Mills Susan W.

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Photographic identification guide to larvae at hydrothermal vents

(Woods Hole Oceanographic Institution, 2009-06) Mills, Susan W. ; Beaulieu, Stace E. ; Mullineaux, Lauren S.

The purpose of this guide is to assist researchers in the identification of larvae of benthic invertebrates at hydrothermal vents. Our work is based on plankton sampling at the East Pacific Rise 9-10°N vent field from 1991-2007, supplemented by benthic collections of juveniles. In addition to images and descriptions of the species, we included frequency data from large-volume plankton pump samples taken between 1998 and 2004 and time-series sediment trap samples from 2004-2005.
Detecting the influence of initial pioneers on succession at deep-sea vents

(Public Library of Science, 2012-12-04) Mullineaux, Lauren S. ; Le Bris, Nadine ; Mills, Susan W. ; Henri, Pauline ; Bayer, Skylar R. ; Secrist, Richard G. ; Siu, Nam

Deep-sea hydrothermal vents are subject to major disturbances that alter the physical and chemical environment and eradicate the resident faunal communities. Vent fields are isolated by uninhabitable deep seafloor, so recolonization via dispersal of planktonic larvae is critical for persistence of populations. We monitored colonization near 9°50′N on the East Pacific Rise following a catastrophic eruption in order to address questions of the relative contributions of pioneer colonists and environmental change to variation in species composition, and the role of pioneers at the disturbed site in altering community structure elsewhere in the region. Pioneer colonists included two gastropod species: Ctenopelta porifera, which was new to the vent field, and Lepetodrilus tevnianus, which had been rare before the eruption but persisted in high abundance afterward, delaying and possibly out-competing the ubiquitous pre-eruption congener L. elevatus. A decrease in abundance of C. porifera over time, and the arrival of later species, corresponded to a decrease in vent fluid flow and in the sulfide to temperature ratio. For some species these successional changes were likely due to habitat requirements, but other species persisted (L. tevnianus) or arrived (L. elevatus) in patterns unrelated to their habitat preferences. After two years, disturbed communities had started to resemble pre-eruption ones, but were lower in diversity. When compared to a prior (1991) eruption, the succession of foundation species (tubeworms and mussels) appeared to be delayed, even though habitat chemistry became similar to the pre-eruption state more quickly. Surprisingly, a nearby community that had not been disturbed by the eruption was invaded by the pioneers, possibly after they became established in the disturbed vents. These results indicate that the post-eruption arrival of species from remote locales had a strong and persistent effect on communities at both disturbed and undisturbed vents.
Active positioning of vent larvae at a mid-ocean ridge

( 2013-03) Mullineaux, Lauren S. ; McGillicuddy, Dennis J. ; Mills, Susan W. ; Kosnyrev, V. K. ; Thurnherr, Andreas M. ; Ledwell, James R. ; Lavelle, J. William

The vertical position of larvae of vent species above a mid-ocean ridge potentially has a strong effect on their dispersal. Larvae may be advected upward in the buoyant vent plume, or move as a consequence of their buoyancy or active swimming. Alternatively, they may be retained near bottom by the topography of the axial trough, or by downward swimming. At vents near 9°50’N on the axis of the East Pacific Rise, evidence for active larval positioning was detected in a comparison between field observations of larvae in the plankton in 2006 and 2007 and distributions of non-swimming larvae in a two-dimensional bio-physical model. In the field, few vent larvae were collected at the level of the neutrally buoyant plume (~75 m above bottom); their relative abundances at that height were much lower than those of simulated larvae from a near-bottom release in the model. This discrepancy was observed for many vent species, particularly gastropods, suggesting that they may actively remain near bottom by sinking or swimming downward. Near the seafloor, larval abundance decreased from the ridge axis to 1000 m off axis much more strongly in the observations than in the simulations, again pointing to behavior as a potential regulator of larval transport. We suspect that transport off axis was reduced by downward-moving behavior, which positioned larvae into locations where they were isolated from cross-ridge currents by seafloor topography, such as the walls of the axial valley – which are not resolved in the model. Cross-ridge gradients in larval abundance varied between gastropods and polychaetes, indicating that behavior may vary between taxonomic groups, and possibly between species. These results suggest that behaviorally mediated retention of vent larvae may be common, even for species that have a long planktonic larval duration and are capable of long-distance dispersal.
Vertical, lateral and temporal structure in larval distributions at hydrothermal vents

(Inter-Research, 2005-06-02) Mullineaux, Lauren S. ; Mills, Susan W. ; Sweetman, Andrew K. ; Beaudreau, A. H. ; Metaxas, Anna ; Hunt, H. L.

We examined larval abundance patterns near deep-sea hydrothermal vents along the East Pacific Rise to investigate how physical transport processes and larval behavior may interact to influence larval dispersal from, and supply to, vent populations. We characterized vertical and lateral distributions and temporal variation of larvae of vent species using high-volume pumps that recovered larvae in good condition (some still alive) and in high numbers (up to 450 individuals sample–1). Moorings supported pumps at heights of 1, 20, and 175 m above the seafloor, and were positioned directly above and at 10s to 100s of meters away from vent communities. Sampling was conducted on 4 cruises between November 1998 and May 2000. Larvae of 22 benthic species, including gastropods, a bivalve, polychaetes, and a crab, were identified unequivocally as vent species, and 15 additional species, or species-groups, comprised larvae of probable vent origin. For most taxa, abundances decreased significantly with increasing height above bottom. When vent sites within the confines of the axial valley were considered, larval abundances were significantly higher on-vent than off, suggesting that larvae may be retained within the valley. Abundances of all vent species varied significantly among sample dates; the variation was not synchronized among taxa, except for consistently low abundances during November 1998. Lateral distributions did not vary among major larval groups (gastropods, polychaetes and bivalves), although polychaetes showed anomalously high abundances off-vent at 1 m above bottom. Lateral patterns also did not vary among species of gastropods, indicating that hydrodynamic processes may be transporting diverse species in similar ways. However, the species-level differences in temporal patterns indicate that there is substantial discontinuity in the abundance of individual species at vent communities, possibly due to timing of spawning and/or behavioral interactions with flow.
Persistent effects of disturbance on larval patterns in the plankton after an eruption on the East Pacific Rise

( 2013-07) Mills, Susan W. ; Mullineaux, Lauren S. ; Beaulieu, Stace E. ; Adams, Diane K.

To predict how benthic communities will respond to disturbance, it is necessary to understand how disturbance affects the planktonic larval supply available to recolonize the area. Deep-sea hydrothermal vent fauna along the East Pacific Rise (EPR) experience frequent local extinctions due to tectonic and magmatic events, but the effects on regional larval abundance and diversity are unknown. We had been monitoring larvae at 9° 50' N on the EPR prior to the 2006 eruption and were able to resume collections shortly afterward. We found that many species that were common before the eruption became significantly less so afterward, whereas a few other species experienced a transient spike in abundance. Surprisingly, overall species richness in the plankton was high 9 mo after the eruption, but then decreased sharply after 1 yr and had not returned to pre-eruption levels after 2 yr. These results suggest that recovery from disturbance may continue to be affected by limited larval supply even several years after a disturbance event. This delay in recovery means that larvae of pioneer species may dominate potential colonists, even after benthic habitats have transitioned to conditions that favor later-successional species. Moreover, the combined effects of natural and anthropogenic disturbance (e.g. mining) would be likely to cause more profound and long-lasting changes than either event alone. Our results indicate that we do not have sufficient data to predict the timing of recovery after disturbance in the deep sea, even in a well-studied vent system.
Biogeochemical exploration of the Pescardero Basin vents

(The Oceanography Society, 2018-03) Michel, Anna P. M. ; Wankel, Scott D. ; Beaulieu, Stace E. ; Soule, Samuel A. ; Mullineaux, Lauren S. ; Coleman, Dwight ; Escobar Briones, Elva ; Gaytan-Caballero, Adriana ; McDermott, Jill M. ; Mills, Susan W. ; Speth, Dan ; Zierenberg, Robert
Prolonged recovery time after eruptive disturbance of a deep-sea hydrothermal vent community

(The Royal Society, 2020-12-23) Mullineaux, Lauren S. ; Mills, Susan W. ; Le Bris, Nadine ; Beaulieu, Stace E. ; Sievert, Stefan M. ; Dykman, Lauren

Deep-sea hydrothermal vents are associated with seafloor tectonic and magmatic activity, and the communities living there are subject to disturbance. Eruptions can be frequent and catastrophic, raising questions about how these communities persist and maintain regional biodiversity. Prior studies of frequently disturbed vents have led to suggestions that faunal recovery can occur within 2–4 years. We use an unprecedented long-term (11-year) series of colonization data following a catastrophic 2006 seafloor eruption on the East Pacific Rise to show that faunal successional changes continue beyond a decade following the disturbance. Species composition at nine months post-eruption was conspicuously different than the pre-eruption ‘baseline' state, which had been characterized in 1998 (85 months after disturbance by the previous 1991 eruption). By 96 months post-eruption, species composition was approaching the pre-eruption state, but continued to change up through to the end of our measurements at 135 months, indicating that the ‘baseline' state was not a climax community. The strong variation observed in species composition across environmental gradients and successional stages highlights the importance of long-term, distributed sampling in order to understand the consequences of disturbance for maintenance of a diverse regional species pool. This perspective is critical for characterizing the resilience of vent species to both natural disturbance and human impacts such as deep-sea mining.
Functional traits provide new insight into recovery and succession at deep-sea hydrothermal vents

(Ecological Society of America, 2021-05-28) Dykman, Lauren ; Beaulieu, Stace E. ; Mills, Susan W. ; Solow, Andrew R. ; Mullineaux, Lauren S.

Investigation of communities in extreme environments with unique conditions has the potential to broaden or challenge existing theory as to how biological communities assemble and change through succession. Deep-sea hydrothermal vent ecosystems have strong, parallel gradients of nutrients and environmental stress, and present unusual conditions in early succession, in that both nutrient availability and stressors are high. We analyzed the succession of the invertebrate community at 9°50′ N on the East Pacific Rise for 11 yr following an eruption in 2006 in order to test successional theories developed in other ecosystems. We focused on functional traits including body size, external protection, provision of habitat (foundation species), and trophic mode to understand how the unique nutritional and stress conditions influence community composition. In contrast to established theory, large, fast-growing, structure-forming organisms colonized rapidly at vents, while small, asexually reproducing organisms were not abundant until later in succession. Species in early succession had high external protection, as expected in the harsh thermal and chemical conditions after the eruption. Changes in traits related to feeding ecology and dispersal potential over succession agreed with expectations from other ecosystems. We also tracked functional diversity metrics over time to see how they compared to species diversity. While species diversity peaked at 8 yr post-eruption, functional diversity was continuing to increase at 11 yr. Our results indicate that deep-sea hydrothermal vents have distinct successional dynamics due to the high stress and high nutrient conditions in early succession. These findings highlight the importance of extending theory to new systems and considering function to allow comparison between ecosystems with different species and environmental conditions.
Habitat associations in gastropod species at East Pacific Rise hydrothermal vents (9°50'N)

(Marine Biological Laboratory, 2007-06) Mills, Susan W. ; Mullineaux, Lauren S. ; Tyler, Paul A.

At deep-sea hydrothermal vents on the East Pacific Rise (9°50'N), distinct megafaunal assemblages are positioned along strong thermal and chemical gradients. We investigated the distribution of gastropod species to determine whether they associate with specific megafaunal zones and to determine the thermal boundaries of their habitats. Gastropods colonized a series of basalt blocks that were placed into three different zones characterized by vestimentiferan tubeworms, bivalves, and suspension-feeders, respectively. Additional gastropods were collected on selected blocks from higher temperature vestimentiferan habitat and from grab samples of alvinellid polychaetes. On the blocks, gastropod species clustered into a "Cool" group (Clypeosectus delectus, Eulepetopsis vitrea, Gorgoleptis spiralis, and Lepetodrilus ovalis) whose species tended to be most abundant in the suspension-feeder zone, and a "Warm" group (Lepetodrilus cristatus, L. elevatus, L. pustulosus, and Cyathermia naticoides) whose species all were significantly more abundant in the vestimentiferan zone than elsewhere. The temperature ranges of Cool species were generally lower than the ranges of Warm ones, although both groups were present at 3 to 6 °C; also present was Bathymargarites symplector, which clustered with neither group. Three additional species, Rhynchopelta concentrica, Neomphalus fretterae, and Nodopelta rigneae, co-occurred with Warm-group species on selected blocks from hotter habitats. Although a few species were found only in alvinellid collections, most species were not exclusive to a specific megafaunal zone. We propose that species in the Cool and Warm groups occupy specific microhabitats that are present in more than one zone.
Expanding dispersal studies at hydrothermal vents through species identification of cryptic larval forms

( 2010-01) Adams, Diane K. ; Mills, Susan W. ; Shank, Timothy M. ; Mullineaux, Lauren S.

The rapid identification of hydrothermal vent-endemic larvae to the species level is a key limitation to understanding the dynamic processes that control the abundance and distribution of fauna in such a patchy and ephemeral environment. Many larval forms collected near vents, even those in groups such as gastropods that often form a morphologically distinct larval shell, have not been identified to species. We present a staged approach that combines morphological and molecular identification to optimize the capability, efficiency, and economy of identifying vent gastropod larvae from the northern East Pacific Rise (NEPR). With this approach, 15 new larval forms can be identified to species. A total of 33 of the 41 gastropod species inhabiting the NEPR, and 26 of the 27 gastropod species known to occur specifically in the 9° 50’ N region, can be identified to species. Morphological identification efforts are improved by new protoconch descriptions for Gorgoleptis spiralis, Lepetodrilus pustulosus, Nodopelta subnoda, and Echinopelta fistulosa. Even with these new morphological descriptions, the majority of lepetodrilids and peltospirids require molecular identification. Restriction fragment length polymorphism digests are presented as an economical method for identification of five species of Lepetodrilus and six species of peltospirids. The remaining unidentifiable specimens can be assigned to species by comparison to an expanded database of 18S ribosomal DNA. The broad utility of the staged approach was exemplified by the revelation of species-level variation in daily planktonic samples and the identification and characterization of egg capsules belonging to a conid gastropod Gymnobela sp. A. The improved molecular and morphological capabilities nearly double the number of species amenable to field studies of dispersal and population connectivity.
Reproduction of gastropods from vents on the East Pacific Rise and the Mid-Atlantic Ridge

(National Shellfisheries Association, 2008-03) Tyler, Paul A. ; Pendlebury, Sophie ; Mills, Susan W. ; Mullineaux, Lauren S. ; Eckelbarger, Kevin J. ; Baker, Maria C. ; Young, Craig M.

The gametogenic biology is described for seven species of gastropod from hydrothermal vents in the East Pacific and from the Mid-Atlantic Ridge. Species of the limpet genus Lepetodrilus (Family Lepetodrilidae) had a maximum unfertilized oocyte size of <90 μm and there was no evidence of reproductive periodicity or spatial variation in reproductive pattern. Individuals showed early maturity with females undergoing gametogenesis at less than one third maximum body size. There was a power relationship between shell length and fecundity, with a maximum of 1,800 oocytes being found in one individual, although individual fecundity was usually <1,000. Such an egg size might be indicative of planktotrophic larval development, but there was never any indication of shell growth in larvae from species in this genus. Cyathermia naticoides (Family Neomphalidea) had a maximum oocyte size of 120 μm and a fecundity of <400 oocytes per individual. Rhynchopelta concentrica (Family Peltospiridae) had a maximum oocyte size of 184 μm and a fecundity <600, whereas in Eulepetopsis vitrea (Family Neolepetopsidae) maximum oocyte size was 232 μm with a fecundity of <200 oocytes per individual. In none of these three species was there any indication of episodicity in oocyte production. From our observations we support the paradigm that there is no reproductive pattern typical of vent systems but is more related to species' phylogeny.
Larvae from afar colonize deep-sea hydrothermal vents after a catastrophic eruption

( 2010-01-29) Mullineaux, Lauren S. ; Adams, Diane K. ; Mills, Susan W. ; Beaulieu, Stace E.

The planktonic larval stage is a critical component of life history in marine benthic species because it confers the ability to disperse, potentially connecting remote populations and leading to colonization of new sites. Larval-mediated connectivity is particularly intriguing in deep-sea hydrothermal vent communities, where the habitat is patchy, transient and often separated by tens or hundreds of kilometers. A recent catastrophic eruption at vents near 9°50’N on the East Pacific Rise created a natural clearance experiment and provided an opportunity to study larval supply in the absence of local source populations. Previous field observations have suggested that established vent populations may retain larvae and be largely self-sustaining. If this hypothesis is correct, the removal of local populations should result in a dramatic change in the flux, and possibly species composition, of settling larvae. Fortuitously, monitoring of larval supply and colonization at the site had been established before the eruption and resumed shortly afterward. We detected a striking change in species composition of larvae and colonists after the eruption, most notably the appearance of the gastropod Ctenopelta porifera, an immigrant from possibly >300 km away, and the disappearance of a suite of species that formerly had been prominent. This switch demonstrates that larval supply can change markedly after removal of local source populations, enabling recolonization via immigrants from distant sites with different species composition. Population connectivity at this site appears to be temporally variable, depending not only on stochasticity in larval supply, but also on the presence of resident populations.
Dispersal of hydrothermal vent larvae at East Pacific Rise 9-10 degrees N segment

(American Geophysical Union, 2018-11-06) Xu, Guangyu ; McGillicuddy, Dennis J. ; Mills, Susan W. ; Mullineaux, Lauren S.

A three‐dimensional, primitive‐equation, ocean circulation model coupled with a Lagrangian particle‐tracking algorithm is used to investigate the dispersal and settlement of planktonic larvae released from discrete hydrothermal habitats on the East Pacific Rise segment at 9–10°N. Model outputs show that mean circulation is anticyclonic around the ridge segment, which consists of a northward flow along the western flank and a southward flow along the eastern flank. Those flank jets are dispersal expressways for the along‐ridge larval transport and strongly affect its overall direction and spatial‐temporal variations. It is evident from model results that the transform faults bounding the ridge segment and off axis topography (the Lamont Seamount Chain) act as topographic barriers to larval dispersal in the along‐ridge direction. Furthermore, the presence of an overlapping spreading center and an adjacent local topographic high impedes the southward along‐ridge larval transport. The model results suggest that larval recolonization within ridge‐crest habitats is enhanced by the anticyclonic circulation around the ridge segment, and the overall recolonization rate is higher for larvae having a short precompetency period and an altitude above the bottom sufficient to avoid influence by the near‐bottom currents Surprisingly, for larvae having a long precompetency period (>10 days), the prolonged travel time allowed some of those larvae to return to their natal vent clusters, which results in an unexpected increase in connectivity among natal and neighboring sites. Overall, model‐based predictions of connectivity are highly sensitive to the larval precompetency period and vertical position in the water column.
Larvae of deep-sea invertebrates harbor low-diversity bacterial communities

(University of Chicago Press, 2021-07-28) Carrier, Tyler ; Beaulieu, Stace E. ; Mills, Susan W. ; Mullineaux, Lauren S. ; Reitzel, Adam M.

Microbial symbionts are a common life-history character of marine invertebrates and their developmental stages. Communities of bacteria that associate with the eggs, embryos, and larvae of coastal marine invertebrates tend to be species specific and correlate with aspects of host biology and ecology. The richness of bacteria associated with the developmental stages of coastal marine invertebrates spans four orders of magnitude, from single mutualists to thousands of unique taxa. This understanding stems predominately from the developmental stages of coastal species. If they are broadly representative of marine invertebrates, then we may expect deep-sea species to associate with bacterial communities that are similar in diversity. To test this, we used amplicon sequencing to profile the bacterial communities of invertebrate larvae from multiple taxonomic groups (annelids, molluscs, crustaceans) collected from 2500 to 3670 m in depth in near-bottom waters near hydrothermal vents in 3 different regions of the Pacific Ocean (the East Pacific Rise, the Mariana Back-Arc, and the Pescadero Basin). We find that larvae of deep-sea invertebrates associate with low-diversity bacterial communities (~30 bacterial taxa) that lack specificity between taxonomic groups. The diversity of these communities is estimated to be ~7.9 times lower than that of coastal invertebrate larvae, but this result depends on the taxonomic group. Associating with a low-diversity community may imply that deep-sea invertebrate larvae do not have a strong reliance on a microbiome and that the hypothesized lack of symbiotic contributions would differ from expectations for larvae of coastal marine invertebrates.
sFDvent: a global trait database for deep-sea hydrothermal-vent fauna

(Wiley, 2019-07-30) Chapman, Abbie S. A. ; Beaulieu, Stace E. ; Colaço, Ana ; Gebruk, Andrey V. ; Hilario, Ana ; Kihara, Terue C. ; Ramirez-Llodra, Eva ; Sarrazin, Jozée ; Tunnicliffe, Verena ; Amon, Diva ; Baker, Maria C. ; Boschen‐Rose, Rachel E. ; Chen, Chong ; Cooper, Isabelle J. ; Copley, Jonathan T. ; Corbari, Laure ; Cordes, Erik E. ; Cuvelier, Daphne ; Duperron, Sébastien ; Du Preez, Cherisse ; Gollner, Sabine ; Horton, Tammy ; Hourdez, Stephane ; Krylova, Elena M. ; Linse, Katrin ; LokaBharathi, P. A. ; Marsh, Leigh ; Matabos, Marjolaine ; Mills, Susan W. ; Mullineaux, Lauren S. ; Rapp, Hans Tore ; Reid, William D. K. ; Rybakova, Elena Goroslavskaya ; Thomas, Tresa Remya A. ; Southgate, Samuel James ; Stöhr, Sabine ; Turner, Phillip J. ; Watanabe, Hiromi K. ; Yasuhara, Moriaki ; Bates, Amanda E.

Motivation Traits are increasingly being used to quantify global biodiversity patterns, with trait databases growing in size and number, across diverse taxa. Despite growing interest in a trait‐based approach to the biodiversity of the deep sea, where the impacts of human activities (including seabed mining) accelerate, there is no single repository for species traits for deep‐sea chemosynthesis‐based ecosystems, including hydrothermal vents. Using an international, collaborative approach, we have compiled the first global‐scale trait database for deep‐sea hydrothermal‐vent fauna – sFDvent (sDiv‐funded trait database for the Functional Diversity of vents). We formed a funded working group to select traits appropriate to: (a) capture the performance of vent species and their influence on ecosystem processes, and (b) compare trait‐based diversity in different ecosystems. Forty contributors, representing expertise across most known hydrothermal‐vent systems and taxa, scored species traits using online collaborative tools and shared workspaces. Here, we characterise the sFDvent database, describe our approach, and evaluate its scope. Finally, we compare the sFDvent database to similar databases from shallow‐marine and terrestrial ecosystems to highlight how the sFDvent database can inform cross‐ecosystem comparisons. We also make the sFDvent database publicly available online by assigning a persistent, unique DOI. Main types of variable contained Six hundred and forty‐six vent species names, associated location information (33 regions), and scores for 13 traits (in categories: community structure, generalist/specialist, geographic distribution, habitat use, life history, mobility, species associations, symbiont, and trophic structure). Contributor IDs, certainty scores, and references are also provided. Spatial location and grain Global coverage (grain size: ocean basin), spanning eight ocean basins, including vents on 12 mid‐ocean ridges and 6 back‐arc spreading centres. Time period and grain sFDvent includes information on deep‐sea vent species, and associated taxonomic updates, since they were first discovered in 1977. Time is not recorded. The database will be updated every 5 years. Major taxa and level of measurement Deep‐sea hydrothermal‐vent fauna with species‐level identification present or in progress. Software format .csv and MS Excel (.xlsx).
Animal community dynamics at senescent and active vents at the 9° N East Pacific Rise after a volcanic eruption

(Frontiers Media, 2020-01-24) Gollner, Sabine ; Govenar, Breea ; Arbizu, P. Martinez ; Mullineaux, Lauren S. ; Mills, Susan W. ; Le Bris, Nadine ; Weinbauer, Markus ; Shank, Timothy M. ; Bright, Monika

In 2005/2006, a major volcanic eruption buried faunal communities over a large area of the 9°N East Pacific Rise (EPR) vent field. In late 2006, we initiated colonization studies at several types of post eruption vent communities including those that either survived the eruption, re-established after the eruption, or arisen at new sites. Some of these vents were active whereas others appeared senescent. Although the spatial scale of non-paved (surviving) vent communities was small (several m2 compared to several km2 of total paved area), the remnant individuals at surviving active and senescent vent sites may be important for recolonization. A total of 46 meio- and macrofauna species were encountered at non-paved areas with 33 of those species detected were also present at new sites in 2006. The animals living at non-paved areas represent refuge populations that could act as source populations for new vent sites directly after disturbance. Remnants may be especially important for the meiofauna, where many taxa have limited or no larval dispersal. Meiofauna may reach new vent sites predominantly via migration from local refuge areas, where a reproductive and abundant meiofauna is thriving. These findings are important to consider in any potential future deep-sea mining scenario at deep-sea hydrothermal vents. Within our 4-year study period, we regularly observed vent habitats with tubeworm assemblages that became senescent and died, as vent fluid emissions locally stopped at patches within active vent sites. Senescent vents harbored a species rich mix of typical vent species as well as rare yet undescribed species. The senescent vents contributed significantly to diversity at the 9°N EPR with 55 macrofaunal species (11 singletons) and 74 meiofaunal species (19 singletons). Of these 129 species associated with senescent vents, 60 have not been reported from active vents. Tubeworms and other vent megafauna not only act as foundation species when alive but provide habitat also when dead, sustaining abundant and diverse small sized fauna.
Macrofauna and larvae collected at the Auka hydrothermal vent field in Pescadero Basin in 2017

(Woods Hole Oceanographic Institution, 2023-01-13) Beaulieu, Stace E. ; Mills, Susan W. ; Fleming, Bethany F. M. ; Angier, Sabine ; Toner, Mary ; Mullineaux, Lauren S.

This data package provides the sampling locations and identifications for macrofauna and larvae collected at the Auka hydrothermal vent field in Pescadero Basin in 2017 and used in a study by Fleming et al. (2022). This data package contains five tables: paired tables for benthic slurps (sampling metadata and specimen counts), paired tables for plankton slurps (sampling metadata and specimen counts), and one table summarizing benthic and plankton specimens with Barcode of Life Data System (BOLD) Barcode Index Numbers (BINs). The paired data tables are partially aligned to Darwin Core event and occurrence tables for future contribution to the Ocean Biodiversity Information System (OBIS). Records for specimens in BOLD are available through the Global Biodiversity Information Facility (GBIF).
Ecological connectivity in Pacific deep-sea hydrothermal vent metacommunities

(Inter-Research Science Publisher, 2024-03-13) Fleming, Bethany F. M. ; Beaulieu, Stace E. ; Mills, Susan W. ; Gaggiotti, Oscar E. ; Mullineaux, Lauren S.

Larval dispersal and connectivity between patchy, transient, deep-sea hydrothermal vent communities are important for persistence and recovery from disturbance. We investigated connectivity in vent metacommunities using the taxonomic similarity between larvae and adults to estimate the extent of exchange between communities and determine the relative roles of larval dispersal and environmental limitations (species sorting) in colonization. Connectivity at vent fields in 3 Pacific regions, Pescadero Basin, northern East Pacific Rise (EPR), and southern Mariana Trough, varied substantially and appeared to be driven by different processes. At Pescadero Basin, larval and adult taxa were similar, despite the existence of nearby (within 75 km) vent communities with different species composition, indicating limited larval transport and low connectivity. At EPR, larval and adult taxa differed significantly, despite the proximity of nearby vents with similar benthic composition, indicating substantial larval transport and potentially strong species sorting, but other factors may also explain these results. At the Mariana Trough, the larvae and adults differed significantly, indicating high larval transport but environmental limitations on colonization. We demonstrate that analysis of routinely collected samples and observations provides an informative indicator of metacommunity connectivity and insights into drivers of community assembly.
Faunal colonists, including mussel settlers, respond to microbial biofilms at deep-sea hydrothermal vents

(Elsevier, 2024-04-29) Ladd, Tanika M. ; Selci, Matteo ; Davis, Dexter J. ; Cannon, Olivia ; Plowman, Caitlin Q. ; Schlegel, Ian ; Inaba, Aila ; Mills, Susan Wier ; Vetriani, Costantino ; Mullineaux, Lauren S. ; Arellano, Shawn M.

Colonization processes at dynamic deep-sea hydrothermal vent ecosystems ultimately determine ecosystem structure, function, resilience, and recovery. Microbial biofilms form rapidly on surfaces near hydrothermal vents and are continuously exposed to the highly variable abiotic environment. Thus, biofilm microbes may provide a temporally integrated signal that can indicate whether the habitat is suitable for faunal colonists. This study explored the role of microbial biofilms in controlling faunal colonization through in-situ colonization experiments at Tica Vent in the 9°50’ N region of the East Pacific Rise (EPR). Short-term experiments (∼2 weeks) were conducted by deploying colonization surfaces (“sandwiches”) either with an established biofilm (developed for >1 year) or a fresh biofilm (developed throughout experiment) in zones characterized by different faunal assemblages. Differences in associated larval settlers, faunal immigrants, and microbial communities according to biofilm age across multiple biogenic zones were investigated. Faunal and microbial community compositions significantly differed according to whether the sandwiches had established or fresh biofilms as well as the biogenic zone they were deployed in. Several faunal colonists, including settlers such as the foundational chemosymbiotic mussel Bathymodiolus thermophilus and the nectochaete Archinome sp., were found associated more with established biofilms than fresh biofilms. Microbial biofilm communities were dominated by putative chemoautotrophic members of the Campylobacterota phylum and Gammaproteobacteria class and several microbial taxa were found to covary with faunal colonists. Overall, these findings show that microbial community composition plays a role in larval settlement and animal migration in hydrothermal vent systems and the detection of microbial and faunal interactions provides a starting point for identifying key microbial characteristics influencing colonization processes at hydrothermal vents.
Vertical distributions of megafauna on inactive vent sulfide features

(Elsevier, 2024-07-16) Meneses, Michael J. ; Beaulieu, Stace E. ; Best, Ayinde C. ; Dykman, Lauren N. ; Mills, Susan Wier ; Wu, Jyun-Nai ; Mullineaux, Lauren S.

The discovery of inactive hydrothermal vent sulfide features near 9°50′N on the East Pacific Rise provides an opportunity to investigate the distribution and feeding ecology of communities inhabiting this type of habitat. We quantify megafaunal distributions on two features, Lucky's Mound and Sentry Spire, to determine how taxonomic composition and feeding traits vary with vertical elevation. Fifty-one morphotypes, categorized by feeding mode, were identified from three levels of the features (spire, apron, and base) and the surrounding flat oceanic rise. About half of the morphotypes (26 of 51) were only observed at the sulfide features. Passive suspension feeders were more abundant on the spires, where horizontal particulate flux is expected to be elevated, than the base or rise. Deposit feeders tended to be more abundant on the base and rise, where deposition is expected to be enhanced, but were unexpectedly abundant higher up on Sentry Spire. Community differences between the two sulfide features suggest that other processes, such as feature-specific chemoautotrophic production, may also influence distributions.

Mills Susan W.

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