PART 1
They are here included many pictures published by the
kindness of his authors to which I forward my more sincere thanks. Especially
to M. Verolet, whose generosity has provide many of the most interesting
pictures shown here. I have also
borrowed drawings of some sites of Internet for which I give the needed notice.
The remaining illustrations were taken (in these times of digital cameras with
5 to 10 Mpx) with a camera of 0.4 Mpx.; certainly they are treated vastly by
PhotoPaint, NetImage Demo, and ACDSee.
I hope that, at least, they would be clear and convey the
information that they are intended to give.
INTRODUCTION
In traditional taxonomy Rotifers are considered a Phyllum
which embraces three Classes: the “Seisonacea”, the “Monogononta”,
and the “Bdelloidea”. Seisonacea
are only marine, epizoïques on Nebalia,
(a genus of crustaceans considered very primitive, benthonic, with some
littoral species), and are represented by only one genus, Seison, with three species, the last
one published in 2007, which have a morphology very distant from that normally
assigned to a rotifer. Monogononta, thus called because
male and female have only one genital gland, were vastly treated by Michele
Verolet, which has even presented in the French Magazine Microscopies a
very complete and much illustrated description of its morphology, and a
splendid key to identify all the genera that the amateur has the possibility of
finding with some frequency in its explorations.
http://www.microscopies.com/DOSSIERS/Magazine/Articles/M%20Verolet-CLE/presentation/presentation.htm
Bdelloidea are all parthenogenetic females, with two genital glands. (During a long
time they were called Digononta).
Monogononta and Seisonacea
have males. But Bdelloids do not have them ....... probably since 80 million
years (http://news.bbc.co.uk/1/hi/sci/tech/7039478.htm)
Its reproduction is
exclusively parthenogenetic, and the genetists break their heads to discover
the mechanisms which made it possible for these animals to maintain its genetic
diversity avoiding to be banned from the list of the living species. Some recent
papers show that they steal genes from other species, to modify her own genome.
Of all the Rotifers the Bdelloidea are the best known by the non
specialized students. The cephalic end of the majority has a “corona” divided
in two retractile “trochal discs”, with ciliated margins. The Cilia in the
“trochas”, that beats metachronically, and seems to rotate, gave the name to
these animals. In fact the trochas are restricted to 15 of the 19 genres of
this Class, but they are so showy that the not informed identifies not only the
Class, but all the Phyllum Rotifera, with the bdeloids.
2
- Magnificent frontal view of the “head” of a Philodina (courtesy of Charles
Krebs). Click on the picture to see a labeled version
2b - Annotations of the author: 1 lines of cils implanted in the trochal edge
(2), 3 inferior lip or Cíngulum, 4 lip of the buccal funnel, 5, buccal funnel,
and 6 sulcus (a furrow between Trocha and Cíngulum) These ciliated fields help
to send the food to the mouth.
They are multicellular organisms with a fairly
complex organization, but its size is in a scale similar to that of the big
protozooa. (Between 150 and 700 microns. Only one species, Rotaria neptunia,
she is excessively long and thin, reaching a measure of 1600 microns. (Ricci and
Melone, 2000)
Practically all the species can endure drying
and can revive when they happens to be submerged in water another time.
Jacobs published in 1909, for the first time,
the description of this amazing capacity of the rotifers, which gives them the
possibility of invading really difficult habitats, like desecables mosses, the
cracks in the crusts of the trees, the ground, and so on.
The rotiferologist Aydn Örstan was able to
find bdeloids in the wall of the tubes of thermites, adhered to a tree in
Puerto Rico, and even to describe a new species from the dust on the soil of a
dessert in México.
Christian Colin experimentally registered the
images of the drying process and the reviviscence of a bdelloid. The article
that describes his technique, process, and results can be read at:
http ://www.microscopies.com/DOSSIERS/Magazine/Articles/CC-PHENIX/Phenix-1.htm
and its second part
http ://www.microscopies.com/DOSSIERS/Magazine/Articles/CC-PHENIX/Phenix-2.htm
another
shorter graphic document on the same phenomenon written by Hugo Baillie-Johnson
can be found al http://www.microscopy-uk.org.uk/mag/artnov02/hbjtrehalose.html
3 - Reviviscencia, after 46 days of anhydrobiosis.
Registered by Christian Colin
In the state of anhydrobiosis (or anabiosis)
they resemble small dust particles and, like them, they could be dispersed
easily by the wind, or the waters that bathe the surfaces where they are.
Notwithstanding there are in fact few registered data on the real (and surely
multiple) methods that the rotifers use, and which have given to many species
the ability for them to become cosmopolitan.
The anydrobiose is not a simple adaptation to
difficult to colonize habitats. As Ricci et all. says (August 2007)
Bdelloids, although aquatic animals, are not only
efficient in tolerating desiccation, but seem somehow dependent on
anhydrobiosis, a circumstance that might represent a key event in their life
cycle. If this is true, life in unpredictable habitats should not be seen as
the result of competitive exclusion from 'easier' habitats, but a requirement
for long-term survival of these parthenogenetic animals”
The name “bdeloids” comes from the Greek and
it talks about the characteristic way of crawling in a leech-like manner,
fixing alternatelly the terminal toes, extending, fixing the rostrum, and
retracting the previously fixed end.
Also in this case all the bdeloids do not show
always the same behavior. For example a rotifer of the genre Adineta
moves normally gliding over surfaces, using the cils that cover the ventral
side of their cephalic end. (see picture xx)
Even the cephalic end, wich gives his name to
the Class, does not have a homogeneous structure, and there exist at least 3
different structures, which allow to divide the class in 3 Orders: Adinetida,
Philodinida and Phylodinavida, and will be treated in detail when
characterizing them.
As
in the Monogononta (See Verolet
http://www.microscopies.com/DOSSIERS/Magazine/Articles/M%20Verolet-Les%20Rotiferes/texte.htm)
the body is compossed of 3 basic portions: the
head, the trunk and the foot. The epidermis that covers it is a sincitial layer
(that is to say, a layer of cytoplasm in which separated cells do not exist,
although these are represented by big dispersed nuclei) is marked by
cross-sectional furrows that separate superficial ring (pseudosegments, since
they do not divide the interior of the body), and that gives to the rotifer the
capacity to be contracted in a “telescopic” way. This is possible thanks to the
existence of a pliable layer of intracytoplasmic cuticule, denominated
“lorica”.
In many Monogononta this lorica is hard and
indeformable, with important specific characteristics. (See Verolet) But in the
Bdelloidea it is almost always thin and folding, although in some species they
can have characteristic thickenings denominated “Cuticular Sculptures” (plates,
furrows, spines, warts, a.s.o).
The “corona”, when it exists, can be closed
inside the head, and this, and the foot at the other end, are contracted within
the trunk, acquiring a compact and rounded form. This form (named “tun” in
English) is the one that they adopt so much as a strategy of defense in any
case of a sudden aggression, or even as a previous step to drying. (see the
above experiment of Christian Colin)
4
- The immediate total retraction of a Philodina as a reaction to an abrupt
contact, like a stroke over the coverslip
It is common that
the head joint to the trunk by a narrowed portion called the “neck” Normally a
sensitive antenna, thin, and more or less long, lodges dorsally in the “neck”.
5 - Lateral view of a Philodina sp. , showing the dorsal antenna.
Below the antenna can be seen the red eyes . The pseudosegmentation and the
ridged trunk cuticle is also shown.
In addition, between the “trochas” it can be seen dorsally
the “Rostrum”, one generally robust structure provided at its end with
cils, sticky cells and sensitive lamellas.
It is retractable in most of the species.
Some species have eyes, which lodge sometimes in the anterior end
of the rostrum (Rotaria), and other times dorsally in the neck, over the brain
(Phillodina). In Adineta oculata they
are described rostral eyes composed of a red spot surmounted by a concave lens.
6 - Dorsal eyes over the brain
in Mniobia. They are pigmented of red.
7 - Rotaria sp., dorsal view
of the head, with dark eyes in the rostrum
8 - Relation between the
antenna and the rostrum. Trochas semiretracted. One specimen with the “Corona”
totally retracted is seen in picture 27
The mouth opens centrally between the “coronas” (see fig 2) at the end
of a buccal funnel, which could be followed by a buccal tube that communicates
with a masticatory organ named mastax.
The length of the buccal funnel and the buccal tube, and the depth of
the mastax is a character that depends on the style of alimentation.
9 - The deep mastax of a Habrotrochidae
You can see a mastax very near the mouth at fig. 30 (Henoceros)
The Mastax is a complex organ formed by strong muscles armed with
several hard, cuticularized pieces, named “trophi”. Its structure is
characteristic of all the class and it really has little variety. The type of
trophi of the bdelloidea is called “ramate”. The ramate trophi lacks the
fulcrum, one unpaired piece characteristic of the remaining types of trophi of
the Phyllum. (Ver Verolet) The trophi consist of 6 pieces, 2 unci (sing.
uncus), 2 manubria (sing. manubrium) and 2 rami (sing. ramus). The plate-like
Unci are armed of teeth, differentiated in 3 groups. The anterior and posterior
ones, are formed by thin teeths. There is a group (generally median) that has
much more heavy teeths. According to the genre and the species these heavy
teeths can be in numbers from 2 to 10.
10 - Typical image of the trophi of the order Philodinida, courtesy of
M.
Verolet.
It can be compared with this image taken with electron microscope, and
colored to differentiate the different pieces. In red the manubria, in blue the
unci and in green the rami, underneath the indented edge of the unci
11- MEG image of the ramate
trophy electronically stained, due to G.Melone.
12 – Something atypical Mastax of
a Philodinavida from Cancún (5 images
composed with combineZ 5.0)
Moreover the trunk lodges the remaining organs in its interior: brain,
sub cerebral and retrocerebral organs, mastax, salivary glands, digestive and
excretory systems, gonads, and muscles.
13a – Mastax, salivary,
glands and intestine with heavily ciliated lumen in a bdeloid. Courtesy of M.
Verolet.
The mastax is followed by a normally very short esophagus, in most of
the cases difficult to see, which opens in the sac-like, stretched, stomach.
Normally the stomach is syncitial, with a ciliate cavity (mostly a tube) which
is followed by a short, mostly bulbous, intestine (some times called “rectum”).
But in the Habrotrochidae the
stomach has no lumen at all and food is worked out in little balls, and
included in vacuoles that give to the stomach a foamy appearance.
13b - The stomach of an Habrotrochidae
Behind the rectum, or underneath, we can see the urinary bladder, which
gathers the liquid retired from the pseudocoel by two long protonephridia
culminated near the neck by some flame bulbs. Blader and rectum both finish in
a common “cloaca” that opens to the outside dorsally in the base of the trunk.
The reproductive apparatus is represented by two germovitellaria, organs
that produce eggs, and at the same time provide it with the vitelo (nutritive
material) necessary for its development.
14 - Germovitellaria
at both sides of the intestine, in the trunk of
Adineta sp.
Most of the bdelloidea are ovipares, 15 - A finished
Egg, inside a bdelloid
But there exists some cases of viviparity. And some times the already
formed embryos, even mobile ones, can be seen within the mother. Even if the
development of the embryo is not advanced, viviparity can be denounced by the
fact that the egg is seen as pluricelular.
16a - Segmented egg (embryo) inside Rotaria sp. Note the
abscence of a Shell and the irregular surface.
16b- embryo and another segmented
egg in Rotaria sp.
The segments after the anus forms the retractable foot, at whose end 2
spurs can be normally seen, and behind these ones, a short structure lodge the
toes, connected to glands, visible in the last portion of the foot, which produce sticky substances, allowing the
rotifer to adhere to the substrate.
In a few cases (Bradiscella, Mniobia, and some epizoics) these toes are
replaced by a sticky disc.
17a – spurs in the foot of Mniobia, note also the big egg
17b – adhesive disc at the end of the foot in Mniobia
17c – toes at the end of the foot of Rotaria (courtoysie of Bertrand
Parres)
I add
here 3 tables with the list of the known genera, ordered under three different
criteria. This can be redundant, but I think that it is an easy form to convey
very interesting information
LIST OF GENERA IN
ALPHABETICAL ORDER (in blue, epizoic genera) Genus, author, publication
date, aproximative species numbers
Abrochtha
|
Bryce
|
1910
|
2
|
Adineta
|
Hudson & Gosse
|
1886
|
12
|
Anomopus
|
Piovanelli
|
1903
|
2
|
Bradyscela
|
Bryce
|
1910
|
2
|
Ceratotrocha
|
Bryce
|
1910
|
4
|
Didymodactylos
|
Milne
|
1916
|
1
|
Dissotrocha
|
Bryce
|
1910
|
7
|
Embata
|
Bryce
|
1910
|
5
|
Habrotrocha
|
Bryce
|
1910
|
100
|
Henoceros
|
Milne
|
1916
|
2
|
Macrotrachela
|
Milne
|
1886
|
100
|
Mniobia
|
Bryce
|
1910
|
50
|
Otostephanos
|
Milne
|
1916
|
9
|
Philodina
|
Ehrenberg
|
1830
|
40
|
Philodinavus
|
Harring
|
1913
|
1
|
Pleuretra
|
Bryce
|
1910
|
14
|
Rotaria
|
Scopoli
|
1777
|
24
|
Scepanotrocha
|
Bryce
|
1910
|
9
|
Zelinkiella
|
Harring
|
1913
|
1
|
A still mentioned (especially
in text books) genus (Callidina) is not valid now, it was very
heterogenous and the described species have been distributed, between many of
the now accepted genera.
2) Genera ranked by number of species
Habrotrocha
|
Bryce
|
1910
|
100
|
Macrotrachela
|
Milne
|
1886
|
100
|
Mniobia
|
Bryce
|
1910
|
50
|
Philodina
|
Ehrenberg
|
1830
|
40
|
Rotaria
|
Scopoli
|
1777
|
24
|
Pleuretra
|
Bryce
|
1910
|
14
|
Adineta+
|
Hudson & Gosse
|
1886
|
12
|
Otostephanos
|
Milne
|
1916
|
9
|
Scepanotrocha
|
Bryce
|
1910
|
9
|
Dissotrocha
|
Bryce
|
1910
|
7
|
Embata
|
Bryce
|
1910
|
5
|
Ceratotrocha
|
Bryce
|
1910
|
4
|
Abrochtha
|
Bryce
|
1910
|
2
|
Anomopus
|
Piovanelli
|
1903
|
2
|
Bradyscela
|
Bryce
|
1910
|
2
|
Henoceros
|
Milne
|
1916
|
2
|
Didymodactylos
|
Milne
|
1916
|
1
|
Philodinavus
|
Harring
|
1913
|
2
|
Zelinkiella
|
Harring
|
1913
|
1
|
3) Genera ordered by the publication year
Rotaria
|
Scopoli
|
1777
|
24
|
Philodina
|
Ehrenberg
|
1830
|
40
|
Macrotrachela
|
Milne
|
1886
|
100
|
Adineta
|
Hudson & Gosse
|
1886
|
12
|
Anomopus
|
Piovanelli
|
1903
|
2
|
Habrotrocha
|
Bryce
|
1910
|
100
|
Mniobia
|
Bryce
|
1910
|
50
|
Pleuretra
|
Bryce
|
1910
|
14
|
Scepanotrocha
|
Bryce
|
1910
|
9
|
Dissotrocha
|
Bryce
|
1910
|
7
|
Embata
|
Bryce
|
1910
|
5
|
Ceratotrocha
|
Bryce
|
1910
|
4
|
Abrochtha
|
Bryce
|
1910
|
2
|
Bradyscela
|
Bryce
|
1910
|
2
|
Philodinavus
|
Harring
|
1913
|
1
|
Zelinkiella
|
Harring
|
1913
|
1
|
Otostephanos
|
Milne
|
1916
|
9
|
Henoceros
|
Milne
|
1916
|
2
|
Didymodactylos
|
Milne
|
1916
|
1
|
No new genera has been described in the last 92 years
|