The R/V Oceanus completed a 9,789 km, 28 day passage from Woods Hole, Massachusetts, in the Atlantic Ocean, through the Panama Canal to Yaquina Bay, Oregon, in the Pacific Ocean on 21 February 2012. The Oceanus had previously operated in the Mediterranean Sea and Atlantic Ocean (including the Caribbean Sea). We document the sequential acquisition of the barnacles Balanus trigonus and Amphibalanus venustus and the oyster Ostrea equestris on the Oceanus on its high and low latitude transoceanic, intra-oceanic, and interoceanic travels before she was surveyed in Yaquina Bay. The close correspondence between hull fouling accumulations and the detailed two year Oceanus working history reveals B. trigonus settlement occurred in every tropical port visited by the Oceanus, that some populations survived through two of three Woods Hole winters, and that some of these populations passed through the freshwater Panama Canal. These results suggest that marine hull-fouling species are continuously transported globally between most ports of call by most ship passages.
Phylogenetic relationships and the timing of evolutionary events are essential for understanding evolution on longer time scales. Cheilostome bryozoans are a group of ubiquitous, species-rich, marine colonial organisms with an excellent fossil record but lack phylogenetic relationships inferred from molecular data. We present genome-skimmed data for 395 cheilostomes and combine these with 315 published sequences to infer relationships and the timing of key events among c. 500 cheilostome species. We find that named cheilostome genera and species are phylogenetically coherent, rendering fossil or contemporary specimens readily delimited using only skeletal morphology. Our phylogeny shows that parental care in the form of brooding evolved several times independently but was never lost in cheilostomes. Our fossil calibration, robust to varied assumptions, indicates that the cheilostome lineage and parental care therein could have Paleozoic origins, much older than the first known fossil record of cheilostomes in the Late Jurassic.
Shipping has contributed strongly to biological invasions in coastal ecosystems, transferring species in ballast tanks and on exposed underwater surfaces (hulls). A long history exists that documents biota associated with ships’ hulls, including some recent analyses of modern ships, but relatively little is known about the associated risks of invasion. In general, the likelihood of invasion is expected to increase with increasing propagule supply, which suggests that high‐density transfers on hulls may pose a relatively high invasion risk. Obsolete vessels are expected to be at an extreme end of the spectrum for biofouling, since they sit at anchorage for long periods and are towed at relatively slow speeds when moved, but this remains largely unexplored. In this paper, we quantified the biofouling communities of two obsolete vessels, one stationary for one decade and the other for two decades, before and after their final transit from California to Texas. Pre‐departure biofouling surveys across both vessels detected 22 species of macroinvertebrates. The biomass was dominated by the introduced bryozoan Conopeum chesapeakensis, which occurred in 98% of samples and created a three‐dimensional structure (2–5 cm thick). Mobile species, inhabiting the vertical biofouling matrix, were more numerous than sessile ones. Interestingly, the non‐native Asian clam Corbula amurensis, not previously associated with hull fouling assemblages, was recorded in 9% of samples. During the 43‐day voyage, organisms encountered salinity variation that ranged between zero (Panama Canal) and at least 37 parts per thousand (Brownsville, Texas) and temperatures that varied between 9.9 °C and 31.6 °C. Upon arrival in Texas, we measured an expected decrease in biofouling extent across both vessels but also a surprising increase in species richness (57 species were recorded), with small compositional differences between ships that did not exist prior to departure. Several species were recorded alive upon arrival, including non‐natives that are not known to be established in Texas waters. The physiological tolerance and associated risk of colonization have not yet been evaluated for these organisms, or for the broader species pool associated with a standing fleet (n > 200 ships) that may undergo similar movements. Nonetheless, a compelling case exists for vector management based on organism flux alone, to reduce the risk of coastwise and inter‐oceanic invasions.
Global trade by merchant ships is a leading mechanism for the unintentional transfer of marine organisms, including non-indigenous species, to bays and estuaries worldwide. To reduce the likelihood of new invasions, ships are increasingly being required to manage their ballast water (BW) prior to discharge in coastal waters. In the United States, most overseas arrivals have been required to manage BW discharge since 2004, primarily through ballast water exchange (BWE), which flushes out ballast tanks in the open ocean (>200 miles from shore). Studies have found BWE to generally reduce the abundance of organisms, and the amount of water exchanged has been estimated at 96–100%. Despite its widespread use, the overall effect of this management strategy on net propagule supply through time has not been explored. Here, temporal changes in zooplankton concentrations and the volume of BW discharged in Chesapeake Bay, U.S. were evaluated, comparing pre-management era and post-management era time periods. Chesapeake Bay is a large port system that receives extensive BW discharge, especially from bulk cargo vessels (bulkers) that export coal overseas. For bulkers arriving from overseas, mean zooplankton concentrations of total and coastal indicator taxa in BW did not decline between pre- (1993–2000) and post management (2012–2013) eras, when controlling for season and sampling method. Moreover, bulkers discharged 21 million tonnes (82% of total for Chesapeake Bay) of overseas BW in 2013, representing a 374% increase in volume when compared to 2005. The combination of BW discharge volume and zooplankton concentration data indicates that (a) net propagule supply by bulkers has increased since BWE began in Chesapeake Bay; and (b) changes in vessel behaviour and trade have contributed strongly to this outcome. Specifically, the coal-driven increase in BW discharge volume from 2005–2013, concurrent with the onset of BWE regulations, worked to counteract intended results from BW management. A long-term analysis of bulker arrivals (1994–2013) reveals a 20-year minimum in arrival numbers in 2000, just when the implementation of BWE began. This study underscores the need to consider shifts in trade patterns, in order to advance and evaluate effective management strategies for biological invasions.
As biological invasions continue to increase globally, eradication programs have been undertaken at significant cost, often without consideration of relevant ecological theory. Theoretical fisheries models have shown that harvest can actually increase the equilibrium size of a population, and uncontrolled studies and anecdotal reports have documented population increases in response to invasive species removal (akin to fisheries harvest). Both findings may be driven by high levels of juvenile survival associated with low adult abundance, often referred to as overcompensation. Here we show that in a coastal marine ecosystem, an eradication program resulted in stage-specific overcompensation and a 30-fold, single-year increase in the population of an introduced predator. Data collected concurrently from four adjacent regional bays without eradication efforts showed no similar population increase, indicating a local and not a regional increase. Specifically, the eradication program had inadvertently reduced the control of recruitment by adults via cannibalism, thereby facilitating the population explosion. Mesocosm experiments confirmed that adult cannibalism of recruits was size-dependent and could control recruitment. Genomic data show substantial isolation of this population and implicate internal population dynamics for the increase, rather than recruitment from other locations. More broadly, this controlled experimental demonstration of stage-specific overcompensation in an aquatic system provides an important cautionary message for eradication efforts of species with limited connectivity and similar life histories.
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