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Helical motion is prevalent in nature and has been shown to confer stability and efficiency in microorganisms. However, the mechanics of helical locomotion in larger organisms (>1 centimeter) remain unknown. In the open ocean, we observed the chain forming salp,Iasis cylindrica, swimming in helices. Three-dimensional imaging showed that helicity derives from torque production by zooids oriented at an oblique orientation relative to the chain axis. Colonies can spin both clockwise and counterclockwise and longer chains (>10 zooids) transition from spinning around a linear axis to a helical swimming path. Propulsive jets are non-interacting and directed at a small angle relative to the axis of motion, thus maximizing thrust while minimizing destructive interactions. Our integrated approach reveals the biomechanical advantages of distributed propulsion and macroscale helical movement. 

Abstract Gelatinous zooplankton are increasingly recognized as key components of pelagic ecosystems, and there have been many recent insights into their ecology and roles in food webs. To examine the trophic ecology of siphonophores (Cnidaria, Hydrozoa), we used bulk (carbon and nitrogen) and compound‐specific (nitrogen) isotope analysis of individual amino acids (CSIA‐AA). We collected samples of 15 siphonophore genera using blue‐water diving, midwater trawls, and remotely operated vehicles in the California Current Ecosystem, from 0 to 3000 m. We examined the basal resources supporting siphonophore nutrition by comparing their isotope values to those of contemporaneously collected sinking and suspended particles (0–500 m). Stable isotope values provided novel insights into siphonophore trophic ecology, indicating considerable niche overlap between calycophoran and physonect siphonophores. However, there were clear relationships between siphonophore trophic positions and phylogeny, and the highest siphonophore trophic positions were restricted to physonects. Bulk and source amino acid nitrogen isotope (δ¹⁵N) values of siphonophores and suspended particles all increased significantly with increasing collection depth. In contrast, siphonophore trophic positions did not increase with increasing collection depth. This suggests that microbially reworked, deep, suspended particles with higher δ¹⁵N values than surface particles, likely indirectly support deep‐pelagic siphonophores. Siphonophores feed upon a range of prey, from small crustaceans to fishes, and we show that their measured trophic positions reflect this trophic diversity, spanning 1.5 trophic levels (range 2.4–4.0). Further, we demonstrate that CSIA‐AA can elucidate the feeding ecology of gelatinous zooplankton and distinguish between nutritional resources across vertical habitats. These findings improve our understanding of the functional roles of gelatinous zooplankton and energy flow through pelagic food webs. 

Siphonophores (Cnidaria: Hydrozoa) are abundant and diverse gelatinous predators in open-ocean ecosystems. Due to limited access to the midwater, little is known about the diets of most deep-dwelling gelatinous species, which constrains our understanding of food-web structure and nutrient flow in these vast ecosystems. Visual gut-content methods can rarely identify soft-bodied rapidly-digested prey, while observations from submersibles often overlook small prey items. These methods have been differentially applied to shallow and deep siphonophore taxa, confounding habitat and methodological biases. DNA metabarcoding can be used to assess both shallow and deep species’ diets under a common methodological framework, since it can detect both small and gelatinous prey. We (1) further characterized the diets of open-ocean siphonophores using DNA metabarcoding, (2) compared the prey detected by visual and molecular methods to evaluate their technical biases, and (3) evaluated tentacle-based predictions of diet. To do this, we performed DNA metabarcoding analyses on the gut contents of 39 siphonophore species across depths to describe their diets, using six barcode regions along the 18S gene. Taxonomic identifications were assigned using public databases combined with local zooplankton sequences. We identified 55 unique prey items, including crustaceans, gelatinous animals, and fish across 47 siphonophore specimens in 24 species. We reported 29 novel predator-prey interactions, among them the first insights into the diets of nine siphonophore species, many of which were congruent with the dietary predictions based on tentilla morphology. Our analyses detected both small and gelatinous prey taxa underrepresented by visual methods in species from both shallow and deep habitats, indicating that siphonophores play similar trophic roles across depth habitats. We also reveal hidden links between siphonophores and filter-feeders near the base of the food web. This study expands our understanding of the ecological roles of siphonophores in the open ocean, their trophic roles within the ‘jelly-web’, and the importance of their diversity for nutrient flow and ecosystem functioning. Understanding these inconspicuous yet ubiquitous predator-prey interactions is critical to predict the impacts of climate change, overfishing, and conservation policies on oceanic ecosystems. 

Predator specialization has often been considered an evolutionary “dead end” due to the constraints associated with the evolution of morphological and functional optimizations throughout the organism. However, in some predators, these changes are localized in separate structures dedicated to prey capture. One of the most extreme cases of this modularity can be observed in siphonophores, a clade of pelagic colonial cnidarians that use tentilla (tentacle side branches armed with nematocysts) exclusively for prey capture. Here we study how siphonophore specialists and generalists evolve, and what morphological changes are associated with these transitions. To answer these questions, we: a) Measured 29 morphological characters of tentacles from 45 siphonophore species, b) mapped these data to a phylogenetic tree, and c) analyzed the evolutionary associations between morphological characters and prey-type data from the literature. Instead of a dead end, we found that siphonophore specialists can evolve into generalists, and that specialists on one prey type have directly evolved into specialists on other prey types. Our results show that siphonophore tentillum morphology has strong evolutionary associations with prey type, and suggest that shifts between prey types are linked to shifts in the morphology, mode of evolution, and evolutionary correlations of tentilla and their nematocysts. The evolutionary history of siphonophore specialization helps build a broader perspective on predatory niche diversification via morphological innovation and evolution. These findings contribute to understanding how specialization and morphological evolution have shaped present-day food webs.