Killingly Revisited (Images of America)
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Although culling the identical data reduced the number of sequences analyzed to 11, trees were redrawn to include the previously excluded, identical sequences depicted as a polytomy, a method that enabled us to represent the results obtained for all accessions in one diagram.
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For comparison, we also measured leaves and internodes on nine Australian specimens of G. To reduce developmental variability, all measurements were restricted to mature leaves in exclusion of any terminally occurring leaves. Means and standard errors were calculated for all measurements.
Differences in mean leaf and rhizome length between North American plants bearing chasmogamous flowers and those having only cleistogamous flowers were evaluated statistically using t tests as implemented with SYSTAT Three analyses were conducted to compare 1 North American plants both emersed and submersed with Australian material of G. Distribution maps were plotted to depict the native Australian ranges of G. We recorded the occurrence of every observable Glossostigma patch for all Connecticut localities by conducting comprehensive field surveys.
Although lakes were not surveyed as extensively at other localities NJ, PA, RI , we obtained GPS data for at least some reference patches and evaluated the relative abundance of patches based on a survey of roughly comparable lengths of shoreline c. A detailed map showing the distribution of patches was made for the Mansfield Hollow, CT site, which had been surveyed comprehensively using GPS. The density of plants per patch was estimated both quantitatively and qualitatively.
In determining our estimates, each leaf pair i. During surveys of each North American site visited, Glossostigma density was estimated in representative patches and assigned visually to one of three categories: high, medium or low. Digital photographs were taken of plants within 0. The number of individuals then was counted on each photograph and used to compute representative values of plant density per square meter.
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For one locality Alexander Lake, CT , the average number of flowers per node and mean number of seeds per fruit were calculated and used along with density data to estimate frequency of cleistogamous flowers in the population and overall plant reproductive potential. Substrate composition by visual inspection and associated species were recorded for each site visited. For CT, we analyzed differences in environmental conditions that existed between five lakes where Glossostigma occurred and lakes surveyed during June—October and where it does not occur Connecticut Agricultural Experiment Station, New Haven Connecticut, ; unpublished data from the Invasive Aquatic Plant Survey Program.
Total P data were not available for one lake where Glossostigma occurred.
The sequences uniformly yielded chromatograms with no evidence of site polymorphisms. For ETS we analyzed a region of contiguous, aligned sites. Again, there was no evidence of any site polymorphisms in the sequence chromatograms. However, aside from their nucleotide identity, the North American Glossostigma ETS sequences did lack a single nucleotide gap that occurred in the published sequence of G. This single gap was the only difference detected between North American Glossostigma accessions and G.
A specimen of G. The maximum parsimony tree clearly showed that all Glossostigma material originating from North America was associated with native material of G. Leaves on submersed and cleistogamous plants of North American Glossostigma were significantly longer 7. Leaves on six specimens of Australian G.
Leaf lengths of three Australian specimens of G. Leaf length varied considerably among G. Within submersed North American populations, mean leaf length varied from a low of Rhizome internode length of North American G. A similar trend was not evident in Australian material of G.
Chasmogamous flowers occurred only on plants collected in emersed condition, whereas submersed plants produced only cleistogamous flowers. Patch distributions ranged from relatively localized to widespread within six intensively surveyed CT populations Table 1. Jacono s. Patch density estimates for all sites are summarized in Table 1.
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Using bathymetry data for the lake, we estimated that the total area occupied by Glossostigma in the Mansfield Hollow reservoir represented 1. Glossostigma grew in association with 29 aquatic species Table 2. Two taxa Eleocharis acicularis , Elatine spp. Inspection of many submerged plants showed cleistogamous flowers to be extremely abundant. Of nodes examined, only 15 2. Seed germination without cold stratification was low after 30 days with only 3 of seeds 1. However, even if the actual germination rate typically is this low, then the denser patches in populations such as Alexander Lake, CT potentially could yield over new individuals genets per meter of plants, depending on their establishment success.
A Mann—Whitney test, which determines whether measured values of variables differed between lakes with and without G.
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Thus, in North America, G. Correct species identification is essential for evaluating the introduction of any nonindigenous organism. This task is complicated by the fact that newly introduced species and even some of their genera, families, etc. The introduction of Glossostigma to North America epitomizes these problems.
The genus is native entirely to the Old World and never has been monographed. Generally, Glossostigma species are quite similar; they are minute, simple morphologically, and have few characters useful for identification. Beardsley and Barker attributed 6—8 Glossostigma species to Australia; however, the genus is not included in Aquatic Plants of Australia or its supplement Aston, , , and no comprehensive taxonomic treatment of the genus exists for the country.
Rather, sets of different species have been treated in various regional floras of the continent e. Barker, an authority on Australian Scrophulariaceae and Glossostigma in particular , who identified the plants as G.
Rhoads, Morris Arboretum, personal observation. However, we found that the published descriptions of G. Primarily, the morphologically similar species G.
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We also considered the possibility that both G. This possibility gained credibility when we established that plants with chasmogamous flowers also possessed significantly shorter leaves 4. These observations were consistent with reports that leaves of G. However, this hypothesis was not substantiated by further morphological analyses. First, our measurements of Australian specimens of G. Furthermore, the North American specimens fell well within the range of leaf length observed for Australian material of G.
Adding to the uncertainty were comments by Barker b who reported that G. He also remarked that pedicels of the cleistogamous flowers in G. Thus, it would be extremely difficult to identify with certainty any Glossostigma plant that possessed both sessile and pedicellate flowers as in several of our cases , especially if plants were in fruiting condition where other floral details would be lacking.
Observing that a number of North American plants possessed both pedicellate, chasmogamous, and nearly sessile, cleistogamous flowers Fig. Fortunately, a genetic means to facilitate species identification was provided by the publication of DNA sequence data for all known Glossostigma taxa occurring in Australia Beardsley and Barker, Because most of the taxa were sufficiently divergent for the loci surveyed nrITS, nrETS, trnL that data set enabled species determinations to be made using a comparative sequencing approach.
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Thus, by analyzing the DNA sequences of Glossostigma material collected from the introduced, North American populations and comparing them to the sequences obtained for the native Australian populations, we were able to achieve an identification that was based on data that were independent from their morphology. We found that the DNA sequences of all North American populations surveyed, including those plants with cleistogamous flowers, chasmogamous flowers, or mixtures of both floral types, matched the sequence reported for G. Although G. In contrast, multiple accessions of G.
Furthermore, the complete identity of DNA sequences obtained from the two loci ETS, ITS surveyed across all North American accessions enabled us to conclude that only one species had been introduced to the continent and that it was G.
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