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Larval settlement into marine soft-sediment systems: Interactions with the meiofauna

The distribution and abundance of organisms exhibiting complex life cycles, like marine invertebrates, reflect ecological processes operating at all stages in the life history. Few studies have yet considered the mortality patterns of larvae and juveniles. To test whether the permanent meiofauna play a role in macrofaunal larval site-selection, and/or if permanent meiofaunal predators or competitors and disturbers are a significant source of mortality for newly settled macrofaunal juveniles, a series of manipulations of meiofauna were performed. Densities of (1) predatory turbellarians alone, and (2) other meiofauna, were increased in small boxes placed on a holding table in a shallow subtidal habitat. Natural settlement into meiofauna manipulations was examined approximately monthly for a year. Other experiments were conducted to examine the relative importance of site-selection and post-settlement mortality and to determine if settlement was density-dependent or density-independent. Densities of spionid, cirratulid, terebellid, capitellid, and maldanid polychaetes, oligochaetes, bivalves, and amphipods were all reduced by high densities of turbellarians, and sometimes by other meiofauna. Turbellarian predation was responsible for significant post-settlement mortality, but spionids, cirratulids, terebellids, and maldanids also appeared to avoid dense turbellarian treatments. Bivalve densities were reduced by high densities of other meiofauna, probably due to their sedimentary disturbance. Syllid polychaete densities were higher in turbellarian treatments, suggesting that they select for areas with high densities of their prey (small turbellarians). The effects of meiofauna on settlement and survival were greatest in spring and summer. The results of these experiments suggest larval settlement patterns and early juvenile survival should be considered in developing general models of life history and community structure for soft-sediment systems.

Journal of Experimental Marine Biology and Ecology↗

Effects of kelp forest removal on associated fish assemblages in central California

Visual surveys along subtidal belt transects were used to compare fish assemblages on an experimental and a control site before and after the removal of a canopy-forming kelp forest. The giant kelp Macrocystis pyrifera (L.) C.A. Agardh was removed at the holdfast from ≈ 1 ha of high relief structurally complex rock substratum. The abundance of seven species of fish, of which five were considered midwater species, significantly declined after the kelp was removed. No increases in abundance were noted within the experimental site. On the control site, two declines in abundance were observed, while two species increased in abundance. These results indicate that the presence of a giant kelp forest may increase the abundance and species diversity of the fish assemblages over a high relief rocky reef in central California, U.S.A.

Journal of Experimental Marine Biology and Ecology↗

Food availability controls seasonal cycle of growth in Macoma balthica (L.) in San Francisco Bay, California

A 2-yr field study of growth in the bivalve Macoma balthica (L.) at four locations in San Francisco Bay, California, U.S., showed that the timing and rate of growth (increase in shell length) were related to food supply. This clam feeds on both planktonic and benthic microalgae, depending on availability. Growth was apparently food-limited during some months, during one year more than the other, and at some locations more than others. Tissue-weight changes were also related to food availability. The heaviest animals were found in that year and at those locations with the highest chl a concentrations. Tissue-weight gains usually coincided with increased shell-growth rate or with reproductive development, although some large animals showed weight gain independent of both of these factors during periods with mid-range chl a levels. Weight losses coincided with spawning or periods of low growth rate, except at one station where, during a period when most animals were reproductively ripe, food concentrations were high, and shell growth was rapid, animals lost weight. This study failed to show a relation between salinity and the timing or rate of change of either shell length or tissue weight. The mild temperatures (10-23 ??C water temperature) of the area studied resulted in no growth inhibition due to low temperature, but there was some indication that the high air temperatures found in these intertidal areas limited growth rates. ?? 1988.

Journal of Experimental Marine Biology and Ecology↗

Effects of experimental overgrowth on survival and change in the turf assemblage of a giant kelp forest

Crustose coralline algae were the prevalent cover among sessile organisms that paved or grew near the substratum, and also the most commonly overgrown species in a giant kelp Macrocystis pyrifera (L.) C.A. Agardh forest located off San Nicolas Island, California. Giant kelp was the largest and most conspicuous species that overgrew large patches of the substrata; overgrowth among turf organisms also appeared common. To determine the effects of giant kelp holdfasts on crustose coralline algae and other turf organisms, “artificial holdfasts” were placed on 0.125-m 2 plots for 5, 8 and 12 months. In these treatments, 50–57% of the crustose coralline algae survived. Because these algae also recruited while covered, the total cover (survivorship plus recruitment) differed by only 7–26% from that sampled at the start of the study. The decline of these algae in control plots was similar to that in the treatment plots mostly because of overgrowth by sessile invertebrates. Bryozoans increased markedly on the control plots, whereas 0–12% survived in the treatment plots. Bryozoans and sponges also recruited under the artificial holdfasts. Some arborescent turf algae survived in the 5- and 8-month treatments; articulated coralline algae survived better than did foliose algae. High survival recruitment of crustose coralline algae while overgrown contributed to their prevalence in benthic communities.

Journal of Experimental Marine Biology and Ecology↗

Can otolith microchemistry chart patterns of migration and habitat utilization in anadromous fishes?

Seasonal and ontogenetic patterns in estuarine and coastal migrations of anadromous fish species have important consequences to their survival, growth, recruitment, and reproduction. We tested the hypothesis that otolith (sagitta) microchemistry can document the environmental history of individual fish across an estuarine salinity gradient. Juvenile striped bass, Morone saxatilis (Walbaum), (80 days posthatch) were reared for 3 wk in aquaria at two temperatures and six salinities. The ratio of strontium/calcium ( Sr Ca "> SrCa ) deposited in the sagittal otoliths of reared juveniles was positively related to salinity. Temperature and growth rate had relatively minor, but significant effects on the Sr Ca "> SrCa ratio. In a second experiment, juveniles (80 days posthatch) were exposed to increasing salinity (0 ppt to 25 ppt) and then decreasing salinity (25 ppt to 0 ppt) over a 20-wk period. Electron microprobe examination of the otoliths from these juveniles showed a gradual rise and decline in Sr Ca "> SrCa during the experimental period which corresponded directly with experimental changes in salinity. Field data on subadult and adult striped bass corroborated the laboratory analyses and indicated a logistic relationship between ambient salinity and otolith Sr Ca "> SrCa ratio. Verification studies support the use of otolith microchemistry to measure migratory schedules and habitat utilization patterns in anadromous striped bass populations.

Journal of Experimental Marine Biology and Ecology↗

Resilience of predators to fishing pressure on coral patch reefs

Numbers and biomass of piscivorous fish and their predation on other fish may often be high in undisturbed coral reef communities. The effects of such predation have sometimes been studied by removal of piscivores (either experimentally or by fishermen). Such perturbations have usually involved removal of large, highly vulnerable, mobile piscivores that are often actively sought in fisheries. The effects of fishing on smaller, demersal, semi-resident piscivores have been little studied. We studied such effects on the fish communities of patch reefs at Midway atoll by experimentally removing major resident, demersal, piscivorous fishes. First, four control reefs and four experimental reefs were selected, their dimensions and habitats mapped, and their visible fish communities censused repeatedly over 1 year. Census of all control and experimental reefs was continued for the following 39 months, during which known piscivores were collected repeatedly by hand spearing. Records were kept of catch and effort to calculate CPUE as an index of predator density. Spearfishing on the experimental reefs removed 2504 piscivorous fish from 12 families and 43 taxa (mostly species). The species richness of the catch did not show an overall change over the duration of the experiment. Spearman rank correlation analysis showed some unexpected positive correlations for density in numbers and biomass of major fished piscivorous groups (especially lizardfish) over the experiment. Only two relatively minor fished piscivorous taxa declined in abundance over the experiment, while the overall abundance of piscivores increased. Visual censuses of fish on the experimental reefs also failed to show reduction of total piscivores over the full experimental period. No significant trend in the abundance of lizardfish censused over the full period was apparent on any of the control reefs. The high resilience of piscivores on these experimental reefs to relatively intense fishing pressure could result from their protracted recruitment seasons, high immigration rates, cryptic habits, or naturally high abundances. A major factor was the high immigration rates of lizardfish, replacing lizardfish and other less mobile piscivores removed from the reefs by spearing. On the fished reefs, the removed lizardfish population replaced itself >20 times during the experiment; other piscivorous taxa replaced themselves only 5 times.

Journal of Experimental Marine Biology and Ecology↗

Biomechanical factors contributing to self-organization in seagrass landscapes

Field observations have revealed that when water flow is consistently from one direction, seagrass shoots align in rows perpendicular to the primary axis of flow direction. In this study, live Zostera marina shoots were arranged either randomly or in rows perpendicular to the flow direction and tested in a seawater flume under unidirectional flow and waves to determine if shoot arrangement: a) influenced flow-induced force on individual shoots, b) differentially altered water flow through the canopy, and c) influenced light interception by the canopy. In addition, blade breaking strength was compared with flow-induced force to determine if changes in shoot arrangement might reduce the potential for damage to shoots. Under unidirectional flow, both current velocity in the canopy and force on shoots were significantly decreased when shoots were arranged in rows as compared to randomly. However, force on shoots was nearly constant with downstream distance, arising from the trade-off of shoot bending and in-canopy flow reduction. The coefficient of drag was higher for randomly-arranged shoots at low velocities (< 30 cm s- 1) but converged rapidly among the two shoot arrangements at higher velocities. Shoots arranged in rows tended to intercept slightly more light than those arranged randomly. Effects of shoot arrangement under waves were less clear, potentially because we did not achieve the proper plant size?row spacing ratio. At this point, we may only suggest that water motion, as opposed to light capture, is the dominant physical mechanism responsible for these shoot arrangements. Following a computation of the Environmental Stress Factor, we concluded that even photosynthetically active blades may be damaged or broken under frequently encountered storm conditions, irrespective of shoot arrangement. We hypothesize that when flow is generally from one direction, seagrass bed patterns over multiple scales of consideration may arise as a cumulative effect of individual shoot self-organization driven by reduced force and drag on the shoots and somewhat improved light capture.

Journal of Experimental Marine Biology and Ecology↗

Ecological observations on the colonial ascidian Didemnum sp. in a New England tide pool habitat

The colonial ascidian Didemnum sp. has colonized northwestern Atlantic coastal habitats from southern Long Island, New York, to Eastport, Maine. It is also present in offshore habitats of the Georges Bank fishing grounds. It threatens to alter fisheries habitats and shellfish aquacultures. Observations in a tide pool at Sandwich, MA from December 2003 to February 2006 show that Didemnum sp. tolerates water temperatures ranging from ≤ 1 to > 24 °C, with daily changes of up to 11 °C. It attaches to pebbles, cobbles, and boulders, and it overgrows other tunicates, seaweeds, sponges, and bivalves. From May to mid July, colonies appear as small patches on the bottoms of rocks. Colonies grow rapidly from July to September, with some growth into December, and they range in color from pink to pale yellow to pale orange. Colony health declines from October through April, presumably in response to changes in water temperatures, and this degenerative process is manifested by color changes, by the appearance of small dark brown spots that represent clumps of fecal pellets in the colony, by scavenging by periwinkles, and by a peeling-away of colonies from the sides of cobbles and boulders. At Sandwich, colonies died that were exposed to air at low tide. The species does not exhibit this seasonal cycle of growth and decline in subtidal habitats (40–65 m) on the Georges Bank fishing grounds where the daily climate is relatively stable and annual water temperatures range from 4 to 15 °C. Experiments in the tide pool with small colony fragments (5 to 9 cm 2 ) show they re-attach and grow rapidly by asexual budding, increasing in size 6- to 11-fold in the first 15 days. Didemnum sp. at Sandwich has no known predators except for common periwinkles ( Littorina littorea ) that graze on degenerating colonies in the October to April time period and whenever colonies are stressed by desiccation. The tendencies of the ascidian (1) to attach to firm substrates, (2) to rapidly overgrow other species, (3) to tolerate a wide temperature range, (4) to be free from predation, and (5) to spread by colony fragmentation combine to make it a potential threat to benthic marine habitats and aquacultures. Didemnum sp. is known to overgrow mussels, oysters, and sea scallops, and it likely envelops other bivalves too.

Massachusetts↗

The occurrence of the colonial ascidian Didemnum sp. on Georges Bank gravel habitat: ecological observations and potential effects on groundfish and scallop fisheries

The colonial ascidian Didemnum sp. is present on the Georges Bank fishing grounds in a gravel habitat where the benthic invertebrate fauna has been monitored annually since 1994. The species was not noted before 2002 when large colonies were first observed; and by 2003 and 2004 it covered large areas of the seabed at some locations. The latest survey in 2005 documented the tunicate's presence in two gravel areas that total more than 67 nm 2 (230 km 2 ). The affected area is located on the Northern Edge of the bank in United States waters near the U.S./Canada boundary ( Fig. 1). This is the first documented offshore occurrence of a species that has colonized eastern U.S. coastal waters from New York to Maine during the past 15–20 years ( U.S. Geological Survey, 2006). Video imagery shows colonies coalescing to form large mats that cover more than 50% of the seabed along some video/photo transects. The affected area is an immobile pebble and cobble pavement that lies at water depths of 40 to 65 m where strong semidiurnal tidal currents reach speeds of 1 to 2 kt (50–100 cm/s). The water column is mixed year round, ensuring a constant supply of nutrients to the seabed. Annual temperatures range from 4 to 15 °C ( Mountain and Holzwarth, 1989). The gravel areas are bounded by sand ridges whose mobile surfaces are moved daily by the strong tidal currents. Studies commenced here in 1994 to characterize the gravel habitat and to document the effects of fishing disturbance on it ( Collie et al., 2005).

Massachusetts↗

Effects of intermittent flow and irradiance level on back reef Porites corals at elevated seawater temperatures

Corals inhabiting shallow back reef habitats are often simultaneously exposed to elevated seawater temperatures and high irradiance levels, conditions known to cause coral bleaching. Water flow in many tropical back reef systems is tidally influenced, resulting in semi-diurnal or diurnal flow patterns. Controlled experiments were conducted to test effects of semi-diurnally intermittent water flow on photoinhibition and bleaching of the corals Porites lobata and P. cylindrica kept at elevated seawater temperatures and different irradiance levels. All coral colonies were collected from a shallow back reef pool on Ofu Island, American Samoa. In the high irradiance experiments, photoinhibition and bleaching were less for both species in the intermittent high-low flow treatment than in the constant low flow treatment. In the low irradiance experiments, there were no differences in photoinhibition or bleaching for either species between the flow treatments, despite continuously elevated seawater temperatures. These results suggest that intermittent flow associated with semi-diurnal tides, and low irradiances caused by turbidity or shading, may reduce photoinhibition and bleaching of back reef corals during warming events. ?? 2006 Elsevier B.V. All rights reserved.

Journal of Experimental Marine Biology and Ecology↗

Biomedical and veterinary science can increase our understanding of coral disease

A balanced approach to coral disease investigation is critical for understanding the global decline of corals. Such an approach should involve the proper use of biomedical concepts, tools, and terminology to address confusion and promote clarity in the coral disease literature. Investigating disease in corals should follow a logical series of steps including identification of disease, systematic morphologic descriptions of lesions at the gross and cellular levels, measurement of health indices, and experiments to understand disease pathogenesis and the complex interactions between host, pathogen, and the environment. This model for disease investigation is widely accepted in the medical, veterinary and invertebrate pathology disciplines. We present standard biomedical rationale behind the detection, description, and naming of diseases and offer examples of the application of Koch's postulates to elucidate the etiology of some infectious diseases. Basic epidemiologic concepts are introduced to help investigators think systematically about the cause(s) of complex diseases. A major goal of disease investigation in corals and other organisms is to gather data that will enable the establishment of standardized case definitions to distinguish among diseases. Concepts and facts amassed from empirical studies over the centuries by medical and veterinary pathologists have standardized disease investigation and are invaluable to coral researchers because of the robust comparisons they enable; examples of these are given throughout this paper. Arguments over whether coral diseases are caused by primary versus opportunistic pathogens reflect the lack of data available to prove or refute such hypotheses and emphasize the need for coral disease investigations that focus on: characterizing the normal microbiota and physiology of the healthy host; defining ecological interactions within the microbial community associated with the host; and investigating host immunity, host-agent interactions, pathology, pathogenesis, and factors that promote the pathogenicity of the causative agent(s) of disease.

Journal of Experimental Marine Biology and Ecology↗

Molecular detection of native and invasive marine invertebrate larvae present in ballast and open water environmental samples collected in Puget Sound

Non-native marine species have been and continue to be introduced into Puget Sound via several vectors including ship's ballast water. Some non-native species become invasive and negatively impact native species or near shore habitats. We present a new methodology for the development and testing of taxon specific PCR primers designed to assess environmental samples of ocean water for the presence of native and non-native bivalves, crustaceans and algae. The intergenic spacer regions (IGS; ITS1, ITS2 and 5.8S) of the ribosomal DNA were sequenced for adult samples of each taxon studied. We used these data along with those available in Genbank to design taxon and group specific primers and tested their stringency against artificial populations of plasmid constructs containing the entire IGS region for each of the 25 taxa in our study, respectively. Taxon and group specific primer sets were then used to detect the presence or absence of native and non-native planktonic life-history stages (propagules) from environmental samples of ballast water and plankton tow net samples collected in Puget Sound. This methodology provides an inexpensive and efficient way to test the discriminatory ability of taxon specific oligonucleotides (PCR primers) before creating molecular probes or beacons for use in molecular ecological applications such as probe hybridizations or microarray analyses. This work addresses the current need to develop molecular tools capable of diagnosing the presence of planktonic life-history stages from non-native marine species (potential invaders) in ballast water and other environmental samples. ?? 2008 Elsevier B.V.

Journal of Experimental Marine Biology and Ecology↗

Derivation of habitat-specific dissolved oxygen criteria for Chesapeake Bay and its tidal tributaries

The Chesapeake 2000 Agreement committed its state and federal signatories to “define the water quality conditions necessary to protect aquatic living resources” in the Chesapeake Bay (USA) and its tidal tributaries. Hypoxia is one of the key water quality issues addressed as a result of the above Agreement. This paper summarizes the protection goals and specific criteria intended to achieve those goals for addressing hypoxia. The criteria take into account the variety of Bay habitats and the tendency towards low dissolved oxygen in some areas of the Bay. Stressful dissolved oxygen conditions were characterized for a diverse array of living resources of the Chesapeake Bay by different aquatic habitats: migratory fish spawning and nursery, shallow-water, open-water, deep-water, and deep-channel. The dissolved oxygen criteria derived for each of these habitats are intended to protect against adverse effects on survival, growth, reproduction and behavior. The criteria accommodate both spatial and temporal aspects of low oxygen events, and have been adopted into the Chesapeake Bay states – Maryland, Virginia, and Delaware – and the District of Columbia's water quality standards regulations. These criteria, now in the form of state regulatory standards, are driving an array of land-based and wastewater pollution reduction actions across the six-watershed.

Maryland, Virginia, Washington, D.C.↗

Fine scale daily movements and habitat use of East Pacific green turtles at a shallow coastal lagoon in Baja California Sur, Mexico

Green turtles spend most of their lives in coastal foraging areas where they face multiple anthropogenic impacts. Therefore, understanding their spatial use in this environment is a priority for conservation efforts. We studied the fine scale daily movements and habitat use of East Pacific green turtles ( Chelonia mydas ) at Laguna San Ignacio, a shallow coastal lagoon in Baja California Sur, Mexico where sea turtles are subject to high levels of gillnet bycatch and directed hunting. Six turtles ranging from 44.6 to 83.5 cm in straight carapace length were tracked for short deployments (1 to 6 d) with GPS-VHF telemetry. Turtles were active throughout diurnal, nocturnal, and crepuscular periods. Although they moved greater total distances during daytime, their speed of travel and net displacement remained consistent throughout 24-h periods. A positive selection for areas of seagrass and moderate water depth (5 to 10 m) was determined using Ivlev's electivity index, with neutral selection for shallow water (< 5 m) and avoidance of deep water (> 10 m). Turtles exhibited two distinct behavioral movement patterns: circular movements with high fidelity to the capture&ndash;release location and meandering movements with low fidelity to the capture&ndash;release location. Our results indicate that green turtles were active throughout the diel cycle while traveling large distances and traversing multiple habitats over short temporal scales.

Baja California Sur↗

Microhabitat associations of a semi-terrestrial fish, Kryptolebias marmoratus (Poey 1880) in a mosquito-ditched mangrove forest, west-central Florida

Mangrove rivulus ( Kryptolebias marmoratus ) is one of the few species of fish that is semi-terrestrial and able to use exposed intertidal and potentially supratidal habitats for prolonged periods of time. Based on previous work demonstrating frequent use of subterranean crab burrows as well as damp leaf litter and logs, we examined the microhabitat associations of rivulus in a mosquito-ditched mangrove forest on the Gulf coast of Florida near the northern limit of its distribution. We captured 161 rivulus on 20 dates between late April and mid-December 2007 using trench traps. Fish ranged in size from 7 to 35 mm SL. Peak abundance in mid-summer coincided with recruitment of a new year-class. The three study sites occurred within 0.5 km of one another, and experienced similar water temperatures and salinities. Nevertheless, they differed in their degree of tidal inundation, standing stock of leaf litter, and density of entrances to fiddler crab burrows. We consistently observed the highest mean catches of rivulus away from permanent subtidal waters of mosquito ditches, at intermediate relative elevations, and where leaf litter was locally abundant. Density of entrances to crab burrows was apparently unrelated to rivulus distribution or abundance in these forests.

Florida↗

Testing the effect of habitat structure and complexity on nekton assemblages using experimental oyster reefs

Structurally complex habitats are often associated with more diverse and abundant species assemblages in both aquatic and terrestrial ecosystems. Biogenic reefs formed by the eastern oyster ( Crassostrea virginica ) are complex in nature and are recognized for their potential habitat value in estuarine systems along the US Atlantic and Gulf of Mexico coasts. Few studies, however, have examined the response of nekton to structural complexity within oyster reefs. We used a quantitative sampling technique to examine how the presence and complexity of experimental oyster reefs influence the abundance, biomass, and distribution of nekton by sampling reefs 4 months and 16 months post-construction. Experimental oyster reefs were colonized immediately by resident fishes and decapod crustaceans, and reefs supported a distinct nekton assemblage compared to mud-bottom habitat. Neither increased reef complexity, nor age of the experimental reef resulted in further changes in nekton assemblages or increases in nekton abundance or diversity. The presence of oyster reefs per se was the most important factor determining nekton usage.

Journal of Experimental Marine Biology and Ecology↗

Experimentally derived salinity tolerance of hatchling Burmese pythons (Python molurus bivittatus) from the Everglades, Florida (USA)

In a laboratory setting, we tested the ability of 24 non-native, wild-caught hatchling Burmese pythons (Python molurus bivittatus) collected in the Florida Everglades to survive when given water containing salt to drink. After a one-month acclimation period in the laboratory, we grouped snakes into three treatments, giving them access to water that was fresh (salinity of 0, control), brackish (salinity of 10), or full-strength sea water (salinity of 35). Hatchlings survived about one month at the highest marine salinity and about five months at the brackish-water salinity; no control animals perished during the experiment. These results are indicative of a "worst-case scenario", as in the laboratory we denied access to alternate fresh-water sources that may be accessible in the wild (e.g., through rainfall). Therefore, our results may underestimate the potential of hatchling pythons to persist in saline habitats in the wild. Because of the effect of different salinity regimes on survival, predictions of ultimate geographic expansion by non-native Burmese pythons that consider salt water as barriers to dispersal for pythons may warrant re-evaluation, especially under global climate change and associated sea-level-rise scenarios.

Florida↗

Shading decreases the abundance of the herbivorous California horn snail, Cerithidea californica

Most of the intertidal zone in estuaries of California, USA and Baja California, Mexico is covered with vascular vegetation. Shading by these vascular plants influences abiotic and biotic processes that shape benthic community assemblages. We present data on the effects of shading on the California horn snail, Cerithidea californica . This species is important because it is the most common benthic macrofaunal species in these systems and acts as an obligate intermediate host of several species of rematode parasites that infect several other species. Using observational and experimental studies, we found a negative effect of shade on the distribution and abundance of the California horn snail. We hypothesized that shading reduces the abundance of the epipelic diatoms that the snails feeds on, causing snails to leave haded areas. We observed a negative relationship between vascular plant cover, sub-canopy light levels, and snail density in Mugu Lagoon. Then we experimentally manipulated light regimes, by clipping vegetation and adding shade structures, and found higher snail densities at higher light levels. In Goleta Slough, we isolated the effect of shade from vegetation by documenting a negative relationship between the shade created by two bridges and diatom and snail densities. We also found that snails moved the greatest distances over shaded channel banks compared to unshaded channel banks. Further, we documented the effect of water depth and channel bank orientation on shading in this system. An additional effect of shading is the reduction of temperature, providing an alternative explanation for some of our results. These results broaden our knowledge of how variation in the light environment influences the ecology of estuarine ecosystems.

California↗