Sunday, November 18, 2007

The Vampire Squid from Hell (Vampyroteuthis infernalis)



Classification of the Vampire Squid:
Phyla: Mollusca
Class: Cephalopoda
Subclass: Coleoidea
Superorder: Octopodiformes
Order: Vampyromorpha
Family: Vampyroteuthidae
Genus: Vampyroteuthis
Species: infernalis

The superorder of the vampire squid puzzled scientists for quite some time. The Decapodiformes (squid and cuttlefish) exhibit the ancestral condition of 10 arms (in squid two arms are modified into long tentacles) while Octopodiformes (octopi and vampire squid) exhibit the derived condition of 8 arms. However, Vampire Squid have 8 arms as well as 2 long filaments of different structure than arms. These could be derived from tentacles found in squid or may have arisen independently after the ancestors of Vampire Squid and Octopi lost 1 pair of arms. In either case, Vampire Squid are considered a relict, and represent a moment in the evolutionary history of Cephalopods and help to examine the evolution Squid/Cuttlefish and Octopi.



The oldest confirmed fossil Vampyromorphid is Vampyronassa rhodanica, from 165-164 mya

Body Form of the Vampire Squid:

Vampire squid are small, with a maximum size of 28cm, but more typical sizes are 8cm-13cm. They are bright red, from which they get their name, and are very gelatinous in composition, almost like that of jellyfish. They are more similar to squid in body shape than octopi. The vampire squid have 8 arms with suckers of the distal end and 2 long, retractible sensory tentacles, known as velar filaments. Between the arms there is extensive membranous webbing which is black on the underside, and two pouches in the webbing can conceal the velar filaments. On the underside of the webbing and arms are spike-like projections called cirri, but they are as soft as the rest of the organism. They have very large eyes and perhaps the largest eye to body ratio in the animal kingdom.

The webbing and cirri associated with the arms of a Vampire Squid.


The have two fins towards the apex of the mantle. These fins are responsible for the primary locomotion in these animals. These fins are not entirely homologous to squid fins in a similar position. Juvenile Vampire Squid have a pair of fins close to the apex of the mantle, and these are homologous to squid fins, but as it matures it undergoes a metamorphosis. A second pair of fins are produced behind the first and the first pair are re-absorbed. Therefore, there is a stage in development in which vampire squid have 4 fins, and this lead to early confusion in regards to number of species of Vampire Squid.


A juvenile Vampire Squid undergoing metamorphosis - this is the 4-fin stage.


Vampire Squid have poorly developed chromatophores and is unable to produce the complex colour changes of squid and cuttlefish. They also lack ink sacs. As they live in deep, dark environments, they instead have complex photophores for display (ie: to scare predators or attract prey). These photophores are located on the tips of each arm as well as at the base of the fins. Additionally, clouds of glowing particles can be ejected from the tips of the arms, functioning similarly to ink in squid (to startle a predator that has come too close). The control of the photophores is finely tuned – the length of the display can be very short or sustained and the intensity of the display can also be controlled. These photophores are involved in complex displays which I will discuss later.



Distribution:

Vampire Squid are distributed across temperate and tropical regions of oceans all around the world. Their vertical distribution in the water column is of more significance than their global distribution. They are found at depths between 600m and 1200m. This range in depth often represents masses of water with extremely low, but very stable, oxygen concentrations(~0.2ml O2/L water) – these are known as Oxygen Minimal Zones. Vampire Squid are some of the few species of marine invertebrates which can live in the zones continuously, and are the only cephalopods to do so. They are able to survive in these zones due to behavioural and physiological adaptations which maximum the efficiency of oxygen uptake and use:
- efficient fin-based swimming
- large gill surface area
- a hemocyanin which very efficiently binds oxygen in the blood
- a low metabolic rate
- prey capture by stationary display and detection (see Behaviours section)
- predator evasion by stationary display (see Behaviours section)
- gelatinous tissue filled with ammonia maintains neutral buoyancy
Vampire Squid Behaviour:

The method by which Vampire Squid detect prey and evade predators is very complex and highly specialized. First, to detect prey Vampire Squid hang motionless in the water column, with its velar filaments hanging down below. When a prey (such as copepods, prawns and cnidarians) bumps into the velar filaments in the dark or when attracted to light produced by photophores, the stimuli is transmitted up the filament and the Vampire Squid responds by rapidly swinging itself around and envelops its prey using its webbed arms.

When approached by a predator, the Vampire Squid goes to great lengths to confuse it. It begins by taking the “pineapple pose” – it inverts its webbing and arms, surrounding most of its head and mantle. The black colouring of the webbing is more difficult to see and the cirri appear sharp and formidable. The Vampire Squid then opens the photophores near its fins, creating the impression of two great glowing eyes. It then shrinks the size of these photophores to give the impression of retreating. Additionally, the photophores on the arm tips light up and are waved in a fashion that makes it appear to be coming closer to the predator. Generally the predator is too confused on the position of its prey to attack. If this display fails, and the predator approaches further, it ejects the glowing particles from its arm tips and makes a rapid escape.


The "pineapple pose" - the webbing and arms have been inverted, exposing the cirri.



The bright points on this image are the arm tip and fin base photophores, lit up in response to the presence of a dangerous predator (the camera).

Vampire Squid Videos:

A documentary by National Geographic on Vampyroteuthis infernalis:
Vampire Squid movement and photophore display (audio is in Japanese, refer to Behavioural section for descriptions):



(The english audio for this video can be heard here - http://youtube.com/watch?v=Q2o6t-0fU10 at 4:53, however embedding has been disabled)

Images of Vampire Squids:

This is the what the Vampire Squid from the original description of this species looked like. It was described as "a very small but terrible octopus, black as night, with ivory white jaws and blood red eyes". As you can see from the following images, this description could not be farther from the truth.


The terrible "Vampire Squid from Hell".


The massive eye of the Vampire Squid. Like other cephalopod eyes, the Vampire Squid's eyes are complex and highly evolved. The are not totally unlike our own eyes.

After detecting movement with its velar filaments, the Vampire Squid swings around and envelops its food. The Vampire Squid takes on such formidable prey as copepods, cnidarians, prawns and small fish.









Annotated bibliography 1:

Seibel, B.A., Chausson, F., Lallier, F.H., Zal, F. And Childress, J.J. 1999. Vampire blood: respiratory physiology of the vampire squid (Cephalopoda: Vampyromorpha: Vampyroteuthidae) in relation to the oxygen minimum layer. Experimental Biology Online. 4:1-10.

The study was conducted to examine the physiological adaptations of the Vampire Squid which allow it to survive in an extreme environment (minimal oxygen) where other species of Cephalopods and Invertebrates cannot. Seibel et al. hypothesized that the hemocyanin of Vampire Squid must be extremely high and also that it’s affinity to oxygen must be much higher than that of other cephalopods, to be able to survive.

The Vampire Squid is the only cephalopod able to carry out its entire life cycle in the core of the oxygen minimal zone. Pelagic crustaceans in the OMA have large ventilatory volume, large gill surface areas, short diffusion distances for oxygen to travel into the blood, and hemocyanin proteins with high oxygen affinities. Vampire Squids have moderate gill surface area by comparison as well as moderate diffusion distances and the connection between ventilation and locomotion prevents consistent high respiratory rates. The Vampire Squid must be physiological adapted in such a way as to be able to survive in these zones, and therefore and therefore the hemocyanin oxygen affinities must be important.

Specimens were collected off the coast of California and blood was collected and frozen. Oxygen dissociation curves were established by adjusting the pH of the blood as well as a diffusion chamber was used to regulate oxygen levels, and the ionic components/concentrations of the blood were determined. The hemocyanin-oxygen affinity measured was extremely high, and higher than any other cephalopod.

The extremely low oxygen content of the waters that the Vampire Squid inhabit (OMZs) requires that this species must have efficient and effective absorption of oxygen from the surrounding environment. Vampire Squid’s hemocyanin was found to have the highest oxygen affinity of any cephalopod. The efficiency of this hemocyanin was found to be one of the most important physiological adaptations of Vampire Squid to the OMZs.

Annotated bibliography 2:

Seibel, B.A., Thuesen, E.V. and Childress, J.J. 1998. Flight of the Vampire: Ontogenetic Gait-Transition in Vampyroteuthis infernalis (Cephalopoda: Vampyromorpha: Vampyroteuthidae). The Journal of Experimental Biology. 201:2413-2424.

The goal of this study was to investigate a change in swimming method over the course of the Vampire Squid’s life cycle and its relation to efficiency in the oxygen minimal zone. It was examined whether or not there was a relationship between the size of the organism and the efficiency of fin versus jet-propulsion swimming as well as whether or such relationships can result in transitions in locomotion through ontogeny.

Fin swimming is more common in adult Vampire Squid than jet-propulsion, due to its energetic efficiency. However, juvenile Vampire Squid may rely heavily on jet-propulsion and eventually undergo a metamorphosis which results in a change in fin position, relative size and shape as the individual becomes an adult and transitions to fin swimming.

Citrate synthase (CS) and octopine dehydrogenase (ODH) are useful indicators of aerobic and anaerobic metabolism and can be used to determine where the main areas of muscle contractions (for locomotion) are concentrated. Vampire Squid specimens were captured and the shape, size and position of the fins was recorded. CS and ODH levels were measured in the fin muscles (fin swimming) and mantle muscles (jet-propulsion) in juveniles and adults. High levels were shown in the fin muscle in adults, and high levels were shown in the mantle muscle in juveniles.

Changes from jet-propulsion to fin swimming were associated with a change in fin shape, size and position due to metamorphosis from juvenile to adult. Therefore there is a relationship between size of an organism and most efficient swimming methods. The transition between swimming styles can be defined as an “ontogenetic gait-transition”.

Annotated bibliography 3:

Bonnaud, L., Boucher-Rodoni, R. and Monnerott, M. 1997. Phylogeny of Cephalopods Inferred from Mitochondrial DNA Sequences. Molecular Phylogenetics and Evolution. 7(1):44-54.

The class Cephalopoda, of the Phylum Mollusca, was divided into groups primarily based on morphological and paleontological characters. Many of the relationships within this group, such as the relationship between Vampire Squid (Vampyromorpha) and Decapods/Octopods. The purpose of this study was to determine the accuracy of previous phylogenetic studies.

Cephalopoda is currently divided into Nautiloidea (Nautiloids) and Coleoidea (Decapods – squid and cuttlefish, Octopods – Octopi). The monophyly of many groups, such as the decapods and groups within the decapods, was not sufficiently supported by morphological evidence alone. For example: Vampire Squid have characters similar to both decapods and octopods, and may be a sister group to either. Mitochondrial DNA variation can be used to infer phylogenetic relationships, and was examined in this study.

Specimens of numerous groups of Cephalopods were collected and the mtDNA was isolated from tissue homogenates. PCR amplification was used on the mtDNA and these samples were sequenced. Phylogenetic analyses were carried out on the mtDNA sequences to determine the relationships between the organisms being studied. The monophyly of decapods was confirmed and Vampyromorpha was identified as being a sister group to all other octopods.

Some of the relationships between Cephalopods (as determined by morphological and paleontological data) were determined to be inaccurate and others were more extensively resolved. The Vampire Squid was placed as a sister group to Octopods, and not within the Decapods, despite its confusing morphological characters.
References and Links:

Bonnaud, L., Boucher-Rodoni, R. and Monnerott, M. 1997. Phylogeny of Cephalopods Inferred from Mitochondrial DNA Sequences. Molecular Phylogenetics and Evolution. 7(1):44-54.

The Cephalopod Page - http://www.thecephalopodpage.org/vampy.php

MarineBio.org - http://marinebio.org/species.asp?id=179

Seibel, B.A., Chausson, F., Lallier, F.H., Zal, F. And Childress, J.J. 1999. Vampire blood: respiratory physiology of the vampire squid (Cephalopoda: Vampyromorpha: Vampyroteuthidae) in relation to the oxygen minimum layer. Experimental Biology Online. 4:1-10.

Seibel, B.A., Thuesen, E.V. and Childress, J.J. 1998. Flight of the Vampire: Ontogenetic Gait-Transition in Vampyroteuthis infernalis (Cephalopoda: Vampyromorpha: Vampyroteuthidae). The Journal of Experimental Biology. 201:2413-2424.

Tree of Life Project - http://www.tolweb.org/Vampyroteuthis_infernalis/20084

Wikipedia - http://en.wikipedia.org/wiki/Vampire_squid