The Diversity of Echinoderm Anuses!

This week, in the category of "one of those posts only Echinoblog could write"! Let us explore the vibrant diversity of echinoderm anal structures!! Perfectly SAFE for Work! even though it contains the word "anal" and "anuses" several times! WOO!!

Echinoderms are pentaradial (aka pentameral). That is they show a form of radial symmetry wherein their body always occurs around a top-down axis in five parts (although arms, etc. may vary).

As a consequence, the overall dynamic of their life mode is different from an animal with bilateral symmetry. No face, no "front" or "back". Thus, the way food moves through the body is different depending on where the mouth is located and consequently, where the anus is located!!

Since each living class of echinoderms has a fairly unique shape and body morphology, each group has a bunch of unusual specializations that function to facilitate the expulsion of poop!

To be honest, not every member of each class below has what is shown below. These are unusually prominent examples..but it still begs the question "What is it used for?"  How many other animals have so many unusual anal adpations??

5. Crinoids: Anal Chimneys & Pyramids
So, crinoids are suspension feeders. They almost kind of resemble plants. Most living ones are known as "feather stars" and are basically cups with arms for filtering water. But earlier forms of crinoids are known as "stalked crinoids" which have a stalk. I had a gallery of the older Paleozoic ones here a few weeks ago..

Here's a living one from the recent Okeanos Expedition to the Marianas region to give you an idea of what they look like..

So, the mouth in stalked crinoids is inside that cup at the base of where all those arms converge. The anus for these animals is ALSO in the cup. So, there's likely a strategy/adaptation for these animals to push the poop/excreta OUT of the anus, such at it does not end up getting "re-eaten" by the mouth...

According to Fossil Crinoids by Hess et al.  there were several fossil (Paleozoic) forms which had fairly straightforward strategies for dealing with ensuring that poop was discharged FAR from the mouth..

For example, here is the Mississippian Uperocrinus nashvillae with an explanatory diagram from Fig. 37 of Hess et al's book. Basically that huge pointed structure on top? That's called the anal tube  (or sometimes in other animals.. an anal pyramid or even an anal chimney!)

These extended structures serve to project the anus (and the excreted poop) well AWAY from the mouth (the feeding arms would come off right at that wide "ledge" around the center of the specimen..

There was a surprising diversity of these structures. Here's another one called Macrocrinus verneuilianus which is a fossil from the Carboniferous (the Paleozoic). The drawing below is taken from the Wooster Geologist blog and they point out that the elongate anal tube (in the lower left hand corner) aka the anal chimney may have even served an additional function beyond simply transporting poop away from the mouth:

The tube allowed waste products to be whisked away far from the mouth of the crinoid, which was at the base of the arms. Some researchers suggest that the long tube served another function as well: it may have helped stabilize and direct the filter-feeding fan of outstretched arms in a stiff current, something like the tail of an airplane or a panel on a weather vane.

One last weird crinoid is this one: Bicidiocrinus wetherbyi
Another Mississippian (i.e Paleozoic) stalked crinoid.. and this is kinda weird. So, there's the cup and the arms and that cone is the anal cone (=tube, pyramid, chimney, etc.) BUT it also has this weird additional protective "spiniferous canopy" around it!!

The diagram on the right shows this fully "reconstructed"..

                            

Image of the fossil from http://louisvillefossils.blogspot.com/2014/08/bicidiocrinus-wetherbyi-crinoid-calyx.html

and the reconstruction from Hess et al. Fossil Crinoids-which is a GREAT book for these useful facts!                              

Ultimately, there is probably a WHOLE blog post or 5 about Paleozoic crinoids and what we know about their paleoecology.. I briefly touched on the snails that parasitize their anuses here.... 

One LAST MINUTE ADDITION: David Clark (@clarkeocrinus) provides this ASTONISHING Proteriocrinus with a very considerable anal chimney!! which looks to extend nearly the length of the cup and arms!

@echinoblog A Poteriocrinus from the Dev Arkona Fm in Ont of the John & Mike Topor collection. Quite a chimney! pic.twitter.com/gHG76PiLR5

— David Clark (@Clarkeocrinus) April 26, 2016

4. Sea Urchins: (Diadematidae) Anal Sac
Probably one of the best known but most poorly recognized of the various echinoderm anal structures is the ANAL SAC in diadematid (diadematoid?) sea urchins.

This includes Diadema, Astropyga, Echinothrix and all of urchins in this family. Usually these are tropical and characterized by long, sharp spines AND a very distinctive "anal sac" present on the TOP of the body.

The problem is that many people see this big eye-shaped ball on the top of the sea urchin body and assume that it is an eye of some kind...

I can tell you most DEFINITIVELY that this structure is NOT the eye. I actually explained in great detail here how this was actually the unusual ANAL SAC which is characteristic of this type of sea urchin for an error in New Scientist.

That said, they DID publish a WONDERFUL picture by David Fleetham of Astropyga radiata venting poop OUT of the anal sac!!

Basically, this is a transparent or translucent bulb or sac extension from the anus through which feces passes on its way out of the body.

See those little round things that look like corn kernals? Sea urchin poop!

Here's another great pic found on Flickr, taken by Eunice Khoo of what looks like a small Astropyga sp. clearly showing poop THROUGH the transparent walls of its anal sac (aka anal cone).. 

Very nice capture. 

But why take my word for it??  Go ahead and watch it here in this video of what looks like either Diadema or Echinothrix..  The poop event takes place at about 0:15 into the video.
               
But its not JUST shallow-water diadematid urchins that have this anal sac. Here's a deep-water Aspidodiadema from the recent Hawaiian Okeanos expedition. I talk a bit about these here

These were videos of these animals from 2000-3000 METERS below the surface. Aspidodiadematid urchins are classified in the same general group as diadematids. and they too seem to have this anal sac or cone...

3. Ophiuroids: Alas..brittle stars got no anus. In fact, food goes in and goes out the SAME hole: the MOUTH. So..bummer.

2. Anal (or Epiproctal) Cone
Sea Stars/Starfish in a larger group called the Paxillosida (the mud and/or sand stars) have a specialized structure which sits right on the center of the disk called the anal or epiproctal cone.

A brief anatomical note- Although historically called an "anal cone" these starfish don't actually have a complete gut and so, the opening on the disk center is not actually the anus since it doesn't connect with the interestine. Hence the name "epiproctal":  EPI is Greek for "upon" and PROCT is Greek for "anus".. hence the cone or structure UPON the anus..

The cone is basically an outpocketing of the body, extending UPWARDS through the sediment

From (Fig. 2 from Shick 1976)

To quote myself from a few years ago:

Observations of Ctenodiscus under hypoxic conditions led to the illustration above. Basically, its thought that the cone gets more enlarged as hypoxia and hydrogen sulfide increases. The extension of the cone extends through the surface, with the tip at the surface. For your typical 6.0 cm diameter animal, these animals can have a cone that can attain 3 to 4 cm and extend 2 to 3 cm above the mud. It can leave this extended for over an hour. As the picture suggests, it can move around and push through sediment as the mud shifts, and etc. So, it can move around.

This also serves to make the top surface of the animal thinner, allowing easier gas exchange and opening up a channel to the surface water above the sediment surface!!

From Shick 1976 Marine Biology: http://link.springer.com/article/10.1007/BF00387613#page-1

So, in truth, the "anal cones" in these mud stars is NOT really an anus. Its where the anus would be located in other species..and so the labelling of this stucture is kind of a misnomer.. as they are not really used for defecation.

Other "mud stars" such as this deep-sea (3000-6000 m) Porcellanaster also have well-developed "anal cones". These starfish sit buried in sediment with these projections sitting up through the sediment. 

In contrast this shallower water Astropecten armatus has a much less developed anal or epiproctal cone. This is also related to the fact that it occurs in shallower water and tends to bury itself in somewhat less sediment compared to the above two species..

from CSU Fullerton: http://biology.fullerton.edu/biol317/ftm/ft_s15_cat_4_25_15.html

1. Sea Cucumbers (Holothuroidea): Anal Teeth
And finally, one of the best known of echinoderm anal defenses: the anal teeth in sea cucumbers!

I've reported on these before. There's at least one interpretation that these structures are defensive in nature and work to keep pearlfishes (and likely other commensals or parasites) from inhabiting the cloaca.

There's quite a few crabs and shrimps that live in and around sea cucumber anuses. See more here.

Sea cucumbers get kind of a special award for using their cloacal and other "ass end" chambers most efficiently..especially since some species can actually FEED and BREATH as water passes through the anal opening!! Read more about that here.

So there you have it! The Astonishing Anuses of the Echinoderm World! 

Ribbon Worms: The Incredible Predators You've Barely Heard About!

This week: Some cool examples of predation by a predator, you've probably NEVER heard of! And yet, they are deadly, efficient and can take down prey or food that is MANY times their body size.

I am of course talking about the Nemertea aka the Ribbon worms (or sometimes the proboscis worms)! This is actually a whole PHYLUM of worms, which in spite of their superficial appearance, are actually VERY different from flatworms and have a very significant number of internal features..

There's somewhere in the neighborhood of 1300 species of ribbon worms. Most are flattened and can be VERY elongate with some (e.g., Lineus longissimus) approaching 60 METERS (nearly 200 feet) long! Most occur in the oceans but there are freshwater and terrestrial species as we'll see.

Here's a nice summary of info from the Smithsonian magazine. 

Relevant to today's post is that many nemertean worms are PREDATORY.

They capture food using an eversible proboscis which, thanks to the internet's fascination with weird, gross stuff, you have undoubtedly seen in a variety of formats. Here's two of the popular ones: the red Thailand species expelling its bizarre, almost fractal proboscis:

This proboscis is one of the primary defining characteristics of this group.

and of course this green one from Taiwan!

According to an interview with Dr. Jon Norenburg in the Invertebrate Zoology Department of the NMNH, the proboscis of nemerteans is ejected from a body cavity and used to capture prey. Often times, they can capture prey that is THREE TO FOUR TIMES larger than their body!!

Here's the thing though, the above two video/gifs? Showed these animals in stressed situations outside of their natural environment. And NOT demonstrating their mastery of their predatory proboscis appeasing their ravenous rhyncocoel!

These animals likely represent a HUGE ecological impact which is probably difficult to study owing to their rather cryptic nature.

Here today is a nice little collection of ribbon worms "strutting their stuff" with some GREAT images showing attacking/devouring prey (thanks to YouTube and Flickr!)

Some ribbon worms such as these huge Antarctic ribbon worms (Parborlasia corrugatus) are both scavengers AND predators and have been reported as feeding on nearly ANYTHING:  fecal pellets, starfish, dead seal meat, fish, sponges, sea anemones, worms amphipods, penguin meat AND sardine meat with tomato sauce! and on and on....

5. POLYCHAETE WORMS! Polychaete worms of various types appear to be a popular food among ribbon worms.

Here's one image by James Zhan showing a large ribbon worm consuming a "paddle worm"

This one is impressive. You can actually watch the proboscis IN ACTION as its used to attack its polychaete prey!

Several ribbon worms eating a polychaete annelid from LabNemertea on Vimeo.

Here's another showing feeding on dead but still, LARGER polychaete worm food..

Here are pics AND video from Florida of the ribbon worm Tubulanus feeding on a feather duster worms (a filter feeding polychaete).  Here's a nice shot showing it attacking the "head"

Image from Wetpixel, photo by lindai http://wetpixel.com/forums/index.php?showtopic=51604

And a separate video showing another interaction with the same two players! 

4. CRUSTACEANS!! 
From two different sources. Marvyn Yeo shows us this ribbon worm seemingly coming up behind a tiny crab..

And this video shows us a ribbon worm feeding on this amphipod!

Ribbon worm (nemertean) eating an amphipod crustacean from LabNemertea on Vimeo.

and even mysid "shrimp"!

3. INSECTS and land arthropods
Most ribbon worms live in the ocean, but a sparing few have gotten into freshwater and even terrestrial habitats! And yes.. with those same predatory habits!

I'll be honest though, some of these can be pretty difficult to ID even to phylum. these were identified as ribbon worms but I suppose they could easily be flatworms. 

2. Snails! Flickr users Marvyn Yeo and "Paul" brings us stunning pictures from Asia and Ecuador showing this ribbon worm attacking this snail! Apparently by working its way into the shell aperature and attacking the animal in its own shell.

1. FISH!! and finally the pièce de ré·sis·tance! To be sure, I'm don't think that these worms actually captured these fish, BUT they are very clearly DEVOURING THEM.

Many ribbon worms have a stylet or thorn which is thought to deliver a toxin against some prey items.. so is it entirely unreasonable to suggest they capture fish, esp. prey much larger than them? Probably not. 

This one is from the Falkland Islands/Malvinas..

Here is another GREAT feeding observation in VIDEO with a professional commentary by ribbon worm expert Dr. Jon Norenburg at the No Bones blog! here  This one shows a ribbon worm devouring a HUGE fish from a reef in the Cayman Islands.

But good grief: LOOK AT THAT! Amazing. 

And of course leave it to the folks at Tokyo Sea Life Park to capture this AWESOME video of Parborlasia corrugatus An Antarctic ribbon worm devouring this mackeral in time lapse! (Although Parborlasia is pretty big relative to that fish-its still impressive to watch)

LOVECRAFTIAN FOOTNOTE

So, I am enough of an old Lovecraft/Cthulu "unnaturalist" to recognize the similarities between aforementioned nemerteans and Lovecraft's famous wormy Dholes! Images shown here from Fig. 8 (Petersen et al. 1988- Petersens' Field Guide to Cthulhu Monsters) on the left and on the right from YogBlogsoth.

The proboscis is close..but I'm thinking.. polychaete instead (dholes do seem to be segmented worms after all)? But that only suggests that if dholes are derived polychaetes than a giant Cthuloid Ribbon Worm may yet be lurking in the darkness....

fr http://yog-blogsoth.blogspot.com/2012/06/dholedoelbhole.html

Crinoid Fossil Forest Revisited!

This week I thought it would be fun to revisit the wonderful world of stalked crinoids!  I've previously looked at photo galleries of deep-sea living stalked crinoids here and a few from the Hawaiian Okeanos expedition.

Here's a hyocrinid "pinwheel" crinoid form the Okeanos dive to Indonesia! (INDEX-SATAL 2010)

For those who might not be familiar with stalked crinoids, they are the ancient ancestors of modern day feather stars (aka comatulid crinoids).

Stalked crinoids are fundamentally composed of three main regions: the calyx (or cup), arms and stalk which is very nicely illustrated by this diagram from the Field Museum in Chicago!

Stalked crinoids feed on food particles in the water column using their arms which they move down to the mouth located at the top of the calyx (or cup). The stalked and unstalked forms have an unusual relationship which you can read about in an earlier post here.

The high point of stalked crinoid diversity was in the Paleozoic, some 250 to 540 million years ago in the time before dinosaurs..there  existed a HUGE diversity of stalked crinoids.. and even DURING the time of the dinosaurs in the Mesozoic there were still quite a few of them (as I'll share below)..

Let's start off with this gorgeous one called Acanthocrinus rex! from the lower Devonian of Germany. This image was reported by crinoid scientist Hans Hess as "certainly one of the most beautiful crinoids ever found.." Sadly, this specimen was lost in World War II.

This image from: https://geo-ebooks.tumblr.com/post/127610816184/acanthocrinus-rex-j%C3%A4kel-from-the-lower-devonian

The images below are a nice set of Paleozoic crinoids by James St. John on Flickr..

Here is the Paleozoic Taxocrinus colletti from the Edwardsville Formation, Lower Mississippian; Crawfordsville area, Montgomery County, Indiana, USA.

A striking one called Platycrinites saffordi from the Edwardsville Formation, Lower Mississippian; Crawfordsville area, Montgomery County, Indiana, USA.

One with some strikingly different arm branching patterns: Onychocrinus exculptus from the Edwardsville Formation, Lower Mississippian; Crawfordsville area, Montgomery County, Indiana, USA.

Here is: Onychocrinus ulrichi a fossil crinoid from the Edwardsville Formation, Lower Mississippian; Crawfordsville area, Montgomery County, Indiana, USA.

David Clark on Twitter brings us this interesting one with large spines emerging from the calyx! The aptly named Acanthocrinus! (from the Devonian of New York)

Christmas star theme for #FossilFriday ? Acanthocrinus spinosus from the Devonian Windom shale of New York. pic.twitter.com/KejobiKyJR

— David Clark (@Clarkeocrinus) December 18, 2015

Still searching for info about my Acanthocrinus spinosus from Dev. Windom shale in NY. #crinoid #fossil pic.twitter.com/CdWOSMWt

— David Clark (@Clarkeocrinus) May 2, 2012 

This next series is from Elrina753 (thanks for the awesome images!) from the Houston Museum of Natural Science. These look like the Mississippian, Platycrinites

Another gorgeous one from the Houston Museum of Natural Science: Onychocrinus exsculptus

And this third one from Houston which I don't have a name for...

An interesting Paleozoic one called Eretmocrinus (no other info, so unsure if ID is correct) but the arms have very unusual morphology. Notice how they become paddle like towards the tips! 

And from the displays/collections in Washington DC at the NMNH.. the massive Jurassic Seirocrinus which was actually thought to be pelagic and might have attached to floating logs! More on these two unusual swimming/floating crinoids here.

The bizarre Uintacrinus which some thought might have actually dragged its arms over the bottom as as it floated by...