Spread of Exotic Cordgrasses and Hybrids (Spartina sp.) in the Tidal Marshes of San Francisco Bay, California, USA

Four species of exotic cordgrass (Spartina sp.) occur in the San Francisco estuary in addition to the California native Spartina foliosa. Our goal was to map the location and extent of all non-native Spartina in the estuary. Hybrids of S. alterniflora and S. foliosa are by far the most numerous exotic and are spreading rapidly. Radiating from sites of deliberate introduction, S. alterniflora and hybrids now cover ca. 190 ha, mainly in the South and Central Bay. Estimates of rate of aerial increase range from a constant value to an accelerating rate of increase. This could be due to the proliferation of hybrid clones capable of rapid expansion and having superior seed set and siring abilities. The total coverage of 195 ha by hybrids and other exotic cordgrass species is slightly less than 1% of the Bay's tidal mudflats and marshes. Spartina anglica has not spread beyond its original 1970s introduction site. Spartina densiflora has spread to cover over 5 ha at 3 sites in the Central Bay. Spartina patens has expanded from 2 plants in 1970 to 42 plants at one site in Suisun Bay. Spartina seed floats on the tide, giving it the potential to export this invasion throughout the San Francisco estuary, and to estuaries outside of the Golden Gate. We found isolated plants of S. alterniflora and S. densiflora in outer coast estuaries north of the Bay suggesting the likelihood for the San Francisco Bay populations to found others on the Pacific coast.     

The rapid evolution of self-fertility in Spartina hybrids (Spartina alterniflora × foliosa) invading San Francisco Bay, CA

Plant hybridization can lead to the evolution of invasiveness. We wished to determine whether hybrids between the largely self-sterile Atlantic Spartina alterniflora and California native S. foliosa had evolved self-fertility during their ca 30 year existence in San Francisco Bay, CA. In pollination experiments we found that neither of the parental species was self-fertile, nor were early generation hybrids. A large fraction of later generation hybrids were profusely self-fertile. Inbreeding depression was high in the parental species and early generation hybrids, but was much reduced in later generation hybrids—some even showed outbreeding depression. We found that populations of later generation hybrids and their seedling progeny were almost two-fold more homozygous than early generation hybrids, consistent with the evidence of increased selfing shown by our parentage analyses based upon 17 microsatellite markers. We posit that evolved self-fertility has contributed substantially to the rapid spread of hybrid Spartina in San Francisco Bay.     

Hybridization between introduced smooth cordgrass (Spartina alterniflora; Poaceae) and native California cordgrass (S. foliosa) in San Francisco Bay, California, USA

Introduced Spartina alterniflora (smooth cordgrass) is rapidly invading intertidal mudflats in San Francisco Bay, California. At several sites, S. alterniflora co-occurs with native S. foliosa (California cordgrass), a species endemic to California salt marshes. In this study, random amplified polymorphic DNA markers (RAPDs) specific to each Spartina species were identified and used to test for hybridization between the native and introduced Spartina species in the greenhouse and in the field. Greenhouse crosses were made using S. alterniflora as the pollen donor and S. foliosa as the maternal plant, and these crosses produced viable seeds. The hybrid status of the crossed offspring was confirmed with the RAPD markers. Hybrids had low self-fertility but high fertility when back-crossed with S. foliosa pollen. Hybrids were also found established at two field sites in San Francisco Bay; these hybrids appeared vigorous and morphologically intermediate between the parental species. Field observations suggested that hybrids were recruiting more rapidly than the native S. foliosa. Previous work identified competition from introduced S. alterniflora as a threat to native S. foliosa. In this study, we identify introgression and the spread of hybrids as an additional, perhaps even more serious threat to conservation of S. foliosa in San Francisco Bay.     

Native and Introduced Ecosystem Engineers Produce Contrasting Effects on Estuarine Infaunal Communities

Cordgrasses in the genus Spartina are good examples of ecosystem engineers that modify habitat structure in estuaries throughout the world. In San Francisco Bay, California, USA, marshes containing native California cordgrass (Spartina foliosa) are being invaded by a hybrid (S. alterniflora × S. foliosa) formed after introduction of S. alterniflora. This study compared vegetation, sediment structure, and infaunal invertebrates in native and invaded marshes. We hypothesized that differences in the physical structure between S. foliosa and hybrid Spartina would be reflected in differences in density, biomass, diversity, and taxonomic composition of infauna. Hybrid Spartina modifies habitat structure more than S. foliosa by producing taller stems, and greater plant biomass both above- and belowground while occupying a much wider tidal range, thereby transforming open mudflats to a vegetated habitat. In general, S. foliosa areas contained significantly higher densities of benthic infauna than adjacent mudflats, while hybrid Spartina areas never contained greater infaunal densities than mudflats. This is because S. foliosa produces a moderate level of structure that can facilitate benthic invertebrates, whereas hybrid Spartina produces so much structure, particularly belowground, that it actually excludes invertebrates. Therefore, we suggest that these two closely related species both act as ecosystem engineers, but with opposing effects on invertebrate communities.     

Control and consequences of <Emphasis Type="Italic">Spartina</Emphasis> spp. invasions with focus upon San Francisco Bay

Maritime Spartina spp. are powerful ecosystem engineers that accrete sediment, define shorelines, create habitat, and generate prodigious primary productivity both where they are native and where they have been introduced. Invasive Spartina spp. can compete vigorously with native species, diminish biota, change hydrology, and confound human uses of estuaries. Herbicides have been effective in controlling several Spartina spp. invasions. One of the most recent successes is a 15-year campaign that has virtually eliminated S. alterniflora from the large, century-old invasion in Willapa Bay, WA, USA. Hybridization between native and introduced Spartina spp. has created new species and hybrid swarms. In San Francisco Bay, CA, USA (SF Bay) a complicated situation continues to play out from the purposeful introduction of S. alterniflora, which hybridized with native California cordgrass, S. foliosa. The hybrids spread rapidly and led to a long list of environmental problems, which led to an herbicide program that was successful in greatly diminishing the hybrid and saving the open mud habitat of migratory shorebirds. However, it was belatedly realized that the non-migratory, endangered Ridgeway’s rail uses the tall, dense hybrid Spartina as a surrogate for habitat that was lost during the twentieth century to urbanization and agricultural transformation of marshes around SF Bay. This realization has made difficult the simultaneous management of hybrid Spartina, wildlife conservation, and marsh restoration in San Francisco Bay. Restoration of native vegetation could satisfy the multiple goals of preserving open mud and conserving Ridgeway’s rail.     

Sexual reproduction of cordgrass hybrids (Spartina foliosa x alterniflora) invading tidal marshes in San Francisco Bay

We genetically analysed cordgrass plants and seedlings throughout the San Francisco, California, USA, estuary and found that hybrids between exotic Spartina alterniflora and native Spartina foliosa are the principal cordgrass invaders and colonizers. We hypothesized that this was due to higher seed set and siring ability by hybrids relative to the native species; too few alien parents remained in San Francisco Bay for our comparative studies. Hybrid seed comprised 91% to 98% of that set in the marsh study plants over the 2 years of the study. Total viable pollen production by hybrid plants was 400 times that of the native plants. Seed and pollen production were highly skewed towards a few hybrid genotypes. In addition to seed produced by hybrid plants, hybrid seed was produced by S. foliosa due to hybrid backcrossing. While the greatest advantage for hybrids was in pollen and seed production, hybrid seeds germinated, and seedlings survived and grew as well or better than the native species. As native S. foliosa becomes increasingly rare, hybrid seed floating on the tides will predominate, overwhelming recruitment sites and resulting in further colonization by hybrids. In an evolutionary context, hybrids with exceptional pollen and seed production will be initially favoured by natural selection, leading to the evolution of even more fertile hybrid genotypes.     

Does invasion of hybrid cordgrass change estuarine food webs?

Studies examining the impacts of introduced species on food webs often focus on the top-down effects of introduced predators. However, marine and estuarine systems have been invaded by plants that have the potential to alter carbon and nitrogen sources available to consumers. In San Francisco Bay, California, USA, hybridized cordgrass Spartina alterniflora × foliosa is adding C₄ carbon biomass to this system. We used natural abundances of stable isotopes of carbon and nitrogen to examine whether infaunal and epifaunal food webs reflected the large detrital input from hybrid Spartina. We compared stable isotope signatures among macrofaunal invertebrate consumers collected in hybrid Spartina, native S. foliosa, or unvegetated mudflats. We found no additional shift towards hybrid Spartina in hybrid areas. Structural changes brought about by an invasive ecosystem engineer, specifically increased biomass and detrital inputs, do not necessarily result in its increased incorporation into the food web.     

Reciprocal hybrid formation of Spartina in San Francisco Bay

Diversity in the tRNALEU1 intron of the chloroplast genome of Spartina was used to study hybridization of native California cordgrass, Spartina foliosa, with S. alterniflora, introduced to San Francisco Bay approximately 25 years ago. We sequenced 544 bases of the tRNALEU1 intron and found three polymorphic sites, a pyrimidine transition at site 126 and transversions at sites 382 and 430. Spartina from outside of San Francisco Bay, where hybridization between these species is impossible, gave cpDNA genotypes of the parental species. S. foliosa had a single chloroplast haplotype, CCT, and this was unique to California cordgrass. S. alterniflora from the native range along the Atlantic coast of North America had three chloroplast haplotypes, CAT, TAA, and TAT. Hybrids were discriminated by random amplified polymorphic DNA (RAPD) phenotypes developed in a previous study. We found one hybrid that contained a cpDNA haplotype unknown in either parental species (TCT). The most significant finding was that hybridization proceeds in both directions, assuming maternal inheritance of cpDNA; 26 of the 36 hybrid Spartina plants from San Francisco Bay contained the S. foliosa haplotype, nine contained haplotypes of the invading S. alterniflora, and one had the cpDNA of unknown origin. Furthermore, cpDNA of both parental species was distributed throughout the broad range of RAPD phenotypes, suggesting ongoing contributions to the hybrid swarm from both. The preponderance of S. foliosa cpDNA has entered the hybrid swarm indirectly, we propose, from F1s that backcross to S. foliosa. Flowering of the native precedes by several weeks that of the invading species, with little overlap between the two. Thus, F1 hybrids would be rare and sired by the last S. foliosa pollen upon the first S. alterniflora stigmas. The native species produces little pollen and this has low viability. An intermediate flowering time of hybrids as well as pollen that is more vigourous and abundant than that of the native species would predispose F1s to high fitness in a vast sea of native ovules. Thus, spread of hybrids to other S. foliosa marshes could be an even greater threat to the native species than introductions of alien S. alterniflora.     

Ascomycete fungal communities associated with early decaying leaves of Spartina spp. from central California estuaries

Ascomycetous fungi play an important role in the early stages of decomposition of Spartina alterniflora, but their role in the decomposition of other Spartina species has not been investigated. Here we use fingerprint (terminal restriction fragment length polymorphism) and phylogenetic analyses of the 18S to 28S internal transcribed spacer region to compare the composition of the ascomycete fungal communities on early decay blades of Spartina species (Spartina alterniflora, Spartina densiflora, Spartina foliosa, and a hybrid (S. alterniflora × S. foliosa)) collected from three salt marshes in San Francisco Bay and one in Tomales Bay, California, USA. Phaeosphaeria spartinicola was found on all samples collected and was often dominant. Two other ascomycetes, Phaeosphaeria halima and Mycosphaerella sp. strain 2, were also common. These three species are the same ascomycetes previously identified as the dominant fungal decomposers on S. alterniflora on the east coast. Ascomycetes appeared to exhibit varying degrees of host specificity, demonstrated by grouping patterns on phylogenetic trees. Neither the exotic S. alterniflora nor the hybrid supported fungal flora different from that of the native S. foliosa. However, S. densiflora had a significantly different fungal community than the other species, and hosted at least two unique ascomycetes. Significant differences in the fungal decomposer communities were also detected within species (two clones of S. foliosa), but these were minor and may be due to morphological differences among the plants.     

Characterization of microsatellite loci in Spartina species (Poaceae)

The cordgrasses in the genus Spartina have become model organisms for studying biological invasions from both ecological and genetic perspectives. Here we characterize 11 disomic loci in Spartina alterniflora that show promise for population studies and for studying hybridization events between S. alterniflora and S. foliosa. Comparisons among invasive and native S. alterniflora populations showed that levels of allelic variation are lower in invasive populations. In addition, nearly all loci that amplified in S. foliosa populations and in a swarm of S. alterniflora×foliosa hybrids were polymorphic. We also found that several loci amplified successfully in other Spartina species.     

Hybridization between invasive Spartina densiflora (Poaceae) and native S. foliosa in San Francisco Bay, California, USA

Rapid evolution in contemporary time can result when related species, brought together through human-aided introduction, hybridize. The significant evolutionary consequences of post-introduction hybridization range from allopolyploid speciation to extinction of species through genetic amalgamation. Both processes are known to occur in the perennial cordgrass genus, Spartina. Here we report the existence of a third recent Spartina hybridization, discovered in 2002, between introduced S. densiflora and native S. foliosa in San Francisco Bay, California, USA. We used nuclear and chloroplast DNA analysis and nuclear DNA content with chromosome counts to examine plants of morphology intermediate between S. densiflora and S. foliosa in a restored marsh in Marin County, California. We found 32 F(1) diploid hybrids and two triploid plants, all having S. densiflora and S. foliosa as parents; there is also evidence of a genetic contribution of S. alterniflora in some hybrids. None of these hybrids set germinable seed. In 2007 we found a hybrid over 30 miles away in a marsh where both parental species occurred, suggesting hybridization may not be a localized phenomenon. The presence of diploid and triploid hybrids is important because they indicate that several avenues existed that may have given rise to a new allopolyploid species. However, such an event is now unlikely because all hybrids are targets of eradication efforts.     

Greater male fitness of a rare invader (Spartina alterniflora, Poaceae) threatens a common native (Spartina foliosa) with hybridization

Hybridization with abundant invaders is a well-known threat to rare native species. Our study addresses mechanisms of hybridization between a rare invader, smooth cordgrass (Spartina alterniflora) and the common native California cordgrass (S. foliosa) in the salt marshes of San Francisco Bay. These species are wind-pollinated and flower in summer. The invader produced 21-fold the viable pollen of the native, and 28% of invader pollen germinated on native stigmas (1.5-fold the rate of the native's own pollen). Invader pollen increased the seed set of native plants almost eightfold over that produced with native pollen, while native pollen failed to increase seed set of the invader. This pollen swamping and superior siring ability by the invader could lead to serial genetic assimilation of a very large native population. Unlike California cordgrass, smooth cordgrass can grow into low intertidal habitats and cover open mud necessary to foraging shorebirds, marine life, navigation, and flood control in channels. To the extent that intertidal range of the hybrids is more similar to the invader than to the native parent, introgression will lead to habitat loss for shore birds and marine life as well to genetic pollution of native California cordgrass.     

Spatial and temporal genetic structure in a hybrid cordgrass invasion

Invasive hybrids and their spread dynamics pose unique opportunities to study evolutionary processes. Invasive hybrids of native Spartina foliosa and introduced S. alterniflora have expanded throughout San Francisco Bay intertidal habitats within the past 35 years by deliberate plantation and seeds floating on the tide. Our goals were to assess spatial and temporal scales of genetic structure in Spartina hybrid populations within the context of colonization history. We genotyped adult and seedling Spartina using 17 microsatellite loci and mapped their locations in three populations. All sampled seedlings were hybrids. Bayesian ordination analysis distinguished hybrid populations from parent species, clearly separated the population that originated by plantation from populations that originated naturally by seed and aligned most seedlings within each population. Population genetic structure estimated by analysis of molecular variance was substantial (F_ST=0.21). Temporal genetic structure among age classes varied highly between populations. At one population, the divergence between adults and 2004 seedlings was low (F_ST=0.02) whereas at another population this divergence was high (F_ST=0.26). This latter result was consistent with local recruitment of self-fertilized seed produced by only a few parental plants. We found fine-scale spatial genetic structure at distances less than ∼200 m, further supporting local seed and/or pollen dispersal. We posit a few self-fertile plants dominating local recruitment created substantial spatial genetic structure despite initial long-distance, human dispersal of hybrid Spartina through San Francisco Bay. Fine-scale genetic structure may more strongly develop when local recruits are dominated by the offspring of a few self-fertile plants.     

Effects of a habitat-altering invader on nesting sparrows: An ecological trap?

Many invading species impact native species through predation, parasitism or competition, while others affect natives indirectly by restructuring their habitat. How invasive plants affect native animals, and to what extent native animals respond to changes in their habitat and the novel selection pressures that follow, is not well known. We investigated the impacts of a habitat-altering invader, the Atlantic cordgrass Spartina alterniflora, on the nesting success of Alameda song sparrows (Melospiza melodia pusillula), a California Species of Special Concern, in tidal marshes in three sites in San Francisco Bay. Date of laying was the most influential factor in determining daily survival rate of nests, but whether the nest was placed in exotic Spartina was the most important ecological variable. Nests placed in exotic Spartina had a success rate that was 30% lower than those placed in native vegetation. Nests in exotic Spartina were significantly more likely to fail due to tidal flooding than were nests placed in native vegetation, because the densest stands of exotic Spartina occurred at significantly lower elevations relative to the tides. Our results suggest that exotic Spartina may be an ecological trap for song sparrows in San Francisco Bay, attracting birds to nest sites that are often destroyed by tidal flooding.     

Tidal and seasonal effects on survival rates of the endangered California clapper rail: does invasive Spartina facilitate greater survival in a dynamic environment?

Invasive species frequently degrade habitats, disturb ecosystem processes, and can increase the likelihood of extinction of imperiled populations. However, novel or enhanced functions provided by invading species may reduce the impact of processes that limit populations. It is important to recognize how invasive species benefit endangered species to determine overall effects on sensitive ecosystems. For example, since the 1990s, hybrid Spartina (Spartina foliosa × alterniflora) has expanded throughout South San Francisco Bay, USA, supplanting native vegetation and invading mudflats. The endangered California clapper rail (Rallus longirostris obsoletus) uses the tall, dense hybrid Spartina for cover and nesting, but the effects of hybrid Spartina on clapper rail survival was unknown. We estimated survival rates of 108 radio-marked California clapper rails in South San Francisco Bay from January 2007 to March 2010, a period of extensive hybrid Spartina eradication, with Kaplan–Meier product limit estimators. Clapper rail survival patterns were consistent with hybrid Spartina providing increased refuge cover from predators during tidal extremes which flood native vegetation, particularly during the winter when the vegetation senesces. Model averaged annual survival rates within hybrid Spartina dominated marshes before eradication (? = 0.466) were greater than the same marshes posttreatment (? = 0.275) and a marsh dominated by native vegetation (? = 0.272). However, models with and without marsh treatment as explanatory factor for survival rates had nearly equivalent support in the observed data, lending ambiguity as to whether hybrid Spartina facilitated greater survival rates than native marshland. Conservation actions to aid in recovery of this endangered species should recognize the importance of available of high tide refugia, particularly in light of invasive species eradication programs and projections of future sea-level rise.     

Geographic structure, genetic diversity and source tracking of Spartina alterniflora

Aim To examine the distribution and structure of genetic variation among native Spartina alterniflora and to characterize the evolutionary mechanisms underlying the success of non‐native S. alterniflora. Location Intertidal marshes along the Atlantic, Gulf and Pacific coasts of North America. Methods amova , parsimony analysis, haplotype networks of chloroplast DNA (cpDNA) sequences, neighbour‐joining analysis, Bayesian analysis of population structure, and individual assignment testing were used. Results Low levels of gene flow and geographic patterns of genetic variation were found among native S. alterniflora from the Atlantic and Gulf coasts of North America. The distribution of cpDNA haplotypes indicates that Atlantic coast S. alterniflora are subdivided into ‘northern’ and ‘southern’ groups. Variation observed at microsatellite loci further suggests that mid‐Atlantic S. alterniflora are differentiated from S. alterniflora found in southern Atlantic and New England coastal marshes. Comparisons between native populations on the Atlantic and Gulf coasts and non‐native Pacific coast populations substantiate prior studies demonstrating reciprocal interspecific hybridization in San Francisco Bay. Our results corroborate historical evidence that S. alterniflora was introduced into Willapa Bay from multiple source populations. However, we found that some Willapa Bay S. alterniflora are genetically divergent from putative sources, probably as a result of admixture following secondary contact among previously allopatric native populations. We further recovered evidence in support of models suggesting that S. alterniflora has secondarily spread within Washington State, from Willapa Bay to Grays Harbor. Main conclusions Underlying genetic structure has often been cited as a factor contributing to ecological variation of native S. alterniflora. Patterns of genetic structure within native S. alterniflora may be the result of environmental differences among biogeographical provinces, of migration barriers, or of responses to historical conditions. Interactions among these factors, rather than one single factor, may best explain the distribution of genetic variation among native S. alterniflora. Comprehensive genetic comparisons of native and introduced populations can illustrate how biological invasions may result from dramatically different underlying factors – some of which might otherwise go unrecognized. Demonstrating that invasions can result from several independent or interacting mechanisms is important for improving risk assessment and future forecasting. Further research on S. alterniflora not only may clarify what forces structure native populations, but also may improve the management of non‐native populations by enabling post‐introduction genetic changes and the rapid evolution of life‐history traits to be more successfully exploited.     

Variable reproductive output among clones of Spartina alterniflora (Poaceae) invading San Francisco Bay,California: the influence of herbivory,pollination, and establishment site

Spartina alterniflora has recently been introduced to San Francisco Bay, California, and is rapidly invading open mud flats, growing in circular patches that we found to be individual genetic clones. We collected spikelet samples from more than 200 clones and observed germination rates ranging from 0% to 59%, indicating substantial variation in reproductive output among clones. Several experiments were performed to explore the cause of variation. Pollination manipulations showed that S. alterniflora is outcrossing, but pollen supplements did not increase spikelet germination rates. Exclusion of the only insect herbivore (a phloem feeder, Prokelisia marginata) from developing inflorescences increased the proportion of spikelets containing seed, but failed to increase germinations per spikelet. Spikelets from Willapa Bay, Washington, grown free of insect herbivores, had germination rates similar to San Francisco Bay. These results suggest herbivory is not limiting reproductive output of S. alterniflora. Spikelet viability was not related to clone size; however, clones located lower in the intertidal or far up a drainage slough averaged fewer germinations per spikelet, suggesting clones in areas with lower genet density may have lower spikelet viabilities. Spikelet samples from different sections of clones growing across wide environmental ranges had similar rates of germination, suggesting some genetic influence on spikelet viability. Differential reproductive output among clones and the novel selective environment of San Francisco Bay are expected to cause gene frequency changes in this rapidly expanding population.     

Genetic and historical evidence disagree on likely sources of the Atlantic amethyst gem clam Gemma gemma (Totten, 1834) in California

Aim Historical information about source populations of invasive species is often limited; therefore, genetic analyses are used. We compared inference about source populations from historical and genetic data for the oyster‐associated clam, Gemma gemma that invaded California from the USA Atlantic coast. Location Mid‐Atlantic (North Carolina, Maryland), Northeastern (New Jersey, New York, Massachusetts) and the California coasts (Elkhorn Slough, San Francisco Bay, Bolinas Lagoon, Tomales Bay, Bodega Harbor). Methods The documented history of transplantation of Eastern oysters to California was reviewed. Cytochrome c oxidase subunit I (COI) sequences from recent and archived clams were examined in a haplotype network. We used AMOVA to detect geographic genetic structure and a permutation test for significant reductions in diversity. Results Chesapeake Bay oysters were transplanted to New York prior to shipment to San Francisco Bay and from there to peripheral bays. Gemma in the Northeastern and Mid‐Atlantic regions were genetically differentiated. In California, populations in Bodega Harbor and Tomales Bay were genetically similar to those in the Mid‐Atlantic area while clams in San Francisco Bay, Elkhorn Slough and Bolinas Lagoon resembled populations in the Northeastern region. In California, genetic variation was not highest in San Francisco Bay despite greater magnitude of oyster plantings. Haplotypes varied over time in native and introduced populations. Main Conclusions Historical records and inferences from genetics agree that both Northeastern and Mid‐Atlantic regions were sources for Gemma in California. Only complex genetic hypotheses reconcile the strong segregation of haplotypes in California to the historical evidence of mixing in their proximate source (New York). These hypotheses include sorting of mixtures of haplotypes or selection in non‐native areas. Haplotype turnover in San Francisco and Massachusetts samples over time suggests that the sorting hypothesis is plausible. We suggest, however, that Gemma was introduced independently and recently to Tomales Bay and Bodega Harbor.     

RESPONSE TO PHOTOPERIOD AND TEMPERATURE BY SPARTINA ALTERNIFLORA (POACEAE) FROM NORTH CAROLINA AND SPARTINA FOLIOSA FROM CALIFORNIA

Seedlings of Spartina alterniflora Loisel. from Oregon Inlet, North Carolina were generally taller and produced significantly more culms and total dry weight than those of S. foliosa Trin. from two California populations (Alameda Beach and Marin County) in eight photoperiod-temperature treatments over a 17-wk period. Seedlings of S. alterniflora produced maximal biomass in 30–26 C whereas those of S. foliosa produced maximal biomass in 22–18 C, both under long-day conditions. The average photosynthesis rate for S. alterniflora (3.0 mg C g dry wt^−- hr^−-1) was 1.6 times higher than that for S. foliosa (1.9 mg C g dry wt^−-1 hr^−-1), but the dark respiration rates (0.3 and 0.4 mg C g dry wt^−-1 hr^−-1, respectively) were not significantly different. The proportion of rhizomes was greater under short than long-day conditions for both species in most temperature regimes. The average shoot soluble carbohydrate and starch concentrations were higher for S. foliosa (8.4 and 0.9%, respectively) than those for S. alterniflora (6.0 and 0.4%, respectively). The average rhizome soluble carbohydrate concentration (18%) for S. alterniflora under long-day conditions was significantly lower than that under short-day conditions (28%) and also lower than those for S. foliosa under both long (26%) and short-day (25%) conditions. Rhizome starch concentrations of S. alterniflora were significantly higher in the short than in most long-day temperatures and were generally higher for S. alterniflora than for S. foliosa under short-day conditions. The root starch concentration of S. alterniflora under short-day conditions (1.3%) was higher than that under long-day conditions (0.2%) and also higher than those of S. foliosa under both long (0.2%) and short-day (0.7%) conditions. The two species exhibited similar patterns of carbohydrate storage in belowground organs, similar flower initiation processes not under strict photoperiod control, and similar respiration rates, but significantly different photosynthetic rates and growth responses with S. alterniflora having the potential to be the more productive species.