Broadband backscattering from scyphozoan jellyfish.

As the ecological importance of gelatinous organisms becomes increasingly appreciated, so has the need for improved knowledge of their abundance and distribution. Acoustic backscattering measurements are routine for fisheries assessments but are not yet widely used to survey populations of gelatinous zooplankton. The use of acoustic backscattering techniques to understand the distribution and abundance of organisms requires an understanding of their target strength (TS). This study presents a framework for a sound scattering model for jellyfish based on the Distorted Wave Born Approximation that incorporates size, shape, and material properties of individual organisms. This model, with a full three-dimensional shape rendition, is applied to a common species of scyphomedusa (Chrysaora chesapeakei) and verified experimentally with broadband (52-90 and 93-161 kHz) laboratory TS measurements of live individuals. Cyclical changes in the organism's shape due to swimming kinematics were examined, as well as averages over swimming position and comparisons with scattering from simpler shapes. The model predicts overall backscattering levels and broad spectral behavior within <2 dB. Measured TS exhibits greater variability than is predicted by scaling the size of the organism in the scattering model, showing that density and sound speed vary among individuals.

Modeling characterization of the vertical and temporal variability of environmental DNA in the mesopelagic ocean.

Increasingly, researchers are using innovative methods to census marine life, including identification of environmental DNA (eDNA) left behind by organisms in the water column. However, little is understood about how eDNA is distributed in the ocean, given that organisms are mobile and that physical and biological processes can transport eDNA after release from a host. Particularly in the vast mesopelagic ocean where many species vertically migrate hundreds of meters diurnally, it is important to link the location at which eDNA was shed by a host organism to the location at which eDNA was collected in a water sample. Here, we present a one-dimensional mechanistic model to simulate the eDNA vertical distribution after its release and to compare the impact of key biological and physical parameters on the eDNA vertical and temporal distribution. The modeled vertical eDNA profiles allow us to quantify spatial and temporal variability in eDNA concentration and to identify the most important parameters to consider when interpreting eDNA signals. We find that the vertical displacement by advection, dispersion, and settling has limited influence on the eDNA distribution, and the depth at which eDNA is found is generally within tens of meters of the depth at which the eDNA was originally shed from the organism. Thus, using information about representative vertical migration patterns, eDNA concentration variability can be used to answer ecological questions about migrating organisms such as what depths species can be found in the daytime and nighttime and what percentage of individuals within a species diurnally migrate. These findings are critical both to advance the understanding of the vertical distribution of eDNA in the water column and to link eDNA detection to organism presence in the mesopelagic ocean as well as other aquatic environments.

A hybrid underwater robot for multidisciplinary investigation of the ocean twilight zone.

Mesobot, an autonomous underwater vehicle, addresses specific unmet needs for observing and sampling a variety of phenomena in the ocean's midwaters. The midwater hosts a vast biomass, has a role in regulating climate, and may soon be exploited commercially, yet our scientific understanding of it is incomplete. Mesobot has the ability to survey and track slow-moving animals and to correlate the animals' movements with critical environmental measurements. Mesobot will complement existing oceanographic assets such as towed, remotely operated, and autonomous vehicles; shipboard acoustic sensors; and net tows. Its potential to perform behavioral studies unobtrusively over long periods with substantial autonomy provides a capability that is not presently available to midwater researchers. The 250-kilogram marine robot can be teleoperated through a lightweight fiber optic tether and can also operate untethered with full autonomy while minimizing environmental disturbance. We present recent results illustrating the vehicle's ability to automatically track free-swimming hydromedusae (Solmissus sp.) and larvaceans (Bathochordaeus stygius) at depths of 200 meters in Monterey Bay, USA. In addition to these tracking missions, the vehicle can execute preprogrammed missions collecting image and sensor data while also carrying substantial auxiliary payloads such as cameras, sonars, and samplers.

Diversity and distribution of nearshore barnacle cyprids in southern California through the 2015-16 El Niño.

Abundance, species diversity, and horizontal distributions of barnacle cyprids offshore of La Jolla, southern California were described from May 2014 to August 2016 to determine how the nearshore barnacle larval assemblage changed before, during, and after the 2015-16 El Niño. The entire water column was sampled at five stations located within one km of shore with water depths of 4, 6, 8, 10, and 12 m during 33 cruises that encompassed the time when El Niño conditions impacted the area. Nearshore temperature and thermal stratification was concurrently measured using a CTD. Six identified cyprid species, including Chthamalus fissus, Pollicipes polymerus, Megabalanus rosa, Tetraclita rubescens, Balanus glandula, and B. trigonus, along with four unknown species, were collected in our samples. DNA barcoding was used to confirm identifications in a subset of the larvae. C. fissus was more than eight times more abundant than any other species, and while abundance varied by species, cyprid density was highest for all species except for M. rosa before and after the El Niño event, and lower during the environmental disturbance. There were significant differences in cross-shore distributions among cyprid species, with some located farther offshore than others, along with variability in cross-shore distributions by season. C. fissus cyprids were closest to shore during spring-summer cruises when waters were the most thermally stratified, which supports previous findings that C. fissus cyprids are constrained nearshore when thermal stratification is high. Relative species proportions varied throughout the study, but there was no obvious change in species assemblage or richness associated with El Niño. We speculate that barnacle cyprid species diversity did not increase at our study site during the 2015-16 El Niño, as it has in other areas during previous El Niño Southern Oscillation events, due to the lack of anomalous northward flow throughout the 2015-16 event.

The highly toxic and cryptogenic clinging jellyfish Gonionemus sp. (Hydrozoa, Limnomedusae) on the Swedish west coast.

The clinging jellyfish Gonionemus sp. is a small hydromedusa species known historically from the Swedish west coast but not reported in recent times. This species is thought to be native to the northwest Pacific where it is notorious for causing severe stings in humans and is considered invasive or cryptogenic elsewhere. This year, unlike in the past, severe stings in swimmers making contact with Gonionemus sp. medusae occurred in Swedish waters from a sheltered eelgrass bed in the inner Skagerrak archipelago. To the best of our knowledge, this is only the second sting record of Gonionemus sp. from the Northeast Atlantic-with the first record occurring off the Belgian coast in the 1970s. Stinging Gonionemus sp. medusae have also been recently reported from the northwestern Atlantic coast, where, like on the Swedish coast, stings were not reported in the past. We analyzed sea surface temperature data from the past 30 years and show that 2018 had an exceptionally cold spring followed by an exceptionally hot summer. It is suggested that the 2018 temperature anomalies contributed to the Swedish outbreak. An analysis of mitochondrial COI sequences showed that Swedish medusae belong to the same clade as those from toxic populations in the Sea of Japan and northwest Atlantic. Gonionemus sp. is particularly prone to human-mediated dispersal and we suggest that it is possible that this year's outbreak is the result of anthropogenic factors either through a climate-driven northward range shift or an introduction via shipping activity. We examined medusa growth rates and details of medusa morphology including nematocysts. Two types of penetrating nematocysts: euryteles and b-mastigophores were observed, suggesting that Gonionemus sp. medusae are able to feed on hard-bodied organisms like copepods and cladocerans. Given the now-regular occurrence and regional spread of Gonionemus sp. in the northwest Atlantic, it seems likely that outbreaks in Sweden will continue. More information on its life cycle, dispersal mechanisms, and ecology are thus desirable.

The distribution and mitochondrial genotype of the hydroid Aglaophenia latecarinata is correlated with its pelagic Sargassum substrate type in the tropical and subtropical western Atlantic Ocean.

The pelagic brown macroalga Sargassum supports rich biological communities in the tropical and subtropical Atlantic region, including a variety of epiphytic invertebrates that grow on the Sargassum itself. The thecate hydroid Aglaophenia latecarinata is commonly found growing on some, but not all, Sargassum forms. In this study, we examined the relationship between A. latecarinata and its pelagic Sargassum substrate across a broad geographic area over the course of 4 years (2015-2018). The distribution of the most common Sargassum forms that we observed (Sargassum fluitans III and S. natans VIII) was consistent with the existence of distinct source regions for each. We found that A. latecarinata hydroids were abundant on both S. natans VIII and S. fluitans III, and also noted a rare observation of A. latecarinata on S. natans I. For the hydroids on S. natans VIII and S. fluitans III, hydroid mitochondrial genotype was strongly correlated with the Sargassum substrate form. We found significant population genetic structure in the hydroids, which was also consistent with the distributional patterns of the Sargassum forms. These results suggest that hydroid settlement on the Sargassum occurs in type-specific Sargassum source regions. Hydroid species identification is challenging and cryptic speciation is common in the Aglaopheniidae. Therefore, to confirm our identification of A. latecarinata, we conducted a phylogenetic analysis that showed that while the genus Aglaophenia was not monophyletic, all A. latecarinata haplotypes associated with pelagic Sargassum belonged to the same clade and were likely the same species as previously published sequences from Florida, Central America, and one location in Brazil (São Sebastião). A nominal A. latecarinata sequence from a second Brazilian location (Alagoas) likely belongs to a different species.

A new species of Gonionemus (Hydrozoa: Limnomedusae) from southern Australia.

A new species of a crawling limnomedusa belonging to the genus Gonionemus (Hydrozoa: Limnomedusae: Olindiidae) was collected from the brown alga Cystophora monilifera in an intertidal rock pool in Western Port, Victoria, Australia. The new species is described and compared with the three known species of Gonionemus. The mitochondrial DNA barcode markers cytochrome oxidase I and 16S rRNA were sequenced and compared to sequences from other olindiid species. The sequencing results corroborate the morphological findings.

Species-specific crab predation on the hydrozoan clinging jellyfish Gonionemus sp. (Cnidaria, Hydrozoa), subsequent crab mortality, and possible ecological consequences.

Here we report a unique trophic interaction between the cryptogenic and sometimes highly toxic hydrozoan clinging jellyfish Gonionemus sp. and the spider crab Libinia dubia. We assessed species-specific predation on the Gonionemus medusae by crabs found in eelgrass meadows in Massachusetts, USA. The native spider crab species L. dubia consumed Gonionemus medusae, often enthusiastically, but the invasive green crab Carcinus maenus avoided consumption in all trials. One out of two blue crabs (Callinectes sapidus) also consumed Gonionemus, but this species was too rare in our study system to evaluate further. Libinia crabs could consume up to 30 jellyfish, which was the maximum jellyfish density treatment in our experiments, over a 24-hour period. Gonionemus consumption was associated with Libinia mortality. Spider crab mortality increased with Gonionemus consumption, and 100% of spider crabs tested died within 24 h of consuming jellyfish in our maximum jellyfish density containers. As the numbers of Gonionemus medusae used in our experiments likely underestimate the number of medusae that could be encountered by spider crabs over a 24-hour period in the field, we expect that Gonionemus may be having a negative effect on natural Libinia populations. Furthermore, given that Libinia overlaps in habitat and resource use with Carcinus, which avoids Gonionemus consumption, Carcinus populations could be indirectly benefiting from this unusual crab-jellyfish trophic relationship.

Mitochondrial diversity in Gonionemus (Trachylina:Hydrozoa) and its implications for understanding the origins of clinging jellyfish in the Northwest Atlantic Ocean.

Determining whether a population is introduced or native to a region can be challenging due to inadequate taxonomy, the presence of cryptic lineages, and poor historical documentation. For taxa with resting stages that bloom episodically, determining origin can be especially challenging as an environmentally-triggered abrupt appearance of the taxa may be confused with an anthropogenic introduction. Here, we assess diversity in mitochondrial cytochrome oxidase I sequences obtained from multiple Atlantic and Pacific locations, and discuss the implications of our findings for understanding the origin of clinging jellyfish Gonionemus in the Northwest Atlantic. Clinging jellyfish are known for clinging to seagrasses and seaweeds, and have complex life cycles that include resting stages. They are especially notorious as some, although not all, populations are associated with severe sting reactions. The worldwide distribution of Gonionemus has been aptly called a "zoogeographic puzzle" and our results refine rather than resolve the puzzle. We find a relatively deep divergence that may indicate cryptic speciation between Gonionemus from the Northeast Pacific and Northwest Pacific/Northwest Atlantic. Within the Northwest Pacific/Northwest Atlantic clade, we find haplotypes unique to each region. We also find one haplotype that is shared between highly toxic Vladivostok-area populations and some Northwest Atlantic populations. Our results are consistent with multiple scenarios that involve both native and anthropogenic processes. We evaluate each scenario and discuss critical directions for future research, including improving the resolution of population genetic structure, identifying possible lineage admixture, and better characterizing and quantifying the toxicity phenotype.

A crab swarm at an ecological hotspot: patchiness and population density from AUV observations at a coastal, tropical seamount.

A research cruise to Hannibal Bank, a seamount and an ecological hotspot in the coastal eastern tropical Pacific Ocean off Panama, explored the zonation, biodiversity, and the ecological processes that contribute to the seamount's elevated biomass. Here we describe the spatial structure of a benthic anomuran red crab population, using submarine video and autonomous underwater vehicle (AUV) photographs. High density aggregations and a swarm of red crabs were associated with a dense turbid layer 4-10 m above the bottom. The high density aggregations were constrained to 355-385 m water depth over the Northwest flank of the seamount, although the crabs also occurred at lower densities in shallower waters (∼280 m) and in another location of the seamount. The crab aggregations occurred in hypoxic water, with oxygen levels of 0.04 ml/l. Barcoding of Hannibal red crabs, and pelagic red crabs sampled in a mass stranding event in 2015 at a beach in San Diego, California, USA, revealed that the Panamanian and the Californian crabs are likely the same species, Pleuroncodes planipes, and these findings represent an extension of the southern endrange of this species. Measurements along a 1.6 km transect revealed three high density aggregations, with the highest density up to 78 crabs/m(2), and that the crabs were patchily distributed. Crab density peaked in the middle of the patch, a density structure similar to that of swarming insects.

Phylogeographic structure and northward range expansion in the barnacle Chthamalus fragilis.

The barnacle Chthamalus fragilis is found along the US Atlantic seaboard historically from the Chesapeake Bay southward, and in the Gulf of Mexico. It appeared in New England circa 1900 coincident with warming temperatures, and is now a conspicuous member of rocky intertidal communities extending through the northern shore of Cape Cod, Massachusetts. The origin of northern C. fragilis is debated. It may have spread to New England from the northern end of its historic range through larval transport by ocean currents, possibly mediated by the construction of piers, marinas, and other anthropogenic structures that provided new hard substrate habitat. Alternatively, it may have been introduced by fouling on ships originating farther south in its historic distribution. Here we examine mitochondrial cytochrome c oxidase I sequence diversity and the distribution of mitochondrial haplotypes of C. fragilis from 11 localities ranging from Cape Cod, to Tampa Bay, Florida. We found significant genetic structure between northern and southern populations. Phylogenetic analysis revealed three well-supported reciprocally monophyletic haplogroups, including one haplogroup that is restricted to New England and Virginia populations. While the distances between clades do not suggest cryptic speciation, selection and dispersal barriers may be driving the observed structure. Our data are consistent with an expansion of C. fragilis from the northern end of its mid-19th century range into Massachusetts.

Phylogenetic analysis with multiple markers indicates repeated loss of the adult medusa stage in Campanulariidae (Hydrozoa, Cnidaria).

The Campanulariidae is a group of leptomedusan hydroids (Hydrozoa, Cnidaria) that exhibit a diverse array of life cycles ranging from species with a free medusa stage to those with a reduced or absent medusa stage. Perhaps the best-known member of the taxon is Obelia which is often used as a textbook model of hydrozoan life history. However, Obelia medusae have several unique features leading to a hypothesis that Obelia arose, in a saltational fashion, from an ancestor that lacked a medusa, possibly representing an example of a rare evolutionary reversal. To address the evolution of adult sexual stages in Campanulariidae, a molecular phylogenetic approach was employed using two nuclear (18S rDNA and calmodulin) and two mitochondrial (16S rDNA and cytochrome c oxidase subunit I) genes. Prior to the main analysis, we conducted a preliminary analysis of leptomedusan taxa which suggests that Campanulariidae as presently considered needs to be redefined. Campanulariid analyses are consistent with morphological understanding in that three major clades are recovered. However, several recognized genera are not monophyletic calling into question some "diagnostic" features. Furthermore, ancestral states were reconstructed using parsimony, and a sensitivity analysis was conducted to investigate possible evolutionary transitions in life-history stages. The results indicate that life-cycle transitions have occurred multiple times, and that Obelia might be derived from an ancestor with Clytia-like features.