Untangling the diversity and evolution of Sea Hares
Aplysia parvula; Føllingen, Norway; Photo by Nils Aukan
Sampling and freezing at Askøy
Dr Carlo M. Cunha from the Metropolitan University of Santos in Brazil (Universidade Metropolitana de Santos), a world expert in the diversity and systematics of Anaspidea heterobranch gastropods, visited the Natural History Museum of Bergen for a month during January/February 2017 to study our scientific collection of these molluscs. The visit was funded by the University of Bergen´s Strategic Programme for International Research and Education (SPIRE).
The Museum holds a large amount of material from the Scandinavian region, but also from the Mediterranean, Macaronesia islands, Caribbean, and western Indian Ocean.
These marine molluscs commonly known by sea hares comprise around 90 currently known species and have long been of major interest to biologists because of their large and easily accessible nervous system, which form the basis of numerous neurophysiological works.
Preserved specimen of Aplysia punctata from Norway
Dissected specimen of Aplysia punctata from Norway
However, the taxonomy of these molluscs and their evolution are still poorly understood. Dr Cunha is using a combination of molecular and morphological tools to learn more about the worldwide diversity of anaspideans and their phylogenetic relationships.
Dr Cunha visit to Bergen has already resulted in the revision and update of the taxonomy of our Anaspidea collection. The Norwegian species of anaspids were revised and redescribed in detail using electron microscopy and DNA barcoding performed in collaboration with Louise Lindblom (University Museum / Biodiversity Labs).
SEM-image of jaws of Phyllaplysia sp from Florida, USA
Additionally several other species from around the world were studied and will be integrated in ongoing taxonomic revisions. Keep tuned!
We’ve also had Lloyd visiting recently, you’ll find a post about that on the Marine Invertebrates of Western Africa blog: click here
A whale recently had to be put down by wildlife management after it had repeatedly beached itself on the island of Sotra outside of Bergen. It was found to be a Cuvier’s beaked whale (Ziphius cavirostris), a species with apparently no official previous records from Norway. The University Museum of Bergen therefore wished to include the whale skeleton in its collections (and future exhibitions, once the remodelling completes).
Arriving at Espegrend
The whale was transported to the Marine Biological Station of Espegrend, and a team of five people from the museum set to work collecting measurements of the whale, taking tissue samples for DNA-barcoding though the NorBOL-project, collecting ectoparasites, and doing photo-documentation.
We then began removing the blubber and muscle tissue off the whale so that the bones can be further treated (they contain a lot of oil which needs to be taken care of once the soft tissue has been removed), before the skeleton can be mounted for display.
Starting the work of removing blubber and muscles
Little did we know that what had so far been a local news matter would soon go viral…
Sadly, it became clear during the autopsy that the whale had been ingesting massive amounts of plastic – as much as 30 plastic bags, and many smaller pieces of plastic. The whale was emaciated, and we believe that the plastic had gathered in such an amount in its stomach that it had created a plug, stopping the digestive process.
The plastic in and from the whale stomach (photos: T. Lislevand, H.Glenner/C.Noever)
The images of all the plastic spread out on the ground became a potent reminder of the tragedies that marine pollution is creating, and has sparked a renewed debate on how we can limit the amount of micro- and macro-plastic that end up in nature.
The news of the whale’s stomach content became international news
What should the Cuvier’s beaked whale have been eating?
Occurring as solitary animals or in small pods, and preferring the deeper open waters, the Cuvier’s beaked whale is not an easy animal to study. We do know that the species have a more or less cosmopolitan distribution, and that it holds the world record for longest and deepest dive for any mammal: one was recorded diving down to 3000 meters.
What data we do have on the species diet comes from beached individuals, and suggests that the species may be a fairly omnivorous predator. From the limited number of Cuvier’s beaked whales that have been examined for stomach content, there are regional differences in the diet, but it seems to consist mainly of cephalopods (squid and octopuses), deep sea fish, and medium sized crustaceans (Santos og andre 2001).
Above are the suckers on the arm of a giant squid, Architeuthis. Below are scars on the skin of a sperm whale. Photo: E.Willassen
The cephalopods appear to be the dominant food source, but this interpretation may be influenced by the longevity of the hard parts of a cephalopod in the stomach.
The tough beaks of a cephalopod consist of chitin, and is used for tearing prey to pieces. Chitin is also found in the suckers of many cephalopods. The beaks can be used to identify the cephalod groups based on their size and shapes. Animals such as jellyfish would be much harder to document as part of the diet, as they would be digested much more rapidly and completely.
We don’t know how well resolved the information produced by the animal’s echo-location is, but it is conceivable that the plastic reflects signals in a way similar to the natural food of the whale, and is therefore “caught” and eaten.
Cephalopod beak, drawing by J.H. Emerton (from Wikimedia commons)
We did find some cephalopod beaks in between the plastic in the whale stomach – so far we have not had the time to attempt to identify these, but we will.
Amongst the plastic there are some cephalopod beaks (dark brown) and a bivalve shell (top left). Photo: C. Noever
The University Museum have extensive cephalopod collections, and long traditions for working with this group – from Dr. Jakob Johan Adolf Appellöf who began working here in 1890, to the material collected in the MAR-ECO project.
MAR-ECO workshop on cephalopoda
From the work of Santos et al 2001 we know that the following species are in the diet of European Curvier’s beaked whales, and are probably amongst the things our whale should have been eating:
Tewuthowenia megalops. Photo: Richard E. Young during MAR-ECO-cruise 2004.
Teuthowenia megalops is an odd squid that floats around in the open water with a propulsion system based on ammoniumchloride that the animal produces by digesting protein. The name “megalops” hints to the huge eyes, which also contain three light producing organs (chromatophores). The species seems to be common in deep water in the north Atlantic (Vecchione et al. 2008). For more information, see Wikipedia.
Mastigoteuthis agassizii was originally registered in whale stomachs as Mastigoteuthis schmidti, but from the work on the MAR-ECO project, three species of Mastigoteuthis were considered to all be M. agassizii. Some ambiguity remains about the species of this genus of oceanic squid with a broad distribution in the world’s oceans in depths ranging from 500 to 1000 meters. They have diurnal migration, and may be found hunting closer to the surface at night.
Taonius pavo seen ventrally (above) and dorsally. Illustration from Wikipedia.
This little squid is not very well known. It has been recorded from the Atlantic Ocean, but it may have a broader distribution. In this link you will find a video from the Bahamas at 850 m depth where the animal releases bio- luminescent “ink” to confuse a predator and escape.
Histioteuthis bonelli Photographed by Richard E.Young during the Mar-Eco-cruises in 2004
Histioteuthis bonelli, drawing by Ernst Haeckel.
Histioteuthis bonnellii has several names in English, one of which is “umbrella squid”. The name is due to the skirt-like membrane between the arms – when it splays its arms it resembles an umbrella. We don’t know much about the biology of H. bonellii, except that it has several close relatives in the world oceans, and that what has hitherto been considered one species (H. bonellii) may well turn out to be several species.
Todarodes sagittatus, the European flying squid, is one of the ten-armed cephalopods that may irregularly occur in schools along the Norwegian coast. T. sagittatus is subject to fisheries.
Vampyroteuthis infernalis – the vampire squid is a deep-sea squid with eight arms and a skirt-like mantle between its arms. It also has moveable wings on its body that it can use to manoeuvre with. The name “vampire squid” is not quite true – this is no blood sucker, but it traps organic material from the water masses using long, sticky threads. If threatened, it can invert the “skirt” over its head, resembling a hedgehog. It also has light producing organs towards the back of the body, and can create clouds of bioluminescence. Even with all these defences, it may end up in the stomach of a Cuvier’s beaked whale.
Pelagic crustaceans and deep sea fish are also amongst the recorded prey from Cuvier’s beaked whales. Amongst these we find the fairly large and shrimplike Gnathophausia, found within the order Lophogastrida, which has been studied extensively at the University of Bergen. We also found a bivalve shell in the stomach of our whale, which as far as we are aware of has not been recorded as part of their diet previously.
Plastic or food?
It may seem strange that the whale should ingest large amounts of plastic – why would it do that? If the whale primarily finds its pray by echolocation in the pitch black of the deep sea, it may well be that it is unable to differentiate between the reflected signal from a sheet of plastic, and that from one of its usual prey animals.
Unlike the sperm whales that hunt cephalopods in a similar way, the beaked does not have teeth to grab its pray. Instead they use a suction to ingest the food. Perhaps it is this feeding mode that becomes very unfortunate for the whales in a natural environment with an incredible amount of human garbage.
Hyalinoecia tubicola from the North Sea (by K. Kongshavn).
Quill worms belong to the annelid family Onuphidae and are called like that because of their unique tubes. The tubes are secreted by their inhabitants and are very light and rigid, resembling a quill, the basal part of a bird’s feather used for writing. Quill worms are epibenthic creatures capable of crawling on the surface of the sea floor carrying their tubes along. Their anterior feet are modified, strengthened and enlarged, bearing thick and stout bristles. These anterior feet are used for locomotion.
Quill worms are widely distributed in the ocean inhabiting mostly slope depths down to 2000 m. Being large in body size (up to 10-20 cm long), they can be quite abundant in some areas. Meyer et al. (2016) reported Hyalinoecia artifex reaching up to 70 ind./m2 in the Baltimore Canyon at 400 m water depth. Another quill worm, H. tubicola, which is very common in Norwegian waters, reached up to 272 ind./m2 at 365 m offshore of Chesapeake Bay (Wigley & Emery 1967).
Quill worms are believed to be motile scavengers. Baited monster camera experiments performed at 2000 m deep site in Baja California demonstrated that Hyalinoecia worms can accumulate in hundreds of specimens five hours after the bait (rotten fish) has been deployed (Dayton & Hessler 1972). Myer et al. (2016) analyzed the stable isotope content in Hyalinoecia artifex tissues confirming its secondary consumer status. Their results supported earlier observations on the gut content of the same species by Gaston (1987) showing the presence of the remains of various benthic invertebrates.
Video 1. Quill worm Hyalinoecia tubicola moving inside its tube (by K. Kongshavn).
Video 2. Quill worm Hyalinoecia tubicola protruding from the tube opening. Three antennae and a pair of palps are seen on the head. The first two pairs of feet are enlarged and strengthened (by K. Kongshavn).
Dayton, P.K., Hessler, R.R., 1972. Role of biological disturbance in maintaining diversity in the deep sea. Deep-Sea Research 19: 199–208.
Meyer, K.S., Wagner, J.K.S., Ball, B., Turner, P.J., Young, C.M., Van Dover, C.L. 2016. Hyalinoecia artifex: Field notes on a charismatic and abundant epifaunal polychaete on the US Atlantic continental margin. Invertebrate Biology 135: 211–224. doi:10.1111/ivb.12132
Gaston, G.R. 1987. Benthic polychaeta of the Middle Atlantic Bight: feeding and distribution. Marine Ecology Progress Series 36: 251–262.
Wigley, R.L., Emery, K.O. 1967. Benthic animals, particularly Hyalinoecia (Annelida) and Ophiomusium (Echinodermata), in sea-bottom photographs from the continental slope. In: Deep-Sea Photography. Hersey JB, ed., pp. 235–250. John Hopkins Press, Baltimore.
To recap, a species’ type is “…the objective standard of reference for the application of zoological names. When a new species or subspecies is described, the specimen(s) on which the author based his/her description become the type(s) (Article 72.1). In this way names are linked to type specimens, which can be referred to later if there is doubt over the interpretation of that name.
Consequently types are sometimes referred to as “onomatophores” which means name bearers.”
The location – sampling site – from which the type specimen is described is known as the type locality.
Michael Sars (image from Wikimedia)
As you have probably noticed, polychaetes (bristle worms) are a focus group in our lab, and several species have type localities close by.
The biologist and theologian Michael Sars (1805-1869) lived in the Bergen region for many years. He was a prolific taxonomist, naming 277 species of marine taxa according to the World Register of Marine Species (WoRMS).
Consequently there are quite a few species that have their type locality within easy daytrip-distance by ship for us.
On the hunt with R/V “Hans Brattstrøm”
One such locality is Glesvær, where Michael Sars described several new species in his work of 1835: Beskrivelser og Iagttagelser over nogle mærkelige eller nye i Havet ved den Bergenske Kyst levende Dyr af Polypernes, Acalephernes, Radiaternes, Annelidernes og Molluskernes Classer* (“Descriptions and Observations of some strange or new animals found off the coast of Bergen, belonging to the Classes …”).
The polychaete Amphicteis gunneri (Ampharetidae) is one of these species. It was first described by Michael Sars as Amphitrite gunneri (the species name is an homage to Johan Ernst Gunnerus (1718-1773) who was an active scientist within botany and zoology, as well as the bishop in Trondheim, and one of the founders of Det Kongelige Norske Videnskapers Selskap) in the publication above. Here are his original illustrations of the species:
Amphicteis gunneri by M. Sars (1835)
We have previously submitted several specimens of Amphicteis gunneri for DNA-barcoding through the NorBOL-project – and found that specimens that according to the keys in the literature should all come out nicely as A. gunneri in fact end up in several barcode-based groupings (BINs), meaning that they genetically different from each other. Then we need to unravel which one is the true A. gunneri, and decide what to do with the others. In such cases, material from type localities is invaluable. By sending in specimens identified by resident taxonomists as A. gunneri from the type locality, we hope to figure out which BIN represent A. gunneri, and which represent potentially new species.
We were also able to photograph live specimens showing the nice coloration of this worm. Fixed specimens loose this colour and become uniformly yellow/white (no dots).
Amphicteis gunneri collected at type locality. Photo: K.Kongshavn
Polina Borisova, a first year master student from the Zoological Department of the Moscow State University (Russia), is coming to the Invertebrate Collections of the University Museum of Bergen with a 1-month research visit in January 2017.
Polina is going to work on the bristle worms from the family Lumbrineridae studying the collection from West Africa and Norway. Her project is jointly supervised by Dr. Nataliya Budaeva from the University Museum of Bergen and Dr. Alexander Tzetlin from the Moscow University.
Various Lumbrineridae from West Africa, scale 1 mm (Photos from BOLD).
Lumbrineridae are the worms with relatively poor external morphology but complex jaw apparatus. The structure of jaws has been traditionally used in the systematics of the family in the generic diagnoses. Polina is utilizing the methods of microCT to study the jaws of lumbrinerids in 3D.
Jaws of Scoletoma fragilis from the White Sea scanned using microCT showing ventral solid mandibles, forceps-like maxillae I and denticulate maxillae II and II, carriers of maxillae are omitted (Photo: P. Borisova)
Polina is also going to sequence several genetic markers to reconstruct the first molecular phylogeny of the family. This will allow testing the current hypothesis on the intergeneric relationships within Lumbrineridae and will aid in tracing the evolution of jaws within the family.
Molluscs come in a variety of shapes and sizes, but some of the least known are perhaps the Aplacophora, or shell-less molluscs. Instead of a shell, these worm-shaped molluscs have a cuticle covered in calcareous spicules, or sclerites, that give them a beautiful, glistening appearance!
The very first species of aplacophoran mollusc, Chaetoderma nitidulum, was collected from the Swedish west coast and described by the Swedish taxonomist Sven Lovén in 1844. At the time, it was not even known what animal group the new, strange animal belonged to. It had spicules– could it be related to the spiny sea urchins? It had a worm-like body– could it be related to other worm-shaped animals? It would be almost 50 years before it was conclusively recognized as part of Mollusca. Since then, many more species have been discovered, and today close to 500 species of aplacophoran molluscs have been described.
A specimen of Chaetoderma nitidulum from the Norwegian West Coast Photo: N. Mikkelsen
Chaetoderma nitidulum is known today as one of the common aplacophoran molluscs in the East Atlantic, with a distribution from the Svalbard archipelago in the north, to the British Isles in the south. However, taxonomist have been debating the identity of Chaetoderma nitidulum since shortly after it was described. Some researchers have suggested that it could in fact consist of up to six different species. Other researchers have synonymized it with other species, or suggested that it is not a separate species, but only part of a larger species which has a distribution that spans the entire North Atlantic.
The shape, size and the patterns on the calcareous sclerites covering the body of the aplacophoran molluscs is unique to each species, making it one of the most important characters we have to distinguish between different species.
Calcareous clerites from Chaetoderma nitidulum. Photo: N. Mikkelsen
Looking at the sclerites through the microscope equipped with a cross-polarizing filter gives us a shiny, colorful view of the sclerites. The light shines with different colors depending on the thickness of the sclerites, helping us get a good view of the structure of the sclerites.
Sclerites from Chaetoderma nitidulum viewed under cross-polarized light. Photo: N. Mikkelsen
We have recently investigated specimens of Chaetoderma nitidulum from different localities from the entire distribution range of the species. Our investigations have revealed a lot of variation between the specimens, both in the calcareous sclerites and in DNA sequences, separating the specimens into at least two different groups. Could it be that Chaetoderma nitidulum actually represents more than one species?
Travelling alone through the water column may be a dangerous business: reaching the final destination is not always guaranteed, the risk of being eaten is high, and even finding food may prove a difficult task… which is why several animals choose to travel comfortably on or inside jellyfish and siphonophores!
Jellyfish are commonly involved in relationships of parasitism and phoresis (i. e., when one organism is mechanically transported by another without any further physiological dependence), and many examples have been observed of these interactions around the world. For instance, it’s not unusual to find hyperiid amphipods and sea-spiders –as well as lobster and crab larvae – piggybacking on the surface of large and tiny jellyfish, and while it’s still not clear whether or not all these passengers feed on their means of transportation, real parasitism and jelly-feeding has been confirmed for at least some of them. Jellyfish may also transport parasitic worms to their final hosts (like the nematode you see in the pictures), acting as carriers of parasites towards fish and mammals, and sometimes, eventually reaching humans as well!
Euphysa aurata medusa with parasitic nematode larva. Korsfjord, February 2016. Credit: Aino Hosia.
A close-up of 2 showing the parasite embedded in the mesoglea (jelly) of the host. Credit: Aino Hosia.
Euphysa aurata medusa with crustacean ectosymbiont. Raunefjord, December 2016. Credit: Luis Martell
These two hydromedusae of Euphysa aurata were collected this year in the fjords south of Bergen, and are only an example of jellyfish harboring other animals in this area. The species is a common and widespread jellyfish around here, but its role in the transmission of parasites and transportation of small crustaceans has never been explored. It might well be that, together with its gelatinous relatives, E. aurata will prove to be involved in many more biological interactions than we previously thought!
Scientific illustrations today are usually formed within quite strict limits. We use photographs or drawings of small details, and these are all connected to one specific specimen that preferably is to be found in a scientific collection.
But can other approaches also help us? The artist Pippip Ferner has long found her inspiration in nature, and especially the (marine) invertebrates. Maybe her pictures can inspire us to examine other details in our study-animals? Maybe a picture can inspire you to think more about nature, the sea, or invertebrates – their lives and lores? These are not pictures that are meant to be scientifically accurate, but rather fabulations inspired by the wild things that happen when evolution gets to do as it pleases…
Some of Pippis drawings are inspired from scientific drawings, both old and new, some are from animals we have looked at together.
Here are some of Pippips pictures from this year, and the animals that inspired them. These three pictures were chosen to be part of the Evolution and Art section of the international science conference Evolution this summer in Austin, TX.
“Tunicate anatomy” (c) Pippip Ferner
Pippip says about this first picture:
“A scientific illustration of a TUNICATE is the inspiration for this work. Tunicates are sort of last stage before vertebrates. Clues for this is found in the larva that has a notochord, comparable to the spine of vertebrates. It has cerebral vesicle equivalent to a vertebrate’s brain, sensory organs that includes an eyespot to detect light and an otolith, which helps the animal orient to the gravity.
Fascinated by the thought of this “slimy blob” having many features similar to humans resulted in this quite complex outcome. The overload of insistent lines has given the tunicate quite a sophisticated system.”
Komodo National Park sea squirt (Polycarpa aurata). Photo: Nick Hobgood (wikipedia)
On the left is a photo of a live tunicate. This photo is from Indonesia, but tunicates are common to find also in our colder waters. They can be solitary as this one, or colonial – where several tunicates form a colony together by budding, so that one large colony basically has the exact same DNA. Most tunicates are sessile (they sit attached to one place), but some live floating around in the water. The best known of these pelagic tunicates are the salps of the southern oceans.
Internal anatomy of a tunicate (Urochordata). Adapted, with permission, from an outline drawing available on BIODIDAC. (Wikipedia)
A scientific illustration of a tunicate in a Biology textbook will look something like this:
Moving to other invertebrates, Pippip has worked with clams:
“Bivalve anatomy” (c) Pippip Ferner
“In this image I question how the clam lives in symbiosis with other species as its shell gets weaker due to climate changes. The drawing might resemble the results of some kind of scientific inquiry with references to the anatomy of a clam (bivalve).
In my work I let my own artistic evolutionary process make the clam into something more abstract.”
(If you wait until door # 22, there might be a story that relates to bivalves that live with others…)
This is a Ctenophore, a comb jelly:
“Comb jelly anatomy” (c) Pippip Ferner
“The starting point of this work was a detailed illustration from biologist Ernst Haeckel’s (Artforms in Nature) of a comb jelly/ctenophorae. The comb jelly differs from other jellyfish with more sophisticated nervous system with both synapses and individual muscle cells.
The outcome of this drawing is a tribute to the beauty of the structure of this organism.”
The comb jelly Pleurobrachia pileus. Photo: Aino Hosia
The comb jelly Mnemiopsis leidyi. Photo: Aino Hosia
The comb jelly Beroe abyssicola. Photo: Aino Hosia
Jelly fishes anf Comb jelly fishes. Illustration: Ernst Haeckel, Kunstformen der Natur 1904, plate 27
Ctenophores are predatory planktonic jellies. The special thing about them, according to our Jelly-specialist Aino, is that they have a rotational symmetry. The diagnostic feature of comb jellies are their comb-rows that they use for swimming. The photos above represent the three groups of comb jellies – all of them are present in Norway.
To the right is the Haeckel-picture she started from, and here is a film of Comb jellies from the Chicago Shedd Aquarium.
Today, I thought I’d introduce to you to a cool group of animals that is ubiquitous in the oceans (including the Norwegian seas), but unfamiliar to most people. Siphonophores (“kolonimaneter” in Norwegian) belong to cnidarians, a group that includes corals, anemones, hydroids and jellyfish, and is characterized by the presence of stinging cells used in prey capture. All siphonophores are predatory, and use their stinging tentacles to catch small crustaceans or, in the case of some species, even small fish.
The most (or only) familiar siphonophore for the majority of people is probably the highly venomous Portuguese Man O’War (Physalia physalis), which can be spotted floating on the surface of the ocean or stranded on beaches. However, it is not really representative of the group as a whole, as most siphonophores live in the water column of the open ocean rather than its surface. There are around 200 described species of siphonophores.
The most fascinating feature of siphonophores is their peculiar body plan. While siphonophores may appear to be a single animal, they are in fact a colony of physiologically connected and genetically identical but morphologically diverse individuals called zooids that have specialized to carry out different tasks for the colony. Siphonophores belong to the class Hydrozoa (“polyppdyr” in Norwegian), which covers two basic body plans: the polyp/hydroid and the medusa.
Schematic of a physonect siphonophore. From http://www.siphonophores.org (CC-by-nc-sa)
The various zooids comprising a siphonophore colony can also be divided into these main groups. For example, the zooids used for swimming, called nectophores, are medusoid, while the feeding zooids, or gastrozooids, are polyp-like. The siphonophore colony can also include specialized defensive, protective and reproductive zooids. All the zooids forming a colony arise by budding from a single fertilized egg. The different zooids are specialized to the degree that they cannot function as individual animals any more, and are only able to perform their specific tasks as parts of the siphonophore colony.
Anterior nectophore, posterior nectophore and eudoxid of the calycophoran siphonophore Dimophyes arctica – a common species in Norwegian waters. Photos by Aino Hosia (cc-by-sa)
The zooids, for example the swimming nectophores, vary in appearance between species, and can be used for species identification. In addition, the various types of zooids in the colony are arranged in a strict species specific pattern, providing the intact colonies of each species with their particular appearance. While the individual zooids are generally small, millimeters to centimeters in size, some siphonophore species, like Praya dubia, may have colonies that reach 40 m in length! Siphonophore colonies generally have a zone of one or more (up to several dozen) swimming nectophores at the front, used to pull the colony through water. Behind this nectosome is the siphosome, which contains the feeding, reproductive and other zooids in a repeating pattern, each iteration of which is called a cormidium. In some species (suborder Calycophorae), these cormidia are released as small free-living reproductive colonies called eudoxids. Unfortunately, siphonophore colonies are extremely fragile and tend to fall apart during standard plankton sampling with nets, leaving behind a bewildering array of small bits and pieces – part of the reason they are relatively poorly known to most people.
Colony of physonect siphonophore Physophora hydrostatica, aka hula skirt siphonophore. Photo by Aino Hosia (cc-by-sa)
Intact siphonophore colonies are beautiful, but often utterly alien in appearance. It is interesting to consider where to draw the line between an individual and a colony. While we as individuals have specialized organs to carry out our various bodily functions, siphonophore colonies are made up of specialized interdependent individuals or zooids similarly carrying out their specific tasks.
As part of project HYPNO we are charting the diversity of pelagic hydrozoans, including siphonophores, in Norway. There are ~15 species observed in Norwegian waters, and some, particularly Dimophyes arctica, Lensia conoidea and Nanomia sp. are extremely common components of marine plankton. However, siphonophores are primarily noticed when they become a nuisance: For example, mass occurrences of Muggiaea atlantica and Apolemia uvaria have in the past killed large numbers of farmed fish in Norway, with resulting losses to aquaculture companies.
When we think about what drives the ecosystems, much of the initial responsibility is put on the sunlight. This is mainly because of the photosynthesis, and thus the basic pieces of almost all food-webs, but light is also important for the animals. Many animals use visual cues to find food, and whether you search for food or do not want to become food, the presence (or absence) of light will help you.
Themisto sp swims up into the dark night. Photo: Geir Johnsen, NTNU
Seawater is a pretty good stopper of light. We don’t need to dive far down before we are in what we consider a dark place, and less and less light finds its way the deeper we come. We tend to call the depths between 200 and 1000 m “the twilight zone”: most light stops way before 200m and the last straggling lumens give up at 1000m.
Most places on earth has a daily division between a dark and a light period: night and day. This is the ultimate reason for what is often called “the largest motion on earth”: Millions of zooplankton hide out in the darker parts of the water column during the day, and then move up to feed on the plants living in the light-affected parts of the water during the night (when predators will have a hard time seeing them). This daily commute up and down is called Diel Vertical Migration (DVM).
Themisto sp among the many smaller particles. (The light in this picture is from a flash). Photo: Geir Johnsen, NTNU
But what about the waters north of the polar circle? These areas will for some time during the winter have days when the sun stays under the horizon the entire day – this is “the Dark time” (Mørketid). At higher latitudes, there will be several days, or even weeks or months when the sun is so far below the horizon that not even the slightest sunset-glow is visible at any time. In this region, we have long thought that the Dark time must be a dead or dormant time.
The acoustic signals that gave the first indications of LVM. Figure 2 from Last et al 2016.
We could not have been more wrong! It turns out that during the polar night, the DVM moves from being on a 24 hr cycle (sunlight-induced), to a 24.8 hour cycle! What is now the driver? The moon !(The lunar day is 24.8 hrs). Another thing that shows us that the moon must give strong enough light that predators can hunt by it, is that every 29.5 days most of the zooplankton sinks down to a depth of 50m: this falls together with the moon being full. Researchers have started to call this LVM (Lunar-day Vertical Migration) to show the difference to the “normal” DVM. There are of course lots of complicated details such as the moons altitude above the horizon and its phase that influences the LVM, but in general we can say that during the polar night (the Very Dark time), the “day” as decided by light has become slightly longer than normal.
The full moon, photographed by the Apollo 11 crew after their visit. Photo: NASA, 1969
Themisto – the werewolf. Note that the whole head is dominated by eyes – this is a visual hunter! Photo: Geir Johnsen, NTNU
Some of the larger animals taking part in the LVM are the amphipods Themisto abyssorum and Themisto libellula. They are hunters – so their reason to migrate up in the water column is not the plants, but the animals eating the plants; their favourite food are copepods of the genus Calanus. These are nice and quite energy-rich small crustaceans that eat the microscopic plants in the upper water column. We have sampled both Themisto-species in the middle of the winter (january), and their guts were filled to the brim with Calanus, so we know that they continue hunting by moon-light. They are such voracious hunters that some researchers have started to call them marine werewolves: the moonlight transforms them from sedate crustaceans to scary killers…
But, if they are the hunters, why do they spend so much time in the deep and dark during the lighter parts of the day? The hunters are of course also hunted. Fish such as polar cod (Boreogadus saida), birds such as little auk (Alle alle) and various seals like to have their fill of the Themisto species. So – life has its ups and downs, and the dance of hunter and hunted continues into the dark polar night…
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