Herbivory and intraspecific competition in a stream caddisfly population

Summary Field experiments were conducted to assess the effects of different population densities of the herbivorous caddisfly Helicopsyche borealis on periphyton biomass and on its own growth rate in Big Sulphur Creek, a third-order stream in northern California, USA. Stream enclosures were used to vary grazer density from one-eighth to twice natural density (1/8X–2X) in two experiments (35 d and 60 d), which spanned the period of most rapid larval growth. Periphyton biomass and chlorophyll a were inversely related to grazer density. Grazer densities of 1/8X–1/2X moderately reduced periphyton when compared to an ungrazed control, whereas densities of 1X–2X greatly depleted periphyton. The growth rate of H. borealis larvae declined with increasing larval density. Growth rate was highest at densities of 1/8X–1/2X; larvae grew more slowly at 1X and showed no growth at 2X. Weekly supplementation of periphyton to 1/4X, 1X, and 2X densities significantly increased the final body mass of larvae at 1X and 2X (by 26% and 50%, respectively, compared to unsupplemented larvae) but did not change the body mass of larvae at 1/4X. These results suggest that periphyton is a limiting resource in Big Sulphur Creek and that H. borealis larvae compete exploitatively for that resource. Intraspecific competition may be an important, but often overlooked, feature of many herbivore populations in streams.     

Periphyton responses to invertebrate grazing and riparian canopy in three northern California coastal streams

SUMMARY. 1. Field experiments were conducted to examine the impact of grazing invertebrates on periphyton biomass in twenty-one pools across three northern California coastal streams (U.S.A.): Big Sulphur Creek, the Rice Fork of the Eel River, and Big Canyon Creek. Periphyton accrual on artificial substrate tiles was compared in each stream between two treatments: those elevated slightly above the stream bottom to reduce access by grazers (= platforms) and those placed directly on the stream bottom to allow access by grazers (=controls).
2. Crawling invertebrate grazers (cased caddisflies and snails) were numerically dominant in each stream (86% of all grazers in Big Sulphur Creek, 61% in the Rice Fork, 84% in Big Canyon Creek). Platforms effectively excluded crawling grazers, but were less effective in excluding swimming mayfly grazers (Baetidae).
3. Periphyton biomass (as AFDM) on tiles was significantly lower on controls compared to platforms for the Rice Fork, an open-canopy stream, and Big Sulphur Creek, a stream with a heterogeneous canopy. In contrast, no grazer impact was found for Big Canyon Creek, a densely shaded stream. Here, extremely low periphyton biomass occurred for both treatments throughout the 60 day study.
4. The influence of riparian canopy on periphyton growth (i.e. accrual on platforms), grazer impact on periphyton, and grazer abundance was examined for Big Sulphur Creek. As canopy increased (15–98% cover), periphyton biomass on platforms decreased. In contrast, canopy had little influence on periphyton accrual on controls; apparently, grazers could maintain low periphyton standing crops across the full range of canopy levels. The abundance of one grazer species, the caddisfly Gumaga nigricula , was highest in open, sunlit stream pools; abundance of two other prominent grazers, Helicopsyche borealis (Trichoptera) and Centroptilum convexum (Ephemeroptera), however, was unrelated to canopy.     

ELECTROPHORETIC ENZYME ANALYSIS OF NORTH AMERICAN AND EASTERN ASIAN POPULATIONS OF AGASTACHE SECT. AGASTACHE (LABIATAE)

Agastache sect. Agastache consists of seven species in North America and one disjunct in eastern Asia. Starch-gel electrophoresis of enzymatic proteins was employed to assess genetic relationships among these species and to estimate the amount of genetic divergence between the North American and Asian populations. Species of the western United States appear to be better adapted for outcrossing than are the others and are much more genetically variable, with higher levels of heterozygosity per individual, more alleles per species, and higher percentages of polymorphic loci per population. Nonmetric multidimensional scaling of Nei's genetic distances among 32 populations partitioned the section into four discrete groups: 1) A. nepetoides (eastern North America), 2) A. scrophulariifolia and A. foeniculum (eastern and central North America), 3) the four species of the western United States (A. urticifolia, A. occidentalis, A. parvifolia and A. cusickii) and 4) A. rugosa (eastern Asia). Asian Agastache, separated from its American congeners for over twelve million years, differed from American populations at two of fifteen loci surveyed. Nei's genetic distances between Asian and North American populations ranged from 0.2877 to 0.6734.     

Description of a New Helicopsyche Species from China (Trichoptera: Helicopsychidae)

Helicopsyche zhejiangensis sp. n. from southeastern China is described and figured, representing the first record of this family in China. The species has a general resemblance with Helicopsyche species known from the Oriental Region, and the East Palaearctic species Helicopsyche coreana Mey from North Korea.     

Description of Two New Neotropical Helicopsyche (Trichoptera: Helicopsychidae)

Helicopsyche flinti sp.n. and Helicopsyche cochleara sp.n. are described from Brazil and Ecuador. H. cochleara sp.n. has certain affinities with H. planorboides Machado from eastern Brazil. H. flinti sp.n. is related with other Neotropical Helicopsyche.     

GENETIC POPULATION STRUCTURE AND LEVELS OF GENE FLOW IN THE STREAM DWELLING WATERSTRIDER,AQUARIUS (=GERRIS) REMIGIS (HEMIPTERA: GERRIDAE)

Gene flow, in combination with selection and drift, determines levels of differentiation among local populations. In this study we estimate gene flow in a stream dwelling, flightless waterstrider, Aquarius remigis. Twenty-eight Aquarius remigis populations from Quebec, Ontario, New Brunswick, Iowa, North Carolina, and California were genetically characterized at 15 loci using starch gel electrophoresis. Sampling over two years was designed for a hierarchical analysis of population structure incorporating variation among sites within streams, streams within watersheds, watersheds within regions, and regions within North America. Hierarchical F statistics indicated that only sites within streams maintained enough gene flow to prevent differentiation through drift (Nm = 27.5). Above the level of sites within streams gene flow is highly restricted (Nm ≤ 0.5) and no correlation is found between genetic and geographic distances. This agrees well with direct estimates of gene flow based on mark and recapture data, yielding an N_e of approximately 170 individuals. Previous assignment of subspecific status to Californian A. remigis is not supported by genetic distances between those populations and other populations in North America. Previous suggestion of specific status for south-eastern A. remigis is supported by genetic distances between North Carolina populations and other populations in North America, and a high proportion of region specific alleles in the North Carolina populations. However, because of the high degree of morphological and genetic variability throughout the range of this species, the assignment of specific or subspecific status to parts of the range may be premature.     

Genetic variation in Batrachospermum helminthosum (Batrachospermales, Rhodophyta) among and within stream reaches using intersimple sequence repeat molecular markers

Intersimple sequence repeat molecular markers were utilized to investigate the genetic relationship among individual gametophyte thalli of the freshwater red alga Batrachospermum helminthosum among distant stream reaches and within a stream reach. Fifteen thalli per stream reach were sampled from 11 streams throughout the known distribution in eastern North America from three locations in Ohio, and one location in Indiana, Michigan, North Carolina, Tennessee, Louisiana, Rhode Island, Massachusetts and Connecticut. The pairwise φ_ST analysis showed significant genetic differences (P<0.05) among all streams. The partitioning of genetic variation was almost equal within and among streams (45% and 55%, respectively). Genetic variation among populations did not reflect geographic distance, suggesting that long distance dispersal may be important in the distribution of this alga. Fifty‐eight individuals from Five Mile Creek, Ohio, were investigated from three distinct segments of the stream (upstream, middle and downstream) to examine small‐scale genetic variation within a stream reach. A much higher proportion of genetic variation was observed within a stream segment (79%) than among the three segments (21%). Among these stream segments within Five Mile Creek, the individuals were variously related, with genetic similarity and spatial distance uncorrelated. A similar result was obtained among individuals within other stream reaches, suggesting that genetic structure within a stream reach may be more complex than just reproduction among closely located individuals.     

Helicopsyche trispina sp. n. (Trichoptera,Helicopsychidae) from New Caledonia

Helicopsyche trispina sp. n. is described from Grande Terre, New Caledonia, based on pharate males, larvae and pupae. The species appears to belong to the monophyletic New Caledonian Helicopsyche clade.     

Herbivorous caddisflies,macroalgae, and epilithic microalgae: dynamic interactions in a stream grazing system

Summary 1. During the low-flow period (April–October) in sunlit pools of Big Sulphur Creek (northern coastal California), the attached algal community predictably changes from an assemblage dominated by lush, upright Cladophora glomerata filaments in spring and early summer to one dominated by epilithic diatoms and blue-green algae (together=microalgae) in late summer through early autumn. Previous studies in this stream indicated that grazing by the caddisflies Helicopsyche borealis and Gumaga nigricula maintain low algal biomass during the latter part of this period. We used a combination of in situ exclusion/enclosure experiments to examine (1) the separate and combined effects of these grazers on Cladophora and microalgal assemblages, and (2) food preferences, growth, and microdistribution patterns of grazers when offered these different algal foods. 2. Grazers exerted strong but divergent effects on algal assemblages. Selective grazing on Cladophora by G. nigricula greatly accelerated the transition from upright Cladophora to epilithic microalgae, whereas selective grazing on microalgae by H. borealis dramatically reduced biomass of these forms. Grazers were largely ineffective at reducing the non-preferred algal food source (i.e. Cladophora by H. borealis, microalgae by G. nigricula). In the case of each grazer, growth was highest on the preferred algal food. Together, the activity of these grazers produced a low-biomass assemblage dominated by microalgal cells. 3. Removal of the Cladophora overstory by G. nigricula resulted in a three-fold increase in the abundance of epilithic microalgae, the preferred food of H. borealis. Elimination of Cladophora by G. nigricula can increase food availability for H. borealis and, in so doing, can indirectly facilitate the growth of this grazer during food-limited conditions. However, microdistribution of G. nigricula shifts from high overlap with H. borealis in spring and early summer when Cladophora is abundant to low overlap in late summer after Cladophora has been eliminated. This may indicate intense competition between these species for limited epilithic algae, and a concomitant movement by G. nigricula to areas in the stream where food resources are more available.     

Lag in Stream Channel Adjustment to Livestock Exclosure, White Mountains, California

Livestock have been excluded from riparian zones along many streams in western North America in an effort to restore aquatic and riparian habitat degraded by livestock grazing. Within these exclosures, channel adjustment to elimination of grazing pressure may lag behind plant recovery because of the time required to deposit sediment along the vegetated banks of the stream channel. Moreover, unless grazing is eliminated from the watershed, the channel within the exclosure must still accommodate increased runoff and sediment loads from upstream. This hydrologic regime may prevent a return to predisturbance channel morphology. Cross sections of the North Fork Cottonwood Creek in the White Mountains of California showed no significant difference in channel width within and downstream of a 24-year-old exclosure, despite a lush growth of stream bank vegetation that gives the impression of a narrower channel within the exclosure.     

Weedy adaptation in Setaria spp. II. Genetic diversity and population genetic structure in S. glauca,S. geniculata,and S. faberii (Poaceae)

Setaria glauca (yellow foxtail), S. geniculata (knotroot foxtail), and S. faberii (giant foxtail) are important cosmopolitan weeds of temperate and tropical regions. Isozyme markers were used to investigate genetic diversity and population genetic structure in 94 accessions of yellow foxtail, 24 accessions of knotroot foxtail, and 51 accessions of giant foxtail, collected mainly from North America and Eurasia. Giant foxtail populations were nearly identical genetically, with only one population exhibiting isozyme polymorphism. Yellow and knotroot foxtail populations had low genetic diversity but marked population differentiation. Although the latter species are similar morphologically, they are readily distinguished electrophoretically, with Nei's genetic identity being 0.83. In both species, genetic divergence between accessions from Eurasia and North America was minimal. Populations from the native ranges had slightly greater genetic diversity than those from the respective introduced ranges. Yellow foxtail populations genetically clustered into Asian, European, and North American groups. Within North America, yellow foxtail populations from Iowa were genetically diverse whereas populations collected from other North American locations were nearly monomorphic for the same multilocus genotype. Knotroot foxtail populations in North America were genetically differentiated into northern and southern groups on either side of a line at ≈37° N latitude. No genetic patterning was evident in knotroot foxtail populations from Eurasia. In both yellow and knotroot foxtail, patterns of population genetic structure have been influenced by several factors, including genetic bottlenecks associated with founder events, genetic drift, and natural selection.     

ISOZYME DIVERGENCE BETWEEN EASTERN ASIAN,NORTH AMERICAN,AND TURKISH SPECIES OF LIQUIDAMBAR (HAMAMELIDACEAE)

The deciduous woody genus Liquidambar has four morphologically similar species in eastern and western Asia, eastern North America, and Central America. Liquidambar styraciflua is found in the eastern United States and Central America, L. orientalis is native only to southwest Turkey, and L. formosana and L. acalycina occur in eastern Asia. This genus is one of many that contributes to the floristic similarities observed between these different regions. Allelic variation was scored at 22 isozyme loci from 41 populations. The level of genetic divergence between species on different continents is high. Nei's genetic identity was 0.431 between L. formosana and L. styraciflua, 0.485 between L. acalycina and L. styraciflua, 0.512 between L. orientalis and L. styraciflua, 0.256 between L. formosana and L. orientalis, and 0.305 between L. acalycina and L. orientalis. Estimates of time of divergence from the isozyme data suggest that the current species diverged before or during the Miocene. The pattern of relationships portrayed by the isozyme data suggest a longer period of separation between the eastern and western Asian forms of this genus. In addition, the eastern North American and Turkish species appear to be the most closely related intercontinental pair of species providing evidence for a North Atlantic land bridge as late as the Miocene. It would appear, therefore, that the North American populations were in contact with the Asian populations over the North Pacific and North Atlantic possibly as late as the Miocene, but that the separation between the two Asian populations occurred much earlier. The time of divergence as measured from the isozyme data correlates with an independent assessment of the origin of these disjuncts as determined from the fossil record.     

Seasonal diversity in aquatic insect communities in an all-year stream system

Summary The general pattern of the data in this study shows that a riffle zone carries more species and, usually, more organisms/m² than a slower current zone. The slower current zone, however, is still apparently able to support a larger number of organisms and species than a pool of standing water. In the West Fork of the Oak Creek system the dominant insects controlling diversity are the Trichoptera, particularly one species of Helicopsyche. Diversity in the fast current zone is high in the summer, and low during other seasons. Diversity in the slow current zone is highest in spring, but is also fairly high in summer. This reflects to a large extent the seasonal fluctuations in abundance of the dominant Trichopterans in the two habitats.     

Molecular and morphological divergence in the butterfly genus Lycaeides (Lepidoptera: Lycaenidae) in North America: evidence of recent speciation

Male genital morphology, allozyme allele frequencies and mtDNA sequence variation were surveyed in the butterfly species Lycaeides idas and L. melissa from across much of their range in North America. Despite clear differences in male genital morphology, wing colour patterns and habitat characteristics, genetic variation was not taxonomically or geographically structured and the species were not identifiable by either genetic data set. Genetic distances (Nei's D=0.002–0.078, calculated from allozyme data) between all populations of both species were within the range commonly observed for conspecific populations of other butterflies. The most frequent mtDNA haplotype was present in individuals of both species in populations from southern California to Wisconsin. We conclude that speciation has probably happened recently and the lack of genetic differentiation between the species is the product of either (1) recent or ongoing gene flow at neutral loci, and/or (2) an insufficiency of time for lineage sorting. The evolution of male genital morphology, wing colour patterns and ecological characteristics has proceeded more rapidly than allozyme or mtDNA evolution.     

Independent and interactive effects of nutrients and grazers on benthic algal community structure

Algal responses to nutrients, grazing by Helicopsyche borealis, and concurrent grazing by Helicopsyche and Baetis tricaudatus were examined in recirculating stream chambers. Alagl communities, dominated by Achnanthes minutissima, Cocconeis placentula, and Synedra ulna, were primarily phosphorus-limited. Algal populations responded after only 6 days of nutrient enrichment. Initially, both the adnate diatom Cocconeis and erect diatom Synedra showed positive response to nutrient enrichment. Accumulation of algal biomass between day 3 and 6 in the P enriched treatment was resulted primarily from the growth of Synedra, an overstory rosette-like diatom colony. Such a shift in dominant growth from adnate to erect diatoms is a general phenomenon in periphyton succession in the absence of disturbance. Algal species showed differential responses to an increase of Helicopsyche densities. The accrual rate of Achnanthes continuously decreased with increasing grazer densities. The accrual rates of both Cocconeis and Synedra declined but reached plateaus between medium and high grazing densities. Baetis effectively and exclusively depressed Synedra and had no significant impact on Cocconeis. After concurrent grazing, algal communities were mainly dominated by Cocconeis (approximately 80% of total algal biovolume). The grazer' s mouth structures, grazing efficiencies, and mobility may account for the differential effects of concurrent grazing on algal communities. Significant interactive effects of P and grazing by Helicopsyche indicated that both nutrient addition and grazing may exert significant impact on algal communities. However, grazing may have a much stronger effect on algae than nutrients. Our results indicate that enhancement of algal biomass by P was dampened by grazing activities and that P had no effect on algal biomass in the presence of grazers.     

PHYLOGEOGRAPHY OF BATRACHOSPERMUM MACROSPORUM (BATRACHOSPERMALES,RHODOPHYTA) FROM NORTH AND SOUTH AMERICA1

Phylogeographic trends in Batrachospermum macrosporum Mont. were investigated using the mitochondrial intergenic spacer between the cytochrome oxidase subunit 2 and 3 genes (cox2‐3). A total of 11 stream segments were sampled with seven in the coastal plain of North America and four in tropical areas of South America. Fifteen thalli were sampled from seven streams, 14 thalli from two streams, and eight thalli from two streams. There were 16 haplotypes detected using 149 individuals. Of the eight haplotypes from locations in North America, all were 334 base pairs (bp) in length, and of those from South America, five were 344 bp, and three were 348 bp. Two individual networks were produced: one for the haplotypes from North America and another for those from South America, and these could not be joined due to the large number of base pair differences. This split between haplotypes from North and South America was confirmed with sequence data of the rbcL gene. There was very little genetic variation among the haplotypes from the North American locations, leading us to hypothesize that these are fairly recent colonization events along the coastal plain. In contrast, there was high variation among haplotypes from South America, and it would appear that the Amazon serves as a center of diversity. We detected considerable variation in haplotypes among streams, but frequently, a single haplotype in an individual stream segment, which is consistent with data from previous studies of other batrachospermalean taxa, may suggest a single colonization event per stream.     

Myriophyllum quitense (Haloragaceae) in the United States

A systematic study of more than 15,000Myriophyllum specimens from 189 North American herbaria yielded 63M. quitense specimens. Late 19th century collections were made from Henry's Fork of the Snake River in Jefferson County, Idaho, in 1872, from Santa Cruz Creek, Tucson, Pima County, Arizona, in 1881, from Yellowstone Lake in 1885, and from Crook County, Oregon, in 1894. Twentieth century collections have been made from Yellowstone National Park, at several sites along the Snake River in Idaho, in Summit County, Utah, in Lassen and Shasta counties of northern California, at numerous sites along the Deschutes River, and in Benton County, Oregon, and at three sites in Washington. We contend that the species was introduced into North America by migratory waterfowl more than 100 years ago. Only in the Yellowstone Lake and numerous Deschutes River habitats in which populations now exist are environmental conditions conducive to full morphological development of the species in North America.     

WEEDY ADAPTATION IN SETARIA SPP. I. ISOZYME ANALYSIS OF GENETIC DIVERSITY AND POPULATION GENETIC STRUCTURE IN SETARIA VIRIDIS

Setaria viridis is an important self-pollinating, cosmopolitan weed of temperate regions worldwide. Allozyme markers were used to investigate genetic diversity and structure in 168 accessions (including four S. italica) collected mainly from North America and Eurasia. Genetic diversity in green foxtail, and its population genetic structure, provided important clues about this weed's evolutionary history. Genetic diversity was low, with marked population differentiation: the percentage of polymorphic loci was 25% (0.95 criterion); mean number of alleles per locus was 1.86; mean panmictic heterozygosity was 0.07; and the coefficient of population genetic differentiation was 0.65. A common genotype occurred in 25 accessions distributed in six countries from both the Old World and New World, in a wide variety of ecological situations. Relatively little genetic divergence occurred between Eurasia and North America, with Nei's unbiased genetic identity between the two regions equaling 1.0. Populations from these two continents also had equivalent genetic diversity. Within North America, regional differentiation was indicated by northern and southern groups separated at 43.5° N latitude. No geographic pattern in genetic diversity was found within Eurasia. The size of the geographic range from which populations were sampled was not an accurate indicator of the extent of genetic diversity found among populations from that region. These results suggest that present patterning among green foxtail populations in North America is the consequence of multiple introductions into the New World followed by local adaptation and regional differentiation. Finally, S. italica and several green foxtail varieties did not differ isozymatically from typical forms of green foxtail. This supports the view that S. italica and S. viridis are conspecific, that the former (foxtail millet) is a domesticated form of the latter, and also questions the taxonomic validity of formally recognizing morphological varieties within green foxtail.     

MOLECULAR DIVERGENCE BETWEEN ASIAN AND NORTH AMERICAN SPECIES OF LIRIODENDRON (MAGNOLIACEAE) WITH IMPLICATIONS FOR INTERPRETATION OF FOSSIL FLORAS

Botanists have long been aware of the floristic similarities between eastern Asia and eastern North America. Most who have considered this classic disjunction pattern have suggested that it arose through range disruption of a flora that was once more widely distributed in the Northern Hemisphere. There is less agreement on the timing of this process, with suggestions ranging from the Paleocene to the Neogene. In this study, molecular markers from two different plant genomes were used to assess the degree of genetic divergence between the two interfertile, morphologically similar species of the genus Liriodendron, i.e., L. tulipifera and L. chinense. Resulting molecular divergence estimates were translated into approximate dates of separation, independent of evidence from the fossil record. Allozyme data (Nei's genetic identity = 0.434) suggested a divergence time of 10–16 million years before present, whereas sequence divergence in the plastid genomes (1.24%) led to an estimate of approximately 11–14 million years before present. A review of the paleobotanical literature indicated that the fossil floras that included, or might have included Liriodendron could not have survived in Beringia after the late Miocene and the onset of southward-migrating Arctic air masses on the North American continent. This interpretation suggests a minimum time of separation of approximately 13 million years before present. Thus, both molecular data sets and the paleobotanical evidence concur in suggesting a divergence time of 10–16 million years before present. Interspecific compatibility and relative morphological stasis must have, therefore, persisted from at least the late Miocene. We emphasize the need for similar studies in other genera, especially those that have both a reasonable Tertiary fossil history and extant species in mesic temperate refugia in Asia, Europe, and western as well as eastern North America.     

GENETIC STRUCTURE AND EVOLUTION OF SPECIES IN THE MANGROVE GENUS AVICENNIA (AVICENNIACEAE) IN THE INDO-WEST PACIFIC

Allozyme variation in species of the mangrove genus Avicennia was screened in 25 populations collected from 22 locations in the Indo-West Pacific and eastern North America using 11 loci. Several fixed gene differences supported the specific status of Avicennia alba, A. integra, A. marina, and A. rumphiana from the Indo-West Pacific, and A. germinans from the Atlantic-East Pacific. The three varieties of A. marina, var. marina, var. eucalyptifolia, and van australasica, had higher genetic similarities (Nei's I) and no fixed gene differences, confirming their conspecific status. Strong genetic structuring was observed in A. marina, with sharp changes in gene frequencies at the geographical margins of varietal distributions. The occurrence of alleles found otherwise in only one variety, in only immediately adjacent populations of another variety, provided evidence of introgession between varieties. The varieties appear to have diverged recently in the Pleistocene and are apparently not of ancient Cretaceous origin, as suggested earlier. Despite evidence of high degrees of outcrossing, gene flow among populations was relatively low (N_em < 1–2), except where populations were geographically continuous, questioning assumptions that these widespread mangrove species achieve high levels of long-distance dispersal.