsuga between 10,500 and 9500 cal yr bp . At 9500–8000 cal yr bp , however, vegetation composition began to differentiate between lowland and upland sites. Lowland sites had higher percentages of Quercus pollen, whereas Tsuga abundance was higher at the upland sites. This spatial heterogeneity strengthened between 8000 and 5500 cal yr bp , when Fagus became abundant in the uplands and Quercus pollen percentages increased further in the lowland records. The differentiation of upland and lowland vegetation zones remained strong during the mid‐Holocene Tsuga decline (5500–3500 cal yr bp ), but the pattern weakened during the late‐Holocene (3500–300 cal yr bp ) and European‐settlement intervals. Within‐group similarity declined in response to the uneven late‐Holocene expansion of Castanea, while between‐group similarity increased due to homogenization of the regional vegetation by forest clearance and ongoing disturbances. Main conclusions The regional gradient of vegetation composition across southern New England was first established between 9500 and 8000 cal yr bp . The spatial heterogeneity of the vegetation may have arisen at that time in response to the development or strengthening of the regional climatic gradient. Alternatively, the differentiation of upland and lowland vegetation types may have occurred as the climate ameliorated and an increasing number of species arrived in the region, arranging themselves in progressively more complex vegetation patterns across relatively stationary environmental gradients. The emergence of a regional vegetational gradient in southern New England may be a manifestation of the increasing number of species and more finely divided resource gradient.     

The biogeography of three Boeckella species (Copepoda: Calanoida) in New Zealand

The New Zealand distributions of three species of Boeckella (Copepoda, Calanoida), B. triarticulata, B. dilatata and B. hamata are mapped. B. triarticulata is primarily a pond dweller but is also found in reservoirs and shallow lakes. B. dilatata is mainly found in the deeper glacial lakes and ponds in the central region of the South Island and B. hamata has a more widespread distribution in lakes and ponds in the South Island and lower half of the North Island. Differences in temperature optima, food requirements and dispersal ability among the three Boeckella species are related to vicariant events to explain their distribution in New Zealand.     

Evolutionary history of Daphnia drives divergence in grazing selectivity and alters temporal community dynamics of producers

Consumers with different seasonal life histories encounter different communities of producers during specific seasonal phases. If consumers evolve to prefer the producers that they encounter, then consumers may reciprocally influence the temporal composition of producer communities. Here, we study the keystone consumer Daphnia ambigua, whose seasonal life history has diverged due to intraspecific predator divergence across lakes of New England. We ask whether grazing preferences of Daphnia have diverged also and test whether any grazing differences influence temporal composition patterns of producers. We reared clonal populations of Daphnia from natural populations representing the two diverged life history types for multiple generations. We conducted short‐term (24 hr) and long‐term (27 days) grazing experiments in equal polycultures consisting of three diatom and two green algae species, treated with no consumer, Daphnia from lakes with anadromous alewife, or from lakes with landlocked alewife. After 24 hr, life history and grazing preference divergence in Daphnia ambigua drove significant differences in producer composition. However, those differences disappeared at the end of the 27‐day experiment. Our results illustrate that, despite potentially more complex long‐term dynamics, a multitrophic cascade of evolutionary divergence from a predator can influence temporal community dynamics at the producer level.     

Bdelloid Rotifers from Lakes above 1700 m in Western Italian Alps,with Taxonomic Notes on Dissotrocha macrostyla

Benthic and periphytic bdelloid communities from 16 alpine lakes from 1700 to 2850 m above sea level in Sesia Valley (Piedmont region, North‐western Italy), sampled during summer 2001 and 2002, were analyzed. Seventeen species were identified from these species‐poor communities, with 1 to 6 species each. Dissotrocha macrostyla and Philodina citrina were the most common species, present in 10 lakes while 9 species were collected from one lake only. New morphological details from S.E.M. pictures of Dissotrocha macrostyla revealed that Dissotrocha macrostyla tuberculata (Gosse , 1886) is only a seasonal morphotype. Its different appearance is due to the presence of locally distributed microscopic mucous bubbles (diameter 1.41 ± 0.18 μm) on the trunk surface, produced by the rotifer itself under stressful conditions.     

Benthic foraminifera associated with the invasive ascidian, Didemnum sp. A

The invasive ascidian, Didemnum sp. A, first appeared in New England bays and harbors in the early 1990s, and in the waters around Cape Cod in 1993. While ship traffic was the likely vector introducing the species, its origin and precise date and location of its introduction are presently unknown. Colony surfaces of Didemnum sp. A appear very clean and not favorable substrates for epibiota settlement, but closer inspection revealed the presence of benthic foraminifera. During 2003 and 2004, 52 samples of Didemnum sp. A and other ascidians were collected to determine whether or not the foraminiferal assemblages might also be non-native and thus provide a potential clue to the place of origin of Didemnum sp. A. Sample locations included the New England coast from Connecticut to Maine (with a concentration in the Cape Cod area), northern California, Zeeland, The Netherlands, and Shakespeare Bay, New Zealand. From New England samples, 18 species of benthic foraminifera were identified. The most common species represented were Cornuspira involvens, C. planorbis, Elphidium galvestonense, E. margaritaceum, Glabratellina lauriei, Miliolinella subrotunda, Quinqueloculina bicornis, and Rosalina floridana. Foraminiferal assemblages on Didemnum sp. A from other regions sampled were composed of the same cosmopolitan species found in New England, plus other species which were indigenous to each region. Because no exotic foraminifera species were found it is concluded that Didemnum sp. A likely did not introduce non-native foraminifera originating from their native habitats into the New England region.     

Predicting spread of invasive macrophytes in New Zealand lakes using indirect measures of human accessibility

1. Models predicting invasive macrophyte spread between lakes provide an important tool for focusing proactive management efforts to lakes deemed susceptible to invasion. However, challenges to forecasting macrophyte spread include wide physiological tolerances of invasive macrophytes and a lack of information on the relative importance of the various human vectors (e.g. boating traffic). In New Zealand, three invasive species that reproduce vegetatively, Ceratophyllum demersum, Lagarosiphon major, Egeria densa, and a single species that reproduces sexually, Utricularia gibba, are currently spreading across the lake landscape at a great cost to the local ecology and economy. 2. In this study, we first examined whether variables that indirectly describe weed spread via human access and use, as well as a lake’s position in the landscape, could describe the distribution of these four species using a boosted regression trees (BRT) modelling approach. Then, as these invasive species have not reached their full invasion potential, we examined how giving more influence to infected lakes at the edge of the invasion front, and including all lakes across New Zealand as background samples, simulating ‘absences beyond the invasion front’, influenced our ability to forecast the potential for new lakes to be invaded. 3. The BRT models identified that variables characterising human access and use, as well as lake position, were associated with the occurrence of the three vegetatively reproducing macrophytes. Weed occurrence was more likely when there was a highway in the vicinity, human population density was high and if the lake was large (c. 55 km²). But in the single case of U. gibba, temperature was the variable that best explained occurrence. This is consistent with the suggestion that U. gibba is predominantly dispersed by waterbirds, rather than human activity. 4. But for all four species, the BRT models based on the recorded observations alone predicted observed invasions with low prediction probabilities and did not forecast further spread. By contrast, when observations at the edge of the invasion front were upweighted, and additional background lakes implemented into the model, recorded observations were predicted and additional lakes were forecast to be at risk, suggesting that these models better captured the current and potential distribution of these macrophyte species. 5. The use of variables that characterise weed spread could provide similar insights into other systems where survey information on the nature, strength and direction of invasion vectors is lacking. Furthermore, when weighting the data, many lakes across New Zealand were forecasted to be at risk of invasion. The advantage of weighing the presence data was that insights into the potential for a species to spread were obtained. The probabilistic estimates of risk, as derived from the models, together with other information for prioritising lakes, can be used to focus surveillance and protection efforts.     

Errors in pH measurement with colorimetric indicators in low alkalinity waters

A series of laboratory and field studies were conducted to evaluate the reliability of colorimetric pH indicators in dilute, acidic surface waters in New England. Laboratory studies consisted of comparing colorimetric and electrometric pH results in synthetic solutions with a range of ionic strength and buffering capacity. Field studies involved simultaneous colorimetric and electrometric pH determinations in 52 New England lakes with differing water chemistry. Colorimetric indicators deviated from electrometric pH results at low alkalinities and near the end of their operating range. They agreed with electrometric values, within 0.2 unit, at alkalinities of 20 µeq 1^-1 and greater, or near the center of their operating range. Changing ionic strength without changing buffering capacity did not affect results. When indicators were selected that bracketed sample pH, colorimetric determinations of pH in New England lakes agreed well with electrometric determinations.     

Habitat management to reduce competitive interactions: case study of native and invading cottontails

Habitat management recommendations are often based on best available science determined through retroductive and inductive hypotheses. Such recommendations are not frequently tested, potentially resulting in the implementation of unreliable practices for management of imperiled species. The New England cottontail (Sylvilagus transitionalis) is an imperiled shrubland-obligate species whose recovery efforts include habitat management and restoration. Researchers suggest former best management practices for the species may result in ecological traps and new recommendations have been developed. We evaluated these newly revised best management practices designed to retain higher tree canopy closure to promote New England cottontails without encouraging eastern cottontails (Sylvilagus floridanus). We compared New England and eastern cottontail density between management plots (tree canopy thinned with all downed trees left on the ground, with or without invasive shrub treatment) and control plots (unmanaged) and examined the influence of management on resource selection and survival. Management strategies retaining higher canopy closure promoted stronger selection by New England cottontails than by eastern cottontails. Catch per unit effort of New England cottontails was greater than for eastern cottontails in management plots (P = 0.002). For both species, the proportion of the 95% home range overlapping managed areas was greater than the proportion of managed area in the habitat patch; however, for the 50% core area of the home range, this was only true for New England cottontails. When post-treatment canopy cover was >75%, New England cottontails selected canopy-thinning treatments without invasive shrub removal over unmanaged areas, but selection by eastern cottontails was unaffected by management treatment or canopy cover. Survival probability of both species was high and uncorrelated with time spent in management areas. Survival probability decreased as the average distance a rabbit moved in a 7-day period increased. Our results illustrate the need to revise management strategies that emphasize eliminating canopy cover when improving New England cottontail habitat is an objective, particularly where they are sympatric with eastern cottontails.     

The submersed macrophyte communities of adirondack lakes (New York,U.S.A.) of varying degrees of acidity

The submersed macrophyte communities of nine high-altitude oligotrophic lakes in the Adirondack Mountain region of upstate New York were characterized in detail in terms of percent cover of each species. The pH of these clear-water lakes ranged from 6.9 to 4.4 and the acid neutralizing capacity ranged from 91 μeq 1^?1 to ?38 μeq l^?1. The degree of acidity covaried with lower total ion content and nutrient status. The dominant species in the circumneutral lakes and in the acidic lakes were the same, with several Utricularia species being the major component of the bottom cover. Species richness and diversity were lower in the acidic lakes (pH < 5.5). The plants found in the acidic lakes were typical of oligotrophic systems in general, regardless of pH, and similar to those found in lake surveys in Scandinavia and Eastern Canada. Potamogeton confervoides Reichenb., Scirpus subterminalis Torr., Sphagnum spp. and Utricularia geminiscapa Benj. were the only species that were primarily exclusive to the acidic lakes surveyed. In contrast to the results of Scandinavian studies, Sphagnum was found to be a major component of the plant cover in only two of the five acidic lakes.     

MORPHOMETRY OF BATRACHOSPERMUM POPULATIONS INTERMEDIATE BETWEEN B. BORYANUM AND B. ECTOCARPUM (RHODOPHYTA)1

Batrachospermum plants collected as part of a comprehensive study of New England streams were found to be intermediate between B. boryanum Sirdt. find B. ectocarpum Sirdt. Sixty-four specimens from ten Rhode Island streams were examined for several characteristics in replicates of two to four hundred. The size ranges obtained were then compared with previous published data for B. boryanum and B. ectocarpum from Europe, Asia and North America. The Rhode Inland populations and individuals exhibited as much or even more variability as that reported for both species combined throughout the world. Therefore, it appears that there is no definite basis for separation of these taxa and it is proposed that they be merged under the name B. boryanum. This combined species ran be distinguished from other species of the section Batrachospermum by compressed mature lateral whorls, little secondary fascicle formation, curved distal branching, terminal hairs on a small proportion of branches, predominantly dioecious state, sessile trichogynes and carposporophytes in outer portions of lateral whorls.     

Trends in abundance of rocky intertidal seaweeds and filter feeders across gradients of elevation,wave exposure,and ice scour in eastern Canada

On NW Atlantic rocky shores, the main basal organisms in intertidal communities are seaweeds (Ascophyllum nodosum, Fucus spp. and Chondrus crispus) and filter feeders (barnacles, Semibalanus balanoides, and mussels, Mytilus spp.). Their ecology has been extensively studied in New England (United States), but knowledge gaps exist for northern shores, which are subjected to stronger environmental stress. Therefore, we studied the above organisms on Canadian shores. We quantified the summer abundance of these seaweeds and filter feeders across full vertical (intertidal elevation) and horizontal (wave exposure and winter ice scour) environmental gradients on the Gulf of St. Lawrence and open Atlantic coasts of Nova Scotia. At the regional scale along the open Atlantic coast, seaweeds showed similar abundances in Nova Scotia than values reported for New England. However, both filter feeders were considerably less abundant in Nova Scotia. At the local scale in Nova Scotia, intense winter ice scour (which only occurs on the Gulf of St. Lawrence coast) was associated with a very low abundance of all species except barnacles. Spatial trends in Nova Scotia were similar to patterns known for certain species elsewhere, such as A. nodosum being almost restricted to sheltered habitats, regardless of elevation, and C. crispus being almost restricted to low elevations, regardless of exposure. Other trends were, however, characteristic of Nova Scotia, such as C. crispus being frequent at low elevations in exposed habitats, unlike in New England, where mussels often predominate there because of competitive advantages. In Nova Scotia, mussels were always restricted to cracks and crevices, unlike in New England, where they form extensive intertidal beds on exposed shores. The direct effects of increased environmental stress and indirect effects through altered interspecific interactions might explain the regional differences in local species distribution, which will require experimental validation. Handling editor: K. Martens     

Predicting invasive success of Hydrilla verticillata (L.f.) Royle in flowing water

Alien aquatic plant species have had spectacular success in invading New Zealand’s freshwaters. Their continued introduction has resulted in few water bodies retaining their natural or original indigenous aquatic vegetation, and associated adverse changes, such as reduced biodiversity and amenity values are well known. This highlights the need to accurately assess aquatic plant species that are already in the country and those outside for their potential to have a negative impact on New Zealand’s lakes and waterways. The study presented in this paper describes an approach to evaluating the invasive weed potential of Hydrilla verticillata (L.f.) Royle compared with Elodea canadensis Michx. under flowing water conditions. The two species were planted in combination and alone, and in a sheltered or exposed position relative to the direction of the water flow. Over the 2-year study, two different waterflow rates were used (0.1 and 0.2 m/s) which represented the gauged flows from a stream of interest that contains E. canadensis and flows from a H. verticillata infested lake. H. verticillata was competitive compared with E. canadensis and able to persist and thrive in the presence of E. canadensis in both exposed and sheltered habitats. These results corroborate observations from other countries of H. verticillata growing in flowing waters and highlight the significant threat posed by H. verticillata to streams and waterways in New Zealand.     

New records of rotifers from the South Island lakes,New Zealand

Thirty-five lakes in the South Island of New Zealand were surveyed for rotifers during 1988–1991. Of 85 taxa identified, 31 are first records for New Zealand, bringing the rotifers recorded from the country to 331. Four species (Keratella australis, K. slacki, Lecane herzigi and L. tasmaniensis), previously recorded as endemic forms only in Australia, are now added to the New Zealand checklist. Several of the new records are photographed, and scanning electron micrographs of the trophi are shown. Comments are made on the Australasian endemics and rotifer biogeography in New Zealand.