Spiders from the deep sea: Pycnogonida

Biodiversity is surprising. Did you know that there exist sea spiders? Pycnogonida, also known as “sea spiders”, form one of the strangest groups of arthropods that have ever existed. They belong to the subphylum Chelicerata (along with spiders) and are a part of the invertebrate fauna from seas and oceans all over the world. Although their number is so scarce and they camouflage so well with the environment they are very difficult to see!  

Do you want to know how to find and recognize them? Keep reading!

So…What are Pycnogonida?

Pycnogonida (from the Greek pykno = ‘lots of’ or ‘thick’ and góny = “knees”), also known as Pantopoda (‘totally made of legs’), is a class of marine benthic arthropods (benthos = organisms that live in association with the sea floor). It belongs to the subphylum Chelicerata, group that also includes the ‘true’ chelicerates or Euchelicerata: arachnids (spiders, scorpions, ticks and mites),  xiphosurans (a relict group of organisms commonly known as horseshoe crabs) and the extinct eurypterids (sea scorpions).

There are about 8-9 families, 86 genera (some of them are fossils) and up to 1000 species of Pycnogonida described worldwide, and all of them are a part of an unique living order: Pantopoda.

Pygnogonids from the Anctartic ocean; this specimen had a lenght of about 30cm (Picture: Keith Martin-Smith).

Pycnogonida live in almost all marine habitats to almost any depth and latitude from the equatorial to Polar Regions both in coastal areas and in the abyssal plains about 6000m depth (although they appear more usually in the Mediterranean Sea, the Caribbean Sea, and Arctic and Antarctic oceans). Although its cryptic appearance and its capacity for camouflage with the environment make them difficult to see at the first sight.

Pycnogonida from the species Nymphon gracile (Picture: Christophe Courteau)

Pycnogonida from the genus Pseudopallene (Picture: Claudia Arango)

Firstly, they were classified as arachnids because of its resemblance to spiders. Due to new anatomical and biological studies, they were reclassified as a new group inside chelicerata which was been related with arachnids. The possibility they were an ancestor of all chelicerates has also been considered, so they will form a very ancient group.

Phylogeny of Arthropoda from the Tree of Life Project (Source: tolweb.org)

External anatomy

Pycnogonida is a morphologically varied group of organisms with a wide range of sizes: from species that barely reach a few centimeters (which are commonly located in coastal benthos) to those that reach 50-70cm lenght (which tend to appear in abyssal depths).

Different species and their morphological differences (Cano E., 2015)

They have a spider-like body divided into two main parts: prosoma (head or cephalon + thorax) and opisthosoma (abdomen). On their head they feature a proboscis, that is, an organ that allows them to suck fluids from soft-bodied invertebrates; they also display 4 eyes on a tubercle and three pair of appendices: a pair of chelifores, a pair of palps and a pair of ovigerous legs (or ovigers), these last being exclusive to pycnogonids. Pycnogonids mainly use ovigerous legs for a self-cleaning function, but these structures seem to be more well developed on males in order to carry the eggs (and even the larvae when they hatch).

Pycngonida usually have 8 legs (even they sometimes suffer polymerization of their body segments and get one or two more pairs of legs, having 10-12 legs in total) which can be as thick as the rest of their body. This usually make them to look like if they were made only of legs (in fact, the term Pantopoda means ‘totally made of legs’).

External anatomy of Pycnogonida (Image source: http://sprott.physics.wisc.edu)

Internal anatomy

Pycnogonida are very strange creatures on their inside: they lack respiratory system (since a very thin cuticle covers their body, the gas exchange takes place through its surface) and excretory system; they have a reduced circulatory system and the nervous system is composed only of a simple brain and two ventral nerve cords. Gonads are located in the prosoma and they extend in the leg cavities; in most species, genital openings are also located on the legs.

Where and how do they live? 

As we said above, Pycnogonida form a group widely distributed in seas and oceans from all over the world. Whether they are located in the deeps or on the surface, they are always a part of the benthos.

Mainly biological components of marine ecosystems (Picture by Castro and Huber, 2007).

After a courtship that is still unknown, both males and females release their gametes in to the environment, where the fertilization occurs. Once fertilized, the eggs are gathered by the male, which will take care of them. To pick them up, it binds them together and sticks them to its body by secreting a sticky substances.

Male of the species Nymphon gracile taking care of a bunch of eggs sticked to its ovigerous legs (Picture by Christophe Courteau)

Tanystylum duospinum (© 2005 California Academy of Sciences, CC)

After hatching, it emerges a free-living larva known as protonymph, which will reach the adulthood by suffering a metamorphosis process (are you interested on metamorphosis? Click here to learn more about it!).

They became carnivorous once they grow; they use chelifora to capture and chop soft-bodied organisms and then use their proboscis to suck their fluids (as spiders do). Generally, they feed on benthic or sessile organisms, like anemones, sponges and bryozoans.

Pycnogonida from the genus Pseudopallene on a bryozoan (Picture by Claudia Arango).

NOTE: Pycnogonida are difficult to see at the first sight, but if you are curious people and you like diving, I encourage you to look into dense algae and sea phanerogams extensions. You will see one if you are lucky!

But it will be more easy to identify them if we have seen them in motion before (video from the Youtube channel Cloud. Tube):

Curiosities of the group

Pycnogonida form a very curious group, both for the morphological external traits that characterize its species and for its biological traits:

• Some species develop a parasitical larval stage that remains in a latent state inside other organisms (e.g. corals) until they reach adulthood, when they leave these organisms and adopt a free-living form.
• The polymerization of their legs due to the increment of corporal segments seems to be an exclusive phenomenon among arthropods.
• Many species of Pycnogonida lose their legs by a process called autotomy, that is, the separation of a body part by self-amputation (e.g. the loss of the tail in lizards).
• Pycnogonida is almost the only group in which the parental generation takes care of its descendants. The male is the one responsible for protecting, cleaning and oxygenating the eggs, even after hatching.

.         .         .

Nowadays, Pycnogonida is a fairly unkown group of organisms on many levels: they are phylogenetically old, scanty and often located so deep in the sea they turn out to be difficult to study. Moreover, no benefits for humans are been found from this group beyond its importance in terms of biodiversity. Unfortunately, this fact usually condemn strange organisms to oblivion.

And you, what do you think about this? Do you think is essential for us, humans, and for all life forms on Earth to preserve biodiversity?

References

• Australian Government. Department of Environment: Australian Antarctic Division. Web: http://www.antarctica.gov.au/science/cool-science/2010/sea-spiders-provide-insights-into-antarctic-evolution.
• Bamber, R. N. & A. El Nagar (Eds.) 2014. Pycnobase: World Pycnogonida Database. Accesible (2014) en: http://www.marinespecies.org/pycnobase/
• Biodiversidad, taxonomía y biogeografía de los Artrópodos de México. Vol. III, Edición: 2002, Capítulo: Pycnogonida (por Tomás Munilla), Publisher: Universidad Nacional Autónoma de México, Editores: J. Llorente y J. Morrone, pp.215-22.
• Blaxter J., Douglas B. (1987). Advances in Marine Biology, Volumen 24. Academic Press.
  
• Cano E., López P.J. (2015). Clase Pycnogonida, Orden Pantopoda. IDE@-SEA, nº 22 (30-06-2015): 1-13.
• Padilla F., Cuesta A. Zoología Aplicada. Ediciones Díaz de Santos, 2003.
  
• Fauna marina circalitoral del sur de la Península Ibérica: resultados de la campaña oceanográfica “Fauna I”. Editorial CSIC – CSIC Press, 1993.

Main image: Colossendeis megalonyx from the deeps of the Anctartic ocean by Norbert Wu/Minden Pictures/FLPA.

Limb regeneration, from the axolotl to human beings

The regeneration of lost or damaged body parts in animals is known from many centuries ago. In 1740 the naturalist Abraham Trembley observed a small cnidarian that could regenerate its head if it was cut off, so he called it Hydra, in reference to the monster from Greek mythology that could grow back its multiple heads if they were cut off. Afterwards, it was discovered that there were many other species of animals with regenerative abilities. In this entry we’ll talk about these animals.

Regeneration in the animal kingdom

Regeneration of body parts is more widespread between the different groups of invertebrates than it is between the vertebrates. This process can be bidirectional, in which both parts of the animal regenerate their missing parts to form two animals (just like the hydra, planarians, earthworms and starfishes) or unidirectional, in which the animal loses an extremity but it just regenerates, without forming two animals (arthropods, molluscs and vertebrates). In vertebrates, fishes and amphibians are the ones that present the greatest regenerative capacities, although many lizards and some mammals are able to regrow their tails.

Image by Matthew McClements about bidirectional regeneration in planarians, hydras and seastars. Extracted from Wolbert's Principles of Development.

Regeneration can be done by two different ways:

• Regeneration without active cellular proliferation or “morphallaxis”. In this type, the absent body part is regrown through remodelling of pre-existing cells. This is what happens in the Hydra, in which lost body parts are regenerated without the creation of new material. So, if a hydra is cut in half, we’ll obtain two smaller versions of the original hydra.

Video about an experiment in which an Hydra has been cut in different pieces. Video by Apnea.

• Regeneration with cellular proliferation or “epimorphosis”. In this type, the lost part is regenerated via cellular proliferation, it is “newly created”. In most cases, it happens through the formation of a specialized structure called blastema, a mass of undifferentiated cells which appears during phenomena of cellular regeneration.

Almost all groups of animals with regenerative capacities present regeneration with blastema formation. Yet the origin of the blastemal stem cells varies between groups. While planarians present pluripotent (that can differentiate to any kind of cell type) stem cells all along their bodies, vertebrates have specific cells in each type of tissue (cartilage, muscle, skin…) that only regenerate cells of the tissue they come from.

In land vertebrates, lizards and urodeles are the ones that present the most powerful regenerative abilities. Down below we’ll see how they regenerate and the applications it has in modern human medicine.

Expendable tails

When you are a small animal that is being chased by a cat or any other predator, it probably is better for you to lose your precious tail than to lose your life. Some terrestrial vertebrates have evolved following this philosophy, and they are able to shed off their tails voluntarily through a process called caudal autotomy. This allows them to escape from their predators, which are entertained with the still moving lost tail.

 Video in which we can see how some lizards like this red-tailed vanzosaur (Vanzosaura rubricauda) have brightly coloured tails to attract the attention of predators. Video by Jonnytropics.

Autotomy or self-amputation, is defined as a behaviour in which the animal can shed off one or more body parts. Caudal autotomy is found in many species of reptiles and in two species of spiny mouse of the genus Acomys. In reptiles we can find caudal autotomy in lacertids, geckos, skinks and tuataras.

Foto of a Cairo spiny mouse (Acomys cahirinus), a mammal which is able to shed and regrow its tail. Photo by Olaf Leillinger.

In reptiles, the fracture of the tail happens in specific areas of the caudal vertebras which are naturally weakened. The autotomy may happen in two different ways: intravertebral autotomy, in which the vertebra at the centre of the tail have transversal fracture planes prepared to break if they are pressed hard enough, and intervertebral autotomy, where the tail breaks between vertebras by muscular constriction.

Tridimensional model of the fracture planes on the tail of a lizard and the regeneration post-autotomy of a cartilaginous tube. Image extracted from Joana D. C. G. de Amorim et al.

Caudal autotomy allows the animal to escape, but it isn’t without cost. Many reptiles use their tails as a reserve of fat and losing this energy store is usually detrimental for the animal. That’s why many lizards, once the threat has disappeared, look for their lost tail and eat it, to at least regain the energy it had as fat. In addition, regenerating a new tail requires a great expenditure of energy.

Photo of a Catalonian wall lizard (Podarcis liolepis) that has shed its tail. Photo by David López Bosch.

The regeneration of the tail in reptiles differs from that of amphibians and fishes in that it happens without the formation of a blastema and instead of an actual regeneration of the caudal vertebras, it forms a cartilaginous tube along it. The new tail is stiffer and shorter than the original one, and it usually regenerates whole some weeks after the amputation. Most lizards can regenerate their tails multiple times, but some species like the slow worm (Anguis fragilis) can only do it once. Sometimes, the original tail isn’t completely broken but the regeneration mechanisms are activated, which can lead to lizards and geckos with more than one tail.

Detail of the tail of a common wall gecko (Tarentola mauritanica) which has regenerated the tail without losing its original tail. Photo by Rafael Rodríguez.

Urodeles, the kings of regeneration

Of all tetrapods, amphibians are the ones that present the more astonishing regenerative capacities. During the larval stage of most species, both the tail and the limbs (if they have them) can be regenerated after its loss. The scientific community thinks that this is due to the fact that in amphibians the development of limbs and other organs is delayed until the moment of metamorphosis. Yet, frogs and toads (anurans) only maintain their regenerative powers during their tadpole stage, losing them when reaching adulthood.

Wood frog tadpole (Rana sylvatica) which, like all amphibians, delays the development of its legs up to the moment of metamorphosis. Photo by Brian Gratwicke.

Instead, many salamanders and newts (urodeles) conserve their regenerative powers their whole life. Even if many species present caudal autotomy, unlike lizards urodeles are able to completely regenerate, not only their tails, but practically any kind of lost body tissue. Of all known species, the axolotl (Ambystoma mexicanum), a neotenic amphibian which reaches adulthood without undergoing metamorphosis, has served as a model organism for the study of the formation of the blastema that precedes regeneration.

 Video about the axolotl, this curious amphibian which is greatly endangered. Video by Zoomin.TV Animals.

Regeneration as it happens in salamanders has stages genetically similar to the ones that occur during the development of the different body tissues and organs during the embryonic development of the rest of vertebrates. In the axolotl (and in the rest of urodeles) regeneration of a limb after amputation goes through three different stages:

• Wound healing: During the first hour after the amputation, epidermal cells migrate to the wound. The closing of the wound usually completes two hours later with the same mechanisms as in the rest of vertebrates. Yet, the complete regeneration of the skin is delayed up until the end of the regeneration.
• Dedifferentiation: This second phase, in which the blastema is formed, starts 24 hours after amputation. This is composed both of cells from the specialized tissues of the amputated zone which lose their characteristics (they obtain the capacity to proliferate and differentiate again) and cells derived from the connective tissue that migrate to the amputation zone. When these cells of different origins accumulate and form the blastema, the cellular proliferation starts.
• Remodelling: For the third stage to start, the formation of the blastema is required. Once the blastema is formed by different dedifferentiated cells, the formation of the new limb follows the same pattern as any kind of vertebrate follows during embryonic development (it even has de same genes intervening).

Diagram about the formation of the blastema in a zebrafish (Danio rerio) another model organism. Image from Kyle A. Gurley i Alejandro Sánchez Alvarado.

Recently fossils have been found from many different groups of primitive tetrapods which present signs of regeneration. Proof has also been found of limb regeneration in temnospondyl (Apateon, Micromelerpeton and Sclerocephalus) and lepospondyl (Microbrachis and Hyloplesion) fossils. This wide variety of basal tetrapod genera presenting regeneration and the fact that many fish also present it, has led many scientists to consider if the different groups of primitive tetrapods had the ability to regenerate, and if it was lost in the ancestors of amniotes (reptiles, birds and mammals).

Photo of an axolotl, by LoKiLeCh.

However, it is believed that the genetic information that forms the blastema could still be found in the DNA of amniotes but in a latent state. Of the three stages of the regeneration process, the only one exclusive to urodeles is the dedifferentiation stage, as the healing stage is the same as in the rest of vertebrates and the remodelling stage is like the one during embryogenesis. Currently many studies are being carried out on the way to reactivate the latent genes that promote the formation of the blastema in other vertebrates, such as humans.

Some human organs like the kidneys and the liver already have some degree of regenerative capacities, but thanks to investigation with stem cells in animals like salamanders and lizards currently it is able to regenerate fingers, toes, genitals and parts of the bladder, the heart and the lungs. As we have seen, the different animals able to regenerate amputated limbs hold the secret that could save thousands of people. Remember this the next time you hear that hundreds of species of amphibians and reptiles are endangered because of human beings.

References

During the writing of this entry the following sources have been consulted:

• Cover photo by Peter Halasz.
• Halliday & Adler (2007). La gran enciclopedia de los Anfibios y Reptiles. Editorial Libsa.
• http://www.eurostemcell.org/de/node/19896
• https://www.ncbi.nlm.nih.gov/pubmed/12455626
• https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3480082/
• http://ftp.eebweb.arizona.edu/courses/Ecol483_583/Bateman%20and%20Fleming%202009.pdf
• http://www.sciencemag.org/content/276/5309/81.short
• http://www.ambystoma.org/
• http://www.nature.com/nature/journal/v527/n7577/full/nature15397.html