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| {{FosterSedgwick1885b header}} | | {{FosterSedgwick1885b header}} |
| ==CHAPTER XX. ECHINODERMATA== | | ==Chapter XX. Echinodermata== |
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| THE development of the Echinodermata naturally falls into | | THE development of the Echinodermata naturally falls into |
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| and the metamorphosis. | | and the metamorphosis. |
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| The Development of the Germinal Layers and of tJie Systems | | The Development of the Germinal Layers and of the Systems of Organs. |
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| of Organs. | |
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| The development of the systems of organs presents no very | | The development of the systems of organs presents no very |
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| An invagination now makes its appearance at the lower | | An invagination now makes its appearance at the lower |
| pole (fig. 247 B), and simultaneously there become budded off | | pole (fig. 247 B), and simultaneously there become budded off |
| from tJie cells undergoing the invagination amoeboid cells, which | | from the cells undergoing the invagination amoeboid cells, which eventually form the muscular system and the connective tissue. |
| | These cells very probably have a bilaterally symmetrical origin. |
| | This stage represents the gastrula stage which is common to all |
| | Echinoderms. The invaginated sack is the archenteron. As it |
| | grows larger one side of the embryo becomes flattened, and the |
| | other more convex. On the flattened side a fresh invagination arises, the opening of which forms the permanent mouth, the |
| | opening of the first invagination remaining as the permanent |
| | anus (fig. 248 A). |
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| 1 The following classification of the Echinodermata is employed in this chapter. | | 1 The following classification of the Echinodermata is employed in this chapter. |
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| ECHINODERMATA. 545
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| eventually form the muscular system and the connective tissue.
| | FIG. 247. TWO STAGES IN THE DEVELOPMENT OF HOLOTHURIA TUBULOSA VIEWED IN OPTICAL SECTION. (After Selenka.) |
| These cells very probably have a bilaterally symmetrical origin.
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| This stage represents the gastrula stage which is common to all
| |
| Echinoderms. The invaginated sack is the archenteron. As it
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| grows larger one side of the embryo becomes flattened, and the
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| other more convex. On the flattened side a fresh invagination
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| FIG. 247. TWO STAGES IN THE DEVELOPMENT OF HOLOTHURIA TUBULOSA | |
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| VIEWED IN OPTICAL SECTION. (After Selenka.)
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| A. Blastosphere stage at the close of segmentation. B. Gastrula stage. | | A. Blastosphere stage at the close of segmentation. B. Gastrula stage. |
| mr. micropyle ; //. chorion; s.c. segmentation cavity; bl. blastoderm; ep. epiblast; | | mr. micropyle ; //. chorion; s.c. segmentation cavity; bl. blastoderm; ep. epiblast; |
| hy. hypoblast; ms. amoeboid cells derived from hypoblast ; a.e. archenteron. | | hy. hypoblast; ms. amoeboid cells derived from hypoblast ; a.e. archenteron. |
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| arises, the opening of which forms the permanent mouth, the
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| opening of the first invagination remaining as the permanent
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| anus (fig. 248 A).
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| These changes give us the means of attaching definite names | | These changes give us the means of attaching definite names |
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| (566) Sir Wyville Thomson. "On the Embryogeny of Antedon rosaceus." | | (566) Sir Wyville Thomson. "On the Embryogeny of Antedon rosaceus." |
| Phil. Trans. 1865. | | Phil. Trans. 1865. |
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| ==CHAPTER XXI. ENTEROPNEUSTA==
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| THE larva of Balanoglossus is known as Tornaria. The prselarval development is not known, and the youngest stage (fig.
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| 272) so far described (Gotte, No. 569) has
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| many remarkable points of resemblance to
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| a young Bipinnaria.
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| A mouth (m\ situated on the ventral
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| surface, leads into an alimentary canal with
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| a terminal anus (an). A prae-oral lobe is
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| well developed, as in Bipinnaria, but there
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| is no post-anal lobe. The bands of cilia
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| have the same general form as in Bipinnaria. There is a prae-oral band, and a
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| longitudinal post-oral band ; and the two
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| bands nearly meet at the apex of the praeoral lobe (fig. 273). A contractile band
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| an
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| FIG. 272. EARLY
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| STAGE IN THE DEVELOPMENT OF TORNARIA.
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| (After Gotte.)
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| W. so-called watervascular vesicle developing as an outgrowth
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| of the mesenteron; m.
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| passes from the oesophagus to the apex of mouth; an. anus,
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| the prae-oral lobe, and a diverticulum (fig. 272, W) from the
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| alimentary tract, directed towards the dorsal surface, is present.
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| Contractile cells are scattered in the space between the body
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| wall and the gut.
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| In the following stage (fig. 274 A) a conspicuous transverse
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| post-oral band of a single row of long cilia is formed, and the
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| original bands become more sinuous. The alimentary diverticulum of the last stage becomes an independent vesicle opening
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| by a pore on the dorsal surface (fig. 274 A, w). The contractile
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| cord is now inserted on this vesicle. Where this cord joins the
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| apex of the prae-oral lobe between the two anterior bands of
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| cilia a thickening of the epiblast (? a ganglion) has become
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| 372
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| 580
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| ENTEROPNEUSTA.
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| C.C.
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| an.
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| FIG. 273. YOUNG TORNARIA.
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| (After Miiller.)
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| m. mouth ; an. anus ; w. watervascular vesicle ; oc. eye-spots ; c.c.
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| contractile cord.
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| established, and on it are placed
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| two eye-spots (fig. 273 oc, and
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| fig. 274 A). A deep bay is
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| formed on the ventral surface of
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| the larva.
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| As the larva grows older the
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| original bands of cilia become
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| more sinuous, and a second
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| transverse band with small cilia
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| is formed (in the Mediterranean
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| larva) between the previous
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| transverse band and the anus.
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| The water-vascular vesicle is
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| prolonged into two spurs, one
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| on each side of the stomach.
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| A pulsating vesicle or heart is
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| also formed (fig. 274 B, ht), and arises, according to Spcngel
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| (No. 572), as a thickening of the epidermis.
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| It subsequently becomes enveloped in a
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| pericardium, and is
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| placed in a depression
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| in the water-vascular
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| vesicle. Two pairs of
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| diverticula, one behind
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| the other, grow out
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| (Agassiz, No. 568) from
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| the gastric region of
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| the alimentary canal.
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| The two parts of each
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| pair form flattened
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| compartments, which
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| together give rise to a
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| complete investment of
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| the adjoining parts of
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| the alimentary tract.
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| The two parts of each
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| coalesce, and thus form
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| FlG. 274. TWO STAGKS IN THK 1 >KY KI.< >I'M KN I
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| OF TORNARIA. (After Metschnikoff.)
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| The black lines represent the ciliated hands.
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| m. mouth; an. anus; br. branchial cleft; ///.
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| heart ; c. Ixxly cavity between splanchnic and
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| somatic mesoblast layers; 7.-'. watcr-vascvilar vesicle:
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| v. circular blood-vessel.
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| ENTEROPNEUSTA.
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| 5 8l
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| a double-walled cylinder round the alimentary tract, but their
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| cavities remain separated by a dorsal and ventral septum.
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| Eventually (Spengel) the cavity of the anterior cylinder
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| forms the section of the body cavity in the collar of the adult,
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| and that of the posterior (fig. 274 B, c) the remainder of the
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| body cavity. The septa, separating the two halves of each,
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| remain as dorsal and ventral mesenteries.
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| The conversion of Tornaria (fig. 274 A) into Balanoglossus
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| (fig. 274 B) is effected in a few hours, and consists mainly in
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| certain changes in configuration, and in the disappearance of
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| the longitudinal ciliated band.
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| The body of the young Balanoglossus (fig. 274 B) is divided
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| into three regions (i) the proboscidian region, (2) the collar,
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| (3) the trunk proper. The proboscidian region is formed by the
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| elongation of the prae-oral lobe into an oval body with the eyespots at its extremity, and provided with strong longitudinal
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| muscles. The heart (hi) and water-vascular vesicle lie near its
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| base, but the contractile cord connected with the latter is no longer
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| present. The mouth is placed on
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| the ventral side at the base of the
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| prae-oral lobe, and immediately behind it is the collar. The remainder
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| of the body is more or less conical,
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| and is still girt with the larval
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| transverse ciliated band, which lies
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| in the middle of the gastric region
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| in the Mediterranean species, but
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| in the cesophageal region in the
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| American one.
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| The whole of the body, including
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| the proboscis, becomes richly ciliated.
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| One of the most important cha- S us WITH FOUR BRANCHIAL
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| racters of the adult Balanoglossus CLEFTS * (After Alex. Agossiz.)
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| r . m. mouth ; an. anus ; br. bran
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| consists in the presence of respira- chial cleft . hL heart ; IV. watertory structures comparable with the vascular vesicle,
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| vertebrate gill slits. The earliest traces of these structures
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| are distinctly formed while the larva is still in the Tornaria
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| FIG. 275. LATE STAGE IN THE
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| DEVELOPMENT OF BALANOGLOS
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| 582 I'N I'KUOl'NKUSTA.
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| condition, as one pair of pouches from the oesophagus in the
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| Mediterranean species, and four pairs in the American one
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| (fig. 275, br).
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| In the Mediterranean Tornaria the two pouches meet the
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| skin dorsally, and in the young Balanoglossus (fig. 274 B, br)
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| acquire an external opening on the dorsal side. In the American
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| species the first four pouches are without external openings
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| till additional pouches have been formed. Fresh gill pouches
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| continue to be formed both in the American and probably
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| the Mediterranean species, but the conversion of the simple
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| pouches into the complicated gill structure of the adult
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| has only been studied by Agassiz (No. 568) in the American
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| species. It would seem in the first place that the structure of
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| the adult gill slits is much less complicated in the American than
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| in the Mediterranean species. The simple pouches of the young
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| become fairly numerous. They are at first circular ; they then
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| become elliptical, and the dorsal wall of each slit becomes folded ;
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| subsequently fresh folds are formed which greatly increase the
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| complexity of the gills. The external openings are not acquired
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| till comparatively late.
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| Our knowledge of the development of the internal organs, mainly
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| derived from Agassiz, is still imperfect. The vascular system appears early
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| in the form of a dorsal and a ventral vessel, both pointed, and apparently
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| ending blindly at their two extremities. The two spurs of the water-vascular
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| vesicle, which in the Tornaria stage rested upon the stomach, now grow
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| round the oesophagus, and form an anterior vascular ring, which Agassiz
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| describes as becoming connected with the heart, though it still communicates
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| with the exterior by the dorsal pore and seems to become connected with the
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| remainder of the vascular system. According to Spengel (No. 572) the
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| dorsal vessel becomes connected with the heart, which remains through life
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| in the proboscis : the cavity of the water-vascular vesicle forms the cavity of
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| the proboscis in the adult, and its pore remains as a dorsal (not, as usually
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| stated, ventral) pore leading to the exterior.
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| The eye-spots disappear.
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| Tornaria is a very interesting larval form, since it is intermediate in structure between the larva of an Echinoderm and
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| trochosphere type common to the Mollusca, Chxtopoda, etc.
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| The shape of the body especially the form of the ventral
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| depression, the character of the longitudinal ciliated band, the
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| structure and derivation of the water-vascular vesicle, and the
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| ENTEROPNEUSTA. 583
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| formation of the walls of the body cavity as gastric diverticula,
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| are all characters which point to a connection with Echinodcrm
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| larvae.
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| On the other hand the eye-spots at the end of the prae-oral
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| lobe 1 , the contractile band passing from the oesophagus to the
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| eye-spots (fig. 273), the two posterior bands of cilia, and the
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| terminal anus are all trochosphere characters.
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| The persistence of the prae-oral lobe as the proboscis is
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| interesting, as tending to shew that Balanoglossus is the surviving representative of a primitive group.
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| *
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| BIBLIOGRAPHY.
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| (567) A. Agassiz. "Tornaria." Ann. Lyceum Nat. Hist.\u\. New York,
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| 1866.
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| (568) A. Agassiz. "The History of Balanoglossus and Tornaria." Mem.
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| Amer. Acad. of Arts and Stien., Vol. IX. 1873.
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| (569) A. Gotte. " Entwicklangsgeschichte d. Comatula Mediterranea." Archiv
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| fur mikr. Anat., Bd. xii., 1876, p. 641.
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| (570) E. Metschnikoff. " Untersuchungen iib d. Metamorphose, etc. (Tornaria)." Zeit.fiir wiss. ZooL, Bd. xx. 1870.
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| (571) J. M tiller. " Ueb. d. Larven u. Metamor. d. Echinodermen." Berlin
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| Akad., 1849 and 1850.
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| (572) J. W. Spengel. "Ban u. Entwicklung von Balanoglossus. Tagebl. d.
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| Naturf. Vers. Miinchen, 1877.
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| 1 It would be interesting to have further information about the fate of the thickening of epiblast in the vicinity of the eye-spots. The thickening should by rights be the
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| supra-oesophageal ganglion, and it does not seem absolutely impossible that it may give
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| rise to the dorso-median cord in the region of the collar, which constitutes, according
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| to Spengel, the main ganglion of the adult.
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| INDEX.
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| Abdominalia, 459, 493, 499
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| Acanthocephala, 379
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| Acanthosoma, 473, 474, 475
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| Acarina, 444, 454
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| Accipenser, 102
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| Achaeta, 319
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| Achelia, 538
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| Achtheres percarum, 490
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| Acineta, 7, 8
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| Acraspeda, 152, 165, 167, 178, 179, 182,
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| 185, 186
| |
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| Actinia, 169, 171, 179
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| Actinophrys, 9
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| Actinotrocha, 315, 318, 363, 364
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| Actinozoa, 26, 102, 152, j66, 170, 171,
| |
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| 172, 176, 178, 179, 181, 182, 186
| |
| Actinula, 155
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| Aculeata, 421
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| ^Egineta flavescens, 158
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| yEginidae, 156, 158
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| ^Eginopsis Mediterranea, 158
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| /Equorea Mitrocoma, 182
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| Agalma, 163
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| Agelena, 436, 450
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| Agelena labyrinthica, 119, 438
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| Alciope, 74
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| Alcippidae, 499
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| Alcyonaria, 152
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| Alcyonidse, 167, 168
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| Alcyonidium mytili, 297, 300, 302
| |
| Alcyonium palmatum, 119, 148, 167, 182
| |
| Alima, 484, 486
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| Amoeba, 19, 20
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| Amphibia, 22, 54, 56, 59, 60, 63, 66, 74,
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| 83, 102
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| Amphilina, 218
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| Amphioxus, 54, 56, 59, 61, 66, 93, 426
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| Amphipoda, 518
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| Amphiporus lactifloreus, 202
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| Amphistomum, 31
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| ,, subclavatum, 205
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| Amphitrochae, 330
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| Amphiura squamata, 565
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| Anchorella, 108, 492, 520
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| Anelasma squalicola, 499
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| Anguillulidse, 371
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| Annelida, 14, 25, 98, 503, 525
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| Anodon, 37, 38, 39, 100, 107, 259, 260,
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| 265, 266, 268
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| Anopla, 189, 202
| |
| Anura, 5
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| | |
| Antedon, 568, 573, 574
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| Aphides, 15, 16, 76, 79, 116, 428, 429
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| Aphrodite, 42
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| | |
| Apis, 402, 407, 408, 412, 413
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| Aplysia, 99, 226, 238, 252, 253
| |
| Aplysinidaa, 146
| |
| Apoda, 459, 493
| |
| Aptera, 395, 420
| |
| Apus, 1 6, 79, 460, 463
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| Arachnida, 22, 114, 119, 413, 4.51, 435,
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| 444, 454, 455, 458, 537, 539
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| Arachnitis, 171
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| Araneina, 50, 51, 436
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| Arbacia, 567
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| Area, 38
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| Archigetes, 218
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| Archizosea gigas, 494
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| Arenicola, 42
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| Argiope, 311, 312, 315, 317
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| Argonauta, 247, 248
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| Argulus, 492
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| Armata, 355
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| Arthropoda, 12, 16, 18, 22, 75, 77,79, 83,
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| 108, no, 221, 382, 383, 434, 448,503,
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| 5 2 5> 534 54', 54 2
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| Articulata, 311, 313, 316, 317
| |
| Ascaridiae, 371
| |
| Ascaris nigrovenosa, 16, 82
| |
| | |
| ,, lumbricoides, 375
| |
| Ascetta, 144
| |
| Ascidia canina, 53
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| Ascidians, 74, 102, 208, 426
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| Asellus aquaticus, 112,120, 516
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| Astacus, 66, 465, 477, 511, 512, 513,
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| 525
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| 586
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| INDKX.
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| Asteracanthion, 69, 70, 561
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| Asterias, 20, 68, 69, 71, 78, 80, 84, 549,
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| 564
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| Asteroidea, 35, 36, 544, 549, 557, 563,
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| | |
| 576
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| Astnea, 169
| |
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| Astroides, 169
| |
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| Atax Bonzi, 445
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| Atlanta, 231, 240
| |
| | |
| Atrochae, 330
| |
| | |
| Aurelia, 167
| |
| | |
| Auricularia, 553, 554, 562, 574
| |
| | |
| Autolytus cornutus, 319, 343
| |
| | |
| Aves, 56, 59, 61, 64, 107. 109
| |
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| Axolotl, 1 6
| |
| | |
| Balanoglossus, 576, 579, 581
| |
| | |
| Balanus balanoides, 75, 493
| |
| | |
| Belemnites, 252, 253
| |
| | |
| Bipinnaria, 557, 563, 574, 576, 579
| |
| | |
| Blatta, 374, 395
| |
| | |
| Bojanus, organ of, 264, 282
| |
| | |
| Bonellia, 20, 43, 44, 98, 324, 355, 358,
| |
| | |
| 359
| |
| Bothriocephalus salmonis, 211
| |
| | |
| ,, proboscideus, 212
| |
| | |
| Brachiella, 492
| |
| Brachiolaria, 558, 564
| |
| Brachiopoda, 311, 317, 318
| |
| Brachyura, 466, 480, 483
| |
| Branchiobdella, 42, 43, 346
| |
| Branchiogasteropoda, 272
| |
| Branchiopoda, 79, 459, 523, 524
| |
| Branchipus, 463, 524
| |
| Branchiura, 459, 492
| |
| Branchionus urceolaris, 221
| |
| Braula, 396
| |
| Uuccinum, 237, 280
| |
| Bulimus citrinus, 229
| |
| Bunodes, 169, 171
| |
| Buthus, 431
| |
| | |
| Calcispongiae, 138, 148
| |
| | |
| Calopteryx, 402
| |
| | |
| Calycophoridce, 152, 159
| |
| | |
| Calyptoblastic Hydroids, 184, 185
| |
| | |
| Calyptraea, 223, 280
| |
| | |
| Campanularidse, 183, 184
| |
| | |
| Capitclla, 330, 332
| |
| | |
| Carabidae, 476
| |
| | |
| Carcinus Mcenas, 481, 483
| |
| | |
| Cardium, 260, 262
| |
| | |
| " pygmaeum, 262
| |
| | |
| Carinaria, 240
| |
| | |
| Caryophyllium, 168, 171
| |
| pea, 165, 167
| |
| | |
| Cecidomyia, 15, 79, 416, 417, 429
| |
| | |
| Cephalopoda, 20, 40, 41, 102, 108, 109,
| |
| 135. "5. 240, 242, 244, 250, 252, 253,
| |
| 270, 271, 272, 274, 279, 282, 287
| |
| | |
| Cephalothrix galatheae, 202
| |
| | |
| Ceratosponguc, 146
| |
| | |
| | |
| | |
| Cercariae, 207, 208, 209
| |
| | |
| Cerianthus, 168, 171
| |
| | |
| Cestodes, 14, 29, 31, 32, 33, 189, 210,
| |
| 212, 218, 313, 425, 541
| |
| | |
| Chsetogaster, 342
| |
| | |
| Chaetopoda, 5, 18, 23, 41, 43, 44 , 54,
| |
| 67, 209, 215, 270, 275, 307, 312, 317,
| |
| 318, 319, 320, 326, 334, 33<S, 342, 346,
| |
| 349, 350, 351, 364. 369. 33, 36, 408,
| |
| 448, 457, 458, 521, 576,582
| |
| | |
| ChiXitopteridte, 333
| |
| | |
| Cha^tosomoidea, 371
| |
| | |
| Chelifer, 434, 436, 442, 446, 454
| |
| | |
| Chermes, 15, 429
| |
| | |
| Chilognatha, 113, 387, 389, 391, 393,
| |
| | |
| 395
| |
| | |
| Chilopoda, 387, 392, 394
| |
| Chilostomata, 292, 297, 298, 304, 305
| |
| Chironomus, 15, 378, 401, 402, 415, 416,
| |
| | |
| 429
| |
| | |
| Chiton, 254, 256, 257, 273
| |
| Chordata, 5
| |
| Chrysaora, 165
| |
| Chthonius, 436
| |
| Cicada, 395
| |
| | |
| Cirripedia, 459, 492, 496,503, 509, 520
| |
| Cladocera, 459, 464, 519
| |
| Clausilia, 239
| |
| Clavella, 520
| |
| Clavularia crassa, 167
| |
| Cleodora, 241
| |
| Clepsine, 73, 346, 347, 349, 351, 352,
| |
| | |
| 353, 354
| |
| Clio, 242, 278
| |
| Clubione, 436
| |
| Clupeidae, 64
| |
| Cobitis barbatula, 378
| |
| Coccida;, 429
| |
| Coccus, 50
| |
| Ccelebogyne, 79
| |
| Coelenterata, 3, 5, 13, 18, 26, 27, 2S, 35,
| |
| | |
| 74, 93, 94, 126, 148, 170, 178,179, 1 80,
| |
| | |
| 181, 191, 342
| |
| | |
| Ccenurus cerebralis, 213, 214
| |
| Coleochaete, 1 1
| |
| | |
| Coleoptera, 396, 402, 409, 412, 420, 421,
| |
| ^5
| |
| | |
| Collembola, 395, 426
| |
| Comatula, 5, 552, 553
| |
| Condracanthus, ill, 120, 520
| |
| Conochilus volvox, 22 1
| |
| Convoluta, 32
| |
| Copepoda, 109, 120, 459, 460, 487, 489,
| |
| | |
| 493, 496, 503, 509, 519
| |
| Corallium rubrum, 168, 182
| |
| Corethra, 422, 423, 424
| |
| Crangoninoe, 476
| |
| Crnniiuhv, 311
| |
| Craniata, 5, 6, 19, 20, 54, 56, 59, 6l, 62,
| |
| | |
| 6 4 , 74, 102
| |
| | |
| Crinoidea, 35, 36, 544, 550, 568, 576
| |
| Criodilus, 321, 324, 328, 341
| |
| | |
| | |
| | |
| INDEX.
| |
| | |
| | |
| | |
| Crisia, 304
| |
| Crocodilia, 63
| |
| | |
| Crustacea, 5, 6, 18, 51, 66, 102, 109, 120,
| |
| 458, 4 6 5> 487* 5<>2, 521, 524, 537, 541
| |
| Cryptophialus, 499, 509
| |
| Crystalloides, 163
| |
| Ctenophora, 26, 93, 102, 152, 173, 175,
| |
| | |
| 177, 178, 179, 180, 181, 182
| |
| Ctenostomata, 292, 297, 298, 304, 305
| |
| Cucullarms elegans, 46, 75, 82, 371, 376
| |
| Cucumaria, 546, 556, 574
| |
| Cumaceae, 459, 465, 486, 506
| |
| Curculio, 421
| |
| | |
| Cyclas, 259, 260, 261, 265
| |
| Cyclops, 376, 377, 418, 489, 503
| |
| Cyclostomata, 102, -292, 304
| |
| Cymbulia, 241, 242
| |
| | |
| Cymothoa, 516, 517, 519, 520,524, 528
| |
| Cynipidae, 15, 421, 428
| |
| Cyphonautes, 297, 301, 304, 306, 308
| |
| Cypridina, 500, 502
| |
| Cysticercus cellulosce, 214, 217
| |
| | |
| ,, fasciolaris, 216
| |
| | |
| ,, limacis, 213
| |
| | |
| Daphnia, 79, 464
| |
| | |
| Dasychone, 331, 336
| |
| | |
| Decapoda, 66, 248, 459, 465, 469, 504,
| |
| | |
| 511
| |
| | |
| Dendroccela, 32, 33, 189, 195, 196
| |
| Dentalium, 258, 576
| |
| Desmacidon, 147
| |
| Desor, type of, 196, 197, 201, 202, 204,
| |
| | |
| 212, 424
| |
| Diastopora, 304
| |
| Dibranchiata, 225, 253
| |
| Dicyema, 9, 131, 134, 135, 136
| |
| Dimya, 225
| |
| Diphyes, 159
| |
| Diplozoon, 11, 209, 210
| |
| Diporpa, 210
| |
| Diptera, 49, 194, 204, 396, 401,402,407,
| |
| | |
| 409, 412, 416, 420, 429
| |
| Discina radiata, 317
| |
| Discinidse, 311
| |
| | |
| Discophora, 18, 42, 165, 346, 383
| |
| Distomese, 189, 205, 425
| |
| Distomum, 31
| |
| | |
| ,, cygnoides, 209
| |
| | |
| ,, globiparum, 207
| |
| | |
| ,, lanceolatum, 205
| |
| Dochmius duodenale, 375
| |
| | |
| ,, trigonocephalus, 375
| |
| Donacia, 401
| |
| Dracunculus, 376, 377
| |
| | |
| Echinaster fallax, 23
| |
| | |
| ,, Sarsii, 102, 561
| |
| Echinodermata, 5, 18, 24, 35, 74, 102,
| |
| | |
| 325, 424, 544, 573, 574, 57 6 > 5 82
| |
| Echinoidea, 35, 36, 544, 549, 565, 576
| |
| Echinorhyncus, 379, 380
| |
| | |
| | |
| | |
| Echinus lividus, 83, 84, 88
| |
| | |
| Echiurus, 44 , 357, 358
| |
| | |
| Ectoprocta, 297, 306
| |
| | |
| Edriophthalmata, 459, 465
| |
| | |
| Elaphocaris, 473
| |
| | |
| Elasmobranchii, 23, 56, 59, 61, 62, 64,
| |
| | |
| 67, 105, 106. 107, 108, 109
| |
| Enopla, 189, 202
| |
| Entoconcha mirabilis, 237
| |
| Entomophaga, 421
| |
| | |
| Entoprocta, 292, 298, 300, 302, 304, 306
| |
| Epeira, 436
| |
| | |
| Ephemera, 395, 409, 420, 422
| |
| Ephyra, 186
| |
| | |
| Epibulia auranliaca, 159, 165
| |
| Erichthus, 484, 507
| |
| Errantia, 319, 336
| |
| Esperia, 147
| |
| Estheria, 463, 464
| |
| Euaxes, lol, 322, 324, 341, 346,349
| |
| Eucharis, 178
| |
| | |
| ,, multicornis, 178
| |
| | |
| Eucopepoda, 459
| |
| Eucope polystyla, 23, 154
| |
| | |
| Eunice sanguinea, 319
| |
| | |
| Eupagurus prideauxii, 112, 113, 115, 511,
| |
| 520
| |
| | |
| Euphausia, 465, 468, 504, 505, 518, 523
| |
| | |
| Eurostomata, 176
| |
| | |
| Eurylepta auriculata, 192
| |
| | |
| Eurynome, 483
| |
| | |
| Euspongia, 146, 147
| |
| | |
| Filaria, 377
| |
| Filaridae, 371
| |
| Firoloidea, 240
| |
| Flagellata, 7, 8
| |
| Flustrella, 301, 303
| |
| Formica, 396
| |
| Fungia, 182, 186
| |
| Fusus, 275, 280, 284, 288
| |
| | |
| Gammarus, 122, 518
| |
| | |
| ,, fluviatilis, 117
| |
| ,, locusta, no, 112
| |
| Ganoids, 54, 102
| |
| Gasteropoda, 39, 41, 98, 225, 226, 229,
| |
| | |
| 230, 232, 233, 240, 258, 260, 261, 270,
| |
| | |
| 272, 275, 279, 283, 324
| |
| Gasterosteus, 64, 210
| |
| Gastrotricha, 370
| |
| Gasterotrochce, 330, 333
| |
| Gecarcinus, 465
| |
| Geophilus, 392, 393
| |
| Gephyrea, 5, 18, 24, 44, 54, 67, 102,
| |
| | |
| 318, 320, 325, 355, 357, 361, 364
| |
| Germogen, 134
| |
| Geryonia hastata, 156
| |
| Geryonidse, 156
| |
| Glochidia, 267, 268
| |
| Gnathobdellidas, 346, 349
| |
| Gordiacea, 94
| |
| | |
| | |
| | |
| 588
| |
| | |
| | |
| | |
| INDEX.
| |
| | |
| | |
| | |
| Cimlioidca, 371, 374, 378
| |
| | |
| ;nia, 168
| |
| Gorgonidce, 181
| |
| Gorgoninrc, 181
| |
| Gregarinidae, 8
| |
| Gryllotalpa, 401, 412, 413
| |
| Gunnnineiv, 147, 148
| |
| Gymnoblastic Hydroids, 184, 185
| |
| Gymnoloemata, 292
| |
| | |
| Gymnosomata, 225, 240, 241, 242, 270
| |
| Gyrodactylus, 210
| |
| | |
| Halichondria, 147
| |
| | |
| Ilalisarca, 22, 66, 145
| |
| | |
| Halistemma, 165
| |
| | |
| Helicidce, 238
| |
| | |
| Helioporidae, 182
| |
| | |
| Helix, 67, 229
| |
| | |
| Hemiptera, 395, 402, 403, 409, 420, 421
| |
| | |
| Hessia, 108, 492
| |
| | |
| Heterakis vermicularis, .374
| |
| | |
| Heteronereis, 343
| |
| | |
| Heteropoda, 71, 72, 225, 226, 231, 278
| |
| | |
| Hexacoralla, 152, 179, 182
| |
| | |
| Hippopodius gleba, 27, 159
| |
| | |
| Hirudinea, 74, 84
| |
| | |
| Hirudo, 350, 351, 352, 353, 354
| |
| | |
| Holometabola, 420, 422
| |
| | |
| Holostomum, 205
| |
| | |
| Holothuria, 19, 25, 35, 549, 558, 576
| |
| | |
| Holothuroidea, 35, 544, 553, 556
| |
| | |
| Homarus, 477
| |
| | |
| Hyaleacea, 273, 275
| |
| | |
| Hyaleidce, 241
| |
| | |
| Hydra, 21, 22, 26, 28, 29, 34, 152, 154,
| |
| | |
| 155. 179, 183
| |
| Hydractinia, 539
| |
| Hydrocoralla, 152, 181, 185
| |
| Hydroidea, 152
| |
| Hydromedusae, 152, 179, 182, 183, 184,
| |
| | |
| 185, 186, 187
| |
| Hydrophilus, 374, 396, 400, 401, 402,
| |
| | |
| 404, 408, 409
| |
| Hydrozoa, 14, 19, 26, 27, 67, 102, 152,
| |
| | |
| 155. 165. 179, 1 80, 181, 182, 539
| |
| Ilymenoptera, 396, 401, 402, 412, 420,
| |
| | |
| 421, 425
| |
| | |
| Ichneumon, 396
| |
| | |
| Inarticulata, 311, 316
| |
| | |
| Incrmi
| |
| | |
| Infusoria, 7, 8
| |
| | |
| Insecta, 5, 15, 18, 19, 25, 46, 395, 396,
| |
| | |
| 4^5, 455. 45
| |
| Intoshia gigas, 136
| |
| Isidimc, 181
| |
| Ixxlyctia, 147
| |
| Isopoda, 109, 515, 519, 521, 523, 527
| |
| | |
| Julus Moneletei, 387, 388, 389
| |
| Kochlorine, 499
| |
| | |
| | |
| | |
| Lacertilia, 64
| |
| Lacinularia, 221, 223
| |
| | |
| socialis, 75
| |
| Lamellibranchiata, 23, 25, 37, 39, 98,
| |
| | |
| 225, 241, 257, 258, 259, 269, 270, 271,
| |
| | |
| 273, 274, 288
| |
| Lepadkue, 498
| |
| | |
| Lepas fascicularis, 224, 493, 494, 495
| |
| Lepidoptera, 79, 396, 402, 407, 408, 412,
| |
| | |
| 413, 415, 417, 420, 421, 423, 415, 426.
| |
| | |
| 455
| |
| | |
| Leptodora, 16, 51
| |
| Leptoplana, 74, 189, 192, 193
| |
| Lernseopoda, 490, 492, 520
| |
| Leucifer, 507
| |
| | |
| Libellulidae, 402, 403, 409, 420
| |
| Limax, 229, 232, 239, 278, 280
| |
| Limnadia, 79, 524
| |
| Limulus, 534
| |
| Lina, 402
| |
| | |
| Lingulidae, 311, 316
| |
| Lithobius, 393
| |
| Lobatse, 178
| |
| | |
| Loligo, 242, 243, 244, 247, 253
| |
| Loricata, 507, 514
| |
| Lota, 105
| |
| | |
| Loxosoma, 292, 294, 296, 306, 307
| |
| Lucernaria, 185
| |
| Lumbricus, 341, 368
| |
| | |
| ,, agricola, 321
| |
| | |
| ,, rubellus, 324
| |
| | |
| trapczoides, 13, 321, 323
| |
| Lumbriconereis, 334
| |
| Lymnseus, 82, 98, 226, 227, 232, 238,
| |
| | |
| 281
| |
| Lycosa, 436
| |
| | |
| Macrostomum, 32, 34
| |
| | |
| Macrura, 476
| |
| | |
| Malacobdella, 203
| |
| | |
| Malacodermata, 171
| |
| | |
| Malacostraca, 66, 459, 462, 465, 504,
| |
| | |
| 505, 506, 511, 523
| |
| Mammalia, 56, 58, 59, 64, 66
| |
| Marsipobranchii, 59
| |
| Mastigopus, 473, 474
| |
| Medusoe, 27, 154, 157, i.^s, 16;, 164, 176,
| |
| | |
| 178, 181, 182, 183, 184, 185, 186
| |
| Megalopa, 482, 483, 484
| |
| Melolontha, 402, 421
| |
| Membranipora, 297, 303
| |
| Mermithido;, 371
| |
| Mesotrochoe, 330
| |
| Metachoetoe, 335
| |
| Metazoa, Q, 10, 12,67, 86, 125, 135, 14^,
| |
| | |
| ISO, 179
| |
| | |
| Millepora, 152, 181
| |
| | |
| Mitraria, 308, 337
| |
| | |
| Molgula, 102
| |
| | |
| Mollusca, 5, 18, 24, 66, 74, 84, 99, 225,
| |
| 247, 248, 251, 256, 257, 262, 271, 285,
| |
| 288, 307, 325, 333, 352, 576, 582
| |
| | |
| | |
| | |
| INDEX.
| |
| | |
| | |
| | |
| 589
| |
| | |
| | |
| | |
| Monomya, -225
| |
| Monostomum capitellum, 205
| |
| | |
| ,, mutabile, 205, 206
| |
| | |
| Monotrochse, 330
| |
| Montacuta, 260, 262
| |
| Musca, 396
| |
| Muscidae, 420, 423
| |
| Myobia, 444, 445
| |
| Myrianida, 343
| |
| Myriapoda, 22, iir, 113, 387, 394, 395,
| |
| | |
| 4i.3 458
| |
| Mynothela, 155
| |
| Myrmeleon, 396
| |
| | |
| Mysis, 120, 469, 472, 486, 504, 509, 525
| |
| Mytilus, 260, 261
| |
| Myxinoids, 5
| |
| Myxispongise, 145
| |
| Myzostomea, 369
| |
| | |
| Nais, 342
| |
| | |
| Nassa mutabilis, 101, 226, 227, 233, 262,
| |
| | |
| 278, 279, 288, 3^4
| |
| Natantia, 487
| |
| Natica, 237, 283
| |
| Nauplius, 5, 16, 460, 461, 463, 465, 466,
| |
| | |
| 469, 473, 490, 491, 493, 497
| |
| Nautilus pompilius, 253, 276
| |
| Nebaliadse, 459, 465, 486
| |
| Nematoda, 45, 46, 50, 66, 74, 75, 371,
| |
| | |
| 373. 374> 376
| |
| Nematogens, 131
| |
| Nematoidea, 18, 84, 94, 371, 374
| |
| Nematus ventricosus, 13, 427
| |
| Nemertea, 94, 189, 196, 202, 204
| |
| Nemertines, 30, 31, 33, 93, 136, 195,
| |
| | |
| 202, 328, 333, 424
| |
| Nephelis, 82, 346, 349, 350, 351, 352,
| |
| | |
| 354
| |
| | |
| Nereis, 343
| |
| | |
| ,, diversicolor, 319
| |
| | |
| ,, Dumerilii, 343
| |
| Neritina, 229, 237
| |
| Neuroptera, 396, 401, 420, 421
| |
| Neuroterus ventricularis, 428
| |
| Notonecta, 395
| |
| Nototrochse, 330, 353
| |
| Nudibranchiata, 229, 241
| |
| | |
| Ocellata, 184
| |
| | |
| Octocoralla, 152, 179
| |
| | |
| Octopus, 248
| |
| | |
| Odontophora, 225, 257, 271
| |
| | |
| Odontosyllis, 333
| |
| | |
| Oedogonium, 1 1
| |
| | |
| Oligochseta, 42, 319, 321, 325, 330, 338,
| |
| | |
| 346, 352
| |
| Olynthus, 144
| |
| Oniscus murarius, 107, 108, 109, 120,
| |
| | |
| 516, 520, 528
| |
| Opercula, 31
| |
| Ophiothryx, 36, 549
| |
| Ophidia, 64
| |
| | |
| | |
| | |
| Ophiuroidea, 136, 544, 553, 562, 565,
| |
| | |
| 576
| |
| | |
| Ophryotrochoe puerilis, 333
| |
| Opisthobranchiata, 225, 232, 237
| |
| Ornithodelphia, 109
| |
| Orthonectidae, 136
| |
| | |
| Orthoptera, 395, 414, 420, 421, 425, 426
| |
| Ostracoda, 459, 500, 510
| |
| Ostrea, 259, 260, 262
| |
| Oxyuridse, 46, 373, 374
| |
| Oxyurus ambigua, 374
| |
| ,, vermicularis, 375
| |
| | |
| PcEcilopoda, 534
| |
| Paguridse, 477
| |
| Pakemon, no
| |
| Palaemonetes, 476
| |
| Pakemoninre, 476, 511, 512
| |
| Palinurus, 478, 480
| |
| | |
| Paludina, 66, 227, 229, 235, 270, 278,
| |
| 280
| |
| | |
| ,, costata, 229
| |
| | |
| ,, vivipara, 226
| |
| Pandorina, n
| |
| Parasita, 489
| |
| Pedalion, 221
| |
| | |
| Pedicellina, 98, 292, 296, 299, 307
| |
| Pelagia, 167, 185
| |
| Penseinse, 476
| |
| Penaeus, no, 113, 465, 469, 473, 474,
| |
| | |
| 504, 518
| |
| | |
| Pennatulidae, 181
| |
| Pentacrinus, 5
| |
| Pentastomida, 539, 540
| |
| Pentastomum denticulatum, 540, 54!
| |
| | |
| tsenoicles, 539, 540, 541
| |
| Percidae, 64
| |
| PerennichaetcE, 335
| |
| Peripatus, 5, 386, 411, 412, 413, 542
| |
| Petromyzon, 61, 63, 64, 74, 83
| |
| Phalangella, 304
| |
| Phalangidse, 436
| |
| Phallusia, 83
| |
| | |
| Phascolosoma, 44, 355, 356, 361
| |
| Pholcus, 436, 442
| |
| Phoronis, 315, 355, 363, 364
| |
| Phoxinus laevis, 378
| |
| Phryganea, 396, 401, 409
| |
| Phylactokemata, 292, 294, 297, 305, 306
| |
| Phyllobothrium, 218
| |
| Phyllodoce, 329
| |
| Phyllopoda, 16, 459, 461, 505
| |
| Phyllosoma, 479, 480
| |
| Phylloxera, 429
| |
| Physophoridoe, 152, 16-2, 164
| |
| Pilidium, type of, 196, 200, 201, 202, 704,
| |
| | |
| 424
| |
| | |
| Pisces, 5
| |
| Piscicola, 20, 43
| |
| Pisidium, 259, 260, 262, 264
| |
| Planaria Neapolitana, 193
| |
| Planorbis, 273, 281, 325
| |
| | |
| | |
| | |
| 590
| |
| | |
| | |
| | |
| INDEX.
| |
| | |
| | |
| | |
| Platyelminthes, 18, 20, 24, 221, 424
| |
| Platygaster, 396, 416, 417, 418, 419
| |
| Pleurohrachia, 176, 177, 238
| |
| Pneumodermon, 242, 576
| |
| Podostomata, 292
| |
| Poduridce, 401, 405
| |
| Polychaeta, 42, 319, 325, 338
| |
| Polydesmus complanatus, 387, 388
| |
| Polygordius, 319, 325, 326, 327, 328,
| |
| | |
| 332, 357 386
| |
| Polynoe, 42, 331
| |
| Polyophthalmus, 328
| |
| Polyplacophora, 225, 254, 270, 271, 288
| |
| Polystomeas, 189, 205, 209
| |
| Polystomum, 209
| |
| | |
| ,, integerrimum, 30, 31, 210
| |
| | |
| Polytrochne, 330, 333
| |
| Polyxenia leucostyla, 158
| |
| Polyxenus lagurus, 387
| |
| Polyzoa, 98, 303, 305, 306. 30 8 > 3 ! 5. 3^
| |
| Porcellana, 483
| |
| Porifera, 102, 138, 148
| |
| Porthesia, 115
| |
| Prorhyncus, 32, 34
| |
| Prosobranchiata, 225, 237, 281
| |
| Prostomum, 32, 34, 38, 196
| |
| Protozoa, 8, 9, lo, n, 86, 135, 149
| |
| Protozoaea, 471
| |
| Protula Dysteri, 342
| |
| Pseudoneuroptera, 426
| |
| Pseudoscorpionid;e, 434
| |
| Psolinus, 556, 574
| |
| Psychidae, 16
| |
| Pteraster miliaris, 561
| |
| Pteropoda, 98, 225, 226, 229, 230, 232,
| |
| | |
| 240, 258, 270, 272, 279, 283
| |
| Pterotrachcea, 71, 229, 240
| |
| Pulex, 396
| |
| | |
| Pulmonata, 39, 225, 232, 238, 281, 282
| |
| I'urpura lapillus, 78
| |
| Pycnogonida, 538
| |
| Pyrosoma, 13, 53, 109
| |
| | |
| Rana temporaria, 210
| |
| Kaspailia, 147
| |
| Rcdia, 206, 207, 208, 209
| |
| Reniera, 147
| |
| | |
| Kcptilia, 56, 59, 60, 61, 62, 64, 109
| |
| Rhabditis dolichura, 82
| |
| Khabdoccela, 32, 33, iSy, ic/>
| |
| Khnbdopleura, 294, 306
| |
| Rhi/occphala, 459, 493, 499, 500
| |
| Klii/.ocrinus, 5
| |
| klii/.ostoma, 167
| |
| Rhomlx>gens, 131, 134
| |
| Khynchoncllidaj, 311
| |
| Rhyncdbddlkbe, 346
| |
| Rotifera, 5, 12, 18, 75, 76, 77, 79, 83,
| |
| 102, 221, 308, 325
| |
| | |
| Saccocirrus, 328, 329, 332, 340
| |
| Sacculina, 500
| |
| | |
| | |
| | |
| Sagartia, 169, 171
| |
| | |
| Sagitta, 33, 74, 94, 130, 366, 367, 368
| |
| | |
| Salmonidrc, 64
| |
| | |
| Salpa, 102
| |
| | |
| Sarcia, 164
| |
| | |
| Seaphopoda, 225, 257, 270, 271
| |
| | |
| Schistocephalus, 2 1 1
| |
| | |
| Schizopoda, 459, 465, 466
| |
| | |
| Scolopendra, 392
| |
| | |
| Scorpio, 120, 43 r, 44 6, 454, 455, 457
| |
| | |
| Scrobicularia, 38, 39
| |
| | |
| Scyllarus, 477
| |
| | |
| Scyphistoma, 179, 185, 186
| |
| | |
| Sedentaria, 319, 336
| |
| | |
| Sepia, 20, 40, 41, 242, 243, 244, 245,
| |
| | |
| 247> 2 49> 253
| |
| Sergestidce, 473, 507
| |
| Serpula, 319. 325, 331
| |
| Sertularia, 152, 183, 184
| |
| Silicispongia.', 147
| |
| Simulia, 401, 415
| |
| Siphonophora, 13, 77, 152, 159, 163,
| |
| | |
| 165, 179, 1 80, 182, 185
| |
| Sipunculida, 24
| |
| Sipunculus, 44
| |
| Sirex, 396
| |
| Sitaris, 42!
| |
| | |
| Spathegaster baccarum, 428
| |
| Spjo, 4 2 > 33 2 > 333
| |
| Spiroptera obtusa, 376
| |
| Spirorbis Pagenstecheri, 319
| |
| | |
| spirillum, 319, 336
| |
| Spirula, 252
| |
| Spirulirostra, 252
| |
| Spongelia, 147
| |
| Spongida, 138, 144, 148, 149
| |
| Spongilla, 147, 150
| |
| Sporocysts, 206, 207, 208, 209
| |
| Squilla, 66, 504, 507
| |
| Stephanomia pictum, 162, 165
| |
| Stomalopoda, 459, 465, 4X4
| |
| Stomodoeum, 413
| |
| Strongylidrc, 371, 375
| |
| Strongylocentrus, 567
| |
| Strongysoloma Guerinii, 3<S7, 388, 390
| |
| Stylasterictae, 152, r8r
| |
| Styliolidic, 24!
| |
| Stylochopsis ponticus, 193
| |
| Sycandra, 93, 138, 144, 145, 147, 150
| |
| | |
| ,, raphanus, i^S, 174
| |
| Syllis, 343
| |
| | |
| vivipara, 319
| |
| Sympodium coralloidcs, 168
| |
| | |
| Taeniatoe, 178
| |
| Tardigrada, 541
| |
| Teoenaria, 436
| |
| | |
| 'I'clcDsti'i, IS, 25, 5^), 59, C>4. 107, io<)
| |
| I'r].)troch;i.-, 330
| |
| Tcndra, 300
| |
| 'I '(.'nth reds, 396
| |
| Tcrcbdla concliilcga, 332, 333, 337
| |
| | |
| | |
| | |
| INDEX.
| |
| | |
| | |
| | |
| 591
| |
| | |
| | |
| | |
| Terebella nebulosa, 332, 333
| |
| Terebratula, 311, 315
| |
| Terebratulina, 311, 315, 316
| |
| | |
| ,, septentrionalis, 315, 316
| |
| | |
| Teredo, larva of, 262
| |
| Tergipes, 232, 238
| |
| | |
| ,, Edwardsii, 238
| |
| ,, lacinulatus, 238
| |
| Tethya, 147
| |
| Tetrabranchiata, 225
| |
| Tetranychus telarius, 116
| |
| Tetrastemma varicolor, 203
| |
| Thalassema, 44, 355, 357
| |
| Thalassinidae, 477
| |
| Thallophytes, n
| |
| Thecidium, 311, 312, 315, 316
| |
| Thecosomata, 225
| |
| Thoracica, 459, 493, 499, 500
| |
| Thysanozoon, 192, 193
| |
| Thysanura, 395, 408, 425, 458
| |
| Tichogonia, 39
| |
| Tipula, 396
| |
| Tipulidae, 420, 421
| |
| Toenia cosnurus, 214
| |
| | |
| ,, echinococcus, 215, 217
| |
| | |
| solium, 217
| |
| Tornaria, 579, 581
| |
| Toxopneustes, 22, 24, 35, 85, 88, 89
| |
| Tracheata, 385, 426, 432, 44 8, 455, 457,
| |
| | |
| 458, 538, 54i
| |
| | |
| Trachymedusae, 152, 156, 179, 185
| |
| Trematodes, 14, 16, 29, 30, 31, 32, 33,
| |
| | |
| 46, 94, 189, 205, 208, 210, 212, 216
| |
| | |
| | |
| | |
| Trichina, 377, 378
| |
| | |
| Trichinidse, 371
| |
| | |
| Trichocepha'lus affinis, 374
| |
| | |
| Trochosphsera aequatorialis, 221
| |
| | |
| TubiporidcE, 182
| |
| | |
| Tubularia, 34, 38, 152, 154, 158
| |
| | |
| Tubularidse, 29, 179, 183
| |
| | |
| Tunicata, 5, I 4 , 53
| |
| | |
| Turbellaria, 5, 30, 31, 33, 74, 98, 102,
| |
| | |
| 136, 179, 189, 193, 333
| |
| Tyroglyphus, 445
| |
| | |
| Unio, 37, 38, 39, 100, 101, 259, 260,265,
| |
| 266, 445
| |
| | |
| Vaginulus luzonicus, 229
| |
| | |
| Vermes, 5, 74, 102, 223, 324, 352
| |
| | |
| Verongia rosea, 146
| |
| | |
| Vertebrata, 14, 18, 19, 24, 59, 64, 83,
| |
| | |
| 272, 349. 397' 4^6
| |
| Vesiculata, 184
| |
| Vitrina, 229
| |
| Vorticella, 8, 9, 10
| |
| | |
| Wilsia, 164
| |
| Xiphoteuthis, 252
| |
| | |
| Zoantharia, 152, 168, 169
| |
| Zooea, 465, 468, 471, 474, 482, 483, 484,
| |
| 486, 504
| |
| | |
| | |
| | |
| BIBLIOGRAPHY.
| |
| | |
| | |
| | |
| THE OVUM.
| |
| | |
| General Works.
| |
| | |
| (1) } Ed. van Beneden. "Recherches sur la composition et la signification de
| |
| ,A T m ' cour ' d ' l Acad " r y- des Sci <<* de Belgique, Vol. xxxiv. 1870.
| |
| | |
| / '%, R- Leuckart. Artikel "Zeugung," R. WagMsfs Handworterbtek d. Physio
| |
| logte, Vol. iv. 1853.
| |
| | |
| (3 ^ Fr ' L/ydig- , " Die Dotterfurchung nach ihrem Vorkommen in d. Thierwelt
| |
| u. n. ihrer Bedeutung." Oken. Isis, 1848.
| |
| | |
| (4) Ludwig. "Ueber d. Eibildung im Thierreiche." Arbeiten a. d. zool.-zoot
| |
| Institiit. Wiirzburg, Vol. I. rSy^.
| |
| | |
| (5) AllenThomson. Article ' ' Ovum " in Todd's Cyclopedia of Anatomy and
| |
| Physiology, Vol. v. 1859.
| |
| | |
| (6) W. Waldeyer. Eierstock u. EL Leipzig, 1870.
| |
| | |
| THE OVUM OF CCELENTERATA.
| |
| | |
| (7) Ed. van Beneden. "De la distinction originelle d. testicule et de
| |
| 1'ovaire." Bull. Acad. roy. Belgique, f serie, Vol. xxxvu. 1874.
| |
| | |
| (8) R. and O. Hertwig. Der Organismus d. Medusen. Jena, 1878.
| |
| | |
| (9) N. Kleinenberg. Hydra. Leipzig, 1872.
| |
| | |
| THE OVUM OF PLATYELMINTHES.
| |
| | |
| (10) P. Hallez. Contributions a fHistoire naturelle des Turbellarih. Lille,
| |
| 1879.
| |
| | |
| (11) S. MaxSchultze. Beitrdge z. Naturgeschichte d. Turbellarien. Greifswald, 1851.
| |
| | |
| (12) C. Th. von Siebold. ' ' Helminthologische Beitrage." Miiller's Archiv,
| |
| 1836.
| |
| | |
| (13) C. Th. von Siebold. Lehrbuch d. vergleich. Anat.d. wirbellosen Thiere.
| |
| Berlin, 1848.
| |
| | |
| (14) E. Zeller. " Weitere Beitrage z. Kenntniss d. Polystomen." Zeit. f.
| |
| wiss. ZooL, Bd. xxvu. 1876.
| |
| | |
| [Vide also Ed. van Beneden (No. i).]
| |
| | |
| THE OVUM OF ECHINODERMATA.
| |
| | |
| (15) C. K. Hoffmann. " Zur Anatomic d. Echiniden u. Spatangen." Niederllindisch. Archiv f. Zoologie, Vol. I. 1871.
| |
| | |
| (16) C. K. Hoffmann. " Zur Anatomic d. Asteriden. Niederldndisch. Ardiiv
| |
| /. Zoologie, Vol. n. 1873.
| |
| | |
| (17) H. Ludwig. "Beitrage zur Anat. d. Crinoiden." Zeil. f. wiss. Zool.,
| |
| Vol. xxvin. 1877.
| |
| | |
| (18) Job. Miiller. "Ueber d. Canal in d. Eiern d. Holothurien." Miiller's
| |
| Archiv, 1854.
| |
| | |
| (19) C. Semper. Holothurien. Leipzig, 1868.
| |
| | |
| (20) E. Selenka. Befruchtung d. Eies v. Toxopneustes variegalits, 1878.
| |
| | |
| [Vide also Ludwig (No. 4), etc.]
| |
| | |
| 1 A very complete and critical account of the literature is contained in this paper.
| |
| B. II. a
| |
| | |
| | |
| | |
| BIBLIOGRAPHY.
| |
| | |
| | |
| | |
| THE OVUM OF MOLLUSC A.
| |
| Lamellibranchiata.
| |
| | |
| (21) II. Lacaze-Duthiers. " Organes genitaux des Acephales Lamellibranches." Ann. Set. Nat., 4 mc serie, Vol. 1 1. 1854.
| |
| | |
| (22) W. F lemming. " Ueb. d. er. Entwick. am Ei d. Teichmuschel." Archiv
| |
| f. mikr. Anat., Vol. x. 1874.
| |
| | |
| (23) W. Flamming. "Studien lib. d. Entwick. d. Najaden." Sitz. d. t: Akad.
| |
| Wiss. men, Vol. LXXI. 1875.
| |
| | |
| (24) Th. von Hassling. " Einige Bemerkungen, etc." Zeit. f. wiss. ZooL,
| |
| Bd. v. 1854.
| |
| | |
| (25) H. von Jhering. "Zur Kenntniss d. Eibildung bei d. Muscheln." Zeit.
| |
| f. wiss. ZooL, Vol. xxix. 1877.
| |
| | |
| (26) Keber. De Introihi Spermatozoorum in ovula, etc. Konigsberg, 1853.
| |
| | |
| (27) Fr. Leydig. " Kleinere Mittheilung etc." Miiller's Archiv, 1854.
| |
| | |
| Gasteropoda.
| |
| | |
| (28) C. Semper. "Beitrage z. Anat. u. Physiol. d. Pulmonaten." Zeit. f.
| |
| wiss. ZooL, Vol. vni. 1857.
| |
| | |
| (29) H. Eisig. " Beitrage z. Anat. u. Entwick. d. Pulmonaten." Zeit.f. wiss.
| |
| ZooL, Vol. xix. 1869.
| |
| | |
| (30) Fr. Leydig. " Ueb. Paludina vivipara." Zeit.f. wiss. ZooL, Vol. u. 1850.
| |
| | |
| Cephalopoda.
| |
| | |
| (31) Al. Kolliker. Entwicklungsgeschichte d. Cephalopoden. Zurich, 1844.
| |
| | |
| (32) E. R. Lankester. "On the Developmental History of the Mollusca."
| |
| Phil. Trans., 1875.
| |
| | |
| THE OVUM OF THE CHJETOPODA.
| |
| | |
| (33) Ed. Claparede. " Les Annelides Chaetopodes d. Golfe de Naples."
| |
| Mem.d. 1. Soctit. phys. eld 1 hist. nat. de Geneve, 1868 9 and 1870.
| |
| | |
| (34) E. Ehlers. Die Borstcnwiirmer nach system, und anat. Untersuchungen.
| |
| Leipzig, 186468.
| |
| | |
| (35) E. Selenka. " Das Gefass-System d. Aphrodite aculeata." Niedcrldndisches Archiv f. ZooL, Vol. n. 1873.
| |
| | |
| THE OVUM OF DISCOPHORA.
| |
| | |
| (36) H. Dorner. " Ueber d. Gattung Branchiobdella." Zeit.f. wiss. ZooL,
| |
| Vol. xv. 1865.
| |
| | |
| (37) R. Leuckart. Die menschlichen Parasiten.
| |
| | |
| (38) Fr. Leydig. "Zur Anatomie v. Piscicola eeometrica, etc." Zeit. f. wiss.
| |
| ZooL, Vol. I. 1849.
| |
| | |
| (30) C. O. Whitman. "Embryology of Clepsine." Quart. 7. of Alter.
| |
| Sci., Vol. xvin. 1878.
| |
| | |
| THE OVUM OF GEPHYREA.
| |
| | |
| (40) Keferstein u. Ehlers. Zoologische Beitrage. Leipzig, 1861.
| |
| | |
| (41) C. Semper. Holothurien, 1868, p. 145.
| |
| | |
| (42) J. W. Spengel. " Beitrage z. Kenntniss d Gephyreen." Beitriigc a. d.
| |
| zool. Stationz. Neapcl, Vol. I. 1879.
| |
| | |
| (43) J. W. Spengel. " Anatomische Mittheilungen lib. Gephyreen." Tagcbl.
| |
| d. Naturf. Vers. Munchen, 1877.
| |
| | |
| THE OVUM OF NEMATODA.
| |
| | |
| (44) Ed. Claparede. De la formation ct de la fccondaiiou dcs- n-uf.\ chcz Ics
| |
| I'crs Ntmatodcs. (ienevc, 1859.
| |
| | |
| (J- r )) K. I. (.-nek art. Hif nirnsf/i lichen Paras! ten.
| |
| | |
| | |
| | |
| BIBLIOGRAPHY. jjj
| |
| | |
| | |
| | |
| d.Nematoden."
| |
| | |
| ^' Nels0n * " On the reproduction of Ascaris mystax, etc." Phil.
| |
| (48) A.Schneider. Monographie d.' Nematoden. Berlin, 1866.
| |
| THE OVUM OF INSECT A.
| |
| | |
| | |
| | |
| Sm ' T? r u n d V Ueb ,?'* a5 Ei u ' seine Bildungsstdtte. Leipzig, 1 878.
| |
| (50) T. H. Huxley. " On the agamic reproduction and morphology of Aphis.
| |
| Ltnnean Trans., Vol. xxn. 1858. Vide also Manual of Invertebrate* Animals, 1877.
| |
| | |
| 1 * ^ ^ ^ *
| |
| | |
| | |
| | |
| (51)
| |
| bei den *,++,*
| |
| | |
| /-a\ ? r ',k ey< MS' Der Eierstock u. die Samentasche d. Insecten. Dresden, 1866.
| |
| tSl ~ ub . bock - " The ov a and pseudova of Insects." Phil. Trans. 1850.
| |
| (o4) Stem. Die weiblichen Geschlcchtsorgane d. Ktifer. Berlin, 1847.
| |
| [Conf. also Glaus, Landois, Weismann, Ludwig (No. 4).]
| |
| | |
| THE OVUM OF ARANEINA.
| |
| | |
| (55) Victor Cams. " Ueb. d. Entwick. d. Spinneneies." Zeit. f. wiss. Zool. t
| |
| Vol. ii. 1850.
| |
| | |
| (56) v. Wittich. "Die Entstehung d. Arachnideneies im Eierstock, etc."
| |
| Miiller s Archiv, 1849.
| |
| | |
| [Conf. Leydig, Balbiani, Ludwig (No. 4), etc.]
| |
| | |
| THE OVUM OF CRUSTACEA.
| |
| | |
| (57) Aug. Weismann. "Ueb. d. Bildung von Wintereiern bei Leptodora
| |
| hyalina." Zeit.f. wiss.ZooL, Vol. xxvn. 1876.
| |
| | |
| [For general literature vide Ludwig, No. 4, and Ed. van Beneden, No. i.]
| |
| | |
| THE OVUM OF CHORD ATA.
| |
| | |
| Urochorda (Tunicata).
| |
| | |
| (58) A. Kowalevsky. " Weitere Studien ii. d. Entwicklung d. Ascidien."
| |
| Archiv f. micr. Anat., Vol. VII. 1871.
| |
| | |
| (59) A. Kowalevsky. "Ueber Entwicklungsgeschichte d. Pyrosoma."
| |
| Arch.f. micr. Anat., Vol. xi. 1875.
| |
| | |
| (60) Kupffer. " Stammverwandtschaft zwischen Ascidien u. Wirbelthieren."
| |
| Arch. f. micr. Anat., Vol. VI. 1870.
| |
| | |
| (61) Giard. " Etudes critiques des travaux, etc. " Archives Zool. experiment.,
| |
| Vol. I. 1872.
| |
| | |
| (62) C. Semper. " Ueber die Entstehung, etc." Arbeiten a. d. zool.-zoot.
| |
| Institut Wiirzburg, Bd. II. 1875.
| |
| | |
| Cephalochorda.
| |
| | |
| (63) P. Langerhans. "Z. Anatomic d. Amphioxus lanceolatus," pp. 330 3.
| |
| Archiv f. mikr. Anat., Vol. xil. 1876.
| |
| | |
| Craniata.
| |
| | |
| (64) F. M. Balfour. "On the structure and development of the Vertebrate
| |
| Ovary." Quart. J. of Micr. Science, Vol. xvm. 1878.
| |
| | |
| (65) Th. Eimer. " Untersuchungen ii. d. Eier d. Reptilien." Arckiv f.
| |
| mikr. Anat., Vol. vni. 1872.
| |
| | |
| (66) Pfliiger. Die Eierstbcke d. Sdugethiere u. d. Menschen. Leipzig, 1863.
| |
| | |
| (67) J. Foulis. " On the development of the ova and structure of the ovary in
| |
| Man and other Mammalia." Quart. J. of Micr. Science, Vol. XVI. 1876.
| |
| | |
| (68) J. Foulis. " The development of the ova, etc." Journal of Anat. and
| |
| Phys., Vol. xni. 18789.
| |
| | |
| a 2
| |
| | |
| | |
| | |
| IV BIBLIOGRAPHY.
| |
| | |
| | |
| | |
| (69) C. Gegenbaur. " Ueb. d. Bau u. d. Entwicklung d. Wirbelthiereier mit
| |
| partieller Dottertheilung." Muller's Archiv, 1861.
| |
| | |
| (70) Alex. Gotte. Entwicklungsgeschichte d. Unke. Leipzig, 1875.
| |
| | |
| (71) W. His. Untersuchungen iib. d. Ei u. d. Eientwicklung bei Knochenfischcn.
| |
| Leipzig, 1873.
| |
| | |
| (72) A. Kolliker. Entwicklungsgeschichte d. Menschen u. hoherer Thicre,
| |
| Leipzig, 1878.
| |
| | |
| (73) J. Miiller. " Ueber d. zahlreichen Porenkanale in d. Eikapsel d. Fische."
| |
| Muller's Archiv, 1854.
| |
| | |
| (74) W. H. Ransom. " On the impregnation of the ovum in the Stickleback."
| |
| Pro. K. Society, Vol. vn. 1854.
| |
| | |
| (75) C. Semper. " Das Urogenitalsystem d. Plagiostomen etc." Arbeiten a.
| |
| d. zool.-zoot. Instit. Wiirzburg, Vol. II. 1875.
| |
| | |
| [Cf. Ludwig, No. 4, Ed. van Beneden, No. i, Waldeyer, No. 6, etc.]
| |
| | |
| | |
| | |
| MATURATION AND IMPREGNATION OF THE OVUM.
| |
| | |
| (76) Auerbach. Organologische Studien, Heft 2. Breslau, 1874.
| |
| | |
| (77) Bambeke. " Recherches s. Embryologie des Batraciens." Bull, de
| |
| royale de Belgique, 2me ser., T. LXI. 1876.
| |
| | |
| (78) E. van Beneden. " La Maturation de 1'CEufdes Mammiferes." Bull,
| |
| de fAcad. royale de Belgique, 2me ser., T. XL. No. 12, 1875.
| |
| | |
| (79) Id em. " Contributions a 1'Histoire de la Vesicule Germinative, &c." Bull,
| |
| de fAcad. royale de Belgique, sme ser., T. XLI. No. i, 1876.
| |
| | |
| (80) O. Biitschli. Eizelle, Zelltheilung, und Conjugation der Infusorien.
| |
| Frankfurt, 1876.
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| 1 The dates in this reference are the dates of publication.
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| [[Category:Historic Embryology]][[Category:1800's]]
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Foster M. and Sedgwick A. The Works of Francis Balfour Vol. II. A Treatise on Comparative Embryology 1. (1885) MacMillan and Co., London.
- The Ovum and Spermatozoon | The Maturation and Impregnation of the Ovum | The Segmentation of the Ovum | Dicyemae and Orthonectidae Dicyema | Porifera | Coelenterata | Platyhelminthes | Rotifera | Mollusca | Polyzoa | Brachiopoda | Chilopoda | Discophora | Gephyrea | Chaetognatha | Nemathelminthes | Tracheata | Crustacea | Pcecilopoda | Echinodermata | Enteropneusta | Bibliography
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Vol II. A Treatise on Comparative Embryology (1885)
Chapter XX. Echinodermata
THE development of the Echinodermata naturally falls into
two sections:
(i) The development of the germinal layers and of the
systems of organs; (2) the development of the larval appendages
and the metamorphosis.
The Development of the Germinal Layers and of the Systems of Organs.
The development of the systems of organs presents no very
important variations within the limits of the group.
Holothuroidea. The Holothurians have been most fully
studied (Selenka, No. 563), and may be conveniently taken as
type.
The segmentation is nearly regular, though towards its close,
and in some instances still earlier, a difference becomes apparent
between the upper and the lower poles.
At the close of segmentation (fig. 247 A) the egg has a
nearly spherical form, and is constituted of a single layer of
columnar cells enclosing a small segmentation cavity. The
lower pole is slightly thickened, and the egg rotates by means of
fine cilia.
An invagination now makes its appearance at the lower
pole (fig. 247 B), and simultaneously there become budded off
from the cells undergoing the invagination amoeboid cells, which eventually form the muscular system and the connective tissue.
These cells very probably have a bilaterally symmetrical origin.
This stage represents the gastrula stage which is common to all
Echinoderms. The invaginated sack is the archenteron. As it
grows larger one side of the embryo becomes flattened, and the
other more convex. On the flattened side a fresh invagination arises, the opening of which forms the permanent mouth, the
opening of the first invagination remaining as the permanent
anus (fig. 248 A).
1 The following classification of the Echinodermata is employed in this chapter.
I. Holothuroidea. IV. Echinoidea.
II. Asteroidea. V. Crinoidea.
III. Ophiuroidea.
FIG. 247. TWO STAGES IN THE DEVELOPMENT OF HOLOTHURIA TUBULOSA VIEWED IN OPTICAL SECTION. (After Selenka.)
A. Blastosphere stage at the close of segmentation. B. Gastrula stage.
mr. micropyle ; //. chorion; s.c. segmentation cavity; bl. blastoderm; ep. epiblast;
hy. hypoblast; ms. amoeboid cells derived from hypoblast ; a.e. archenteron.
These changes give us the means of attaching definite names
to the various parts of the embryo. It deserves to be noted in
the first place that the embryo has assumed a distinctly bilateral
form. There is present a more or less concave surface extending from the mouth to near the anus, which will be spoken
of as the ventral surface. The anus is situated at the posterior
extremity. The convex surface opposite the ventral surface
forms the dorsal surface, which terminates anteriorly in a
rounded prse-oral prominence.
It will be noticed in fig. 248 A that in addition to the
primitive anal invagination there is present a vesicle (?/.).
This vesicle is directly formed by a constriction of the primitive
B. II. 35
54 6
HOLOTHUROIDEA.
archenteron (fig. 249 Vpv.), and is called by Selenka the vasoperitoneal vesicle. It gives origin to the epithelioid lining of
the body cavity and water-vascular system of the adult 1 . In the
parts now developed we have the rudiments of all the adult organs.
The mouth and anal involutions (after the separation of the
vaso-peritoneal vesicle) meet and unite, a constriction indicating
their point of junction (fig. 248 B). Eventually the former gives
FIG. 248. THREE STAGES IN THE DEVELOPMENT OF HOLOTHURIA TUBULOSA
VIEWED FROM THE SIDE IN OPTICAL SECTION. (After Selenka.)
tn. mouth; oe. oesophagus; st. stomach; i. intestine; a. anus; I.e. longitudinal
ciliated band; v.p. vaso-peritoneal vesicle; p.v. peritoneal vesicle; p.r. right peritoneal vesicle ; //. left peritoneal vesicle ; w.v. water- vascular vesicle ; p. dorsal pore
of water- vascular system ; ms. muscle cells.
rise to the mouth and cesophagus, and the latter to the remainder of the alimentary canal 2 .
The vaso-peritoneal vesicle undergoes a series of remarkable
changes. After its separation from the archenteron it takes
up a position on the left side of this, elongates in an anteroposterior direction, and from about its middle sends a narrow
diverticulum towards the dorsal surface of the body, where an
1 The origin of the vaso-peritoneal vesicle is not quite the same in all the species.
In Holothuria tubulosa it is separated from the csecal end of the archenteron; the
remainder of which then grows towards the oral invagination. In Cucumaria the
archenteron forks (fig. 249) ; and one fork forms the vaso-peritoneal vesicle, and the
other the major part of the mesenteron.
2 There appears to be some uncertainty as to how much of the larval cesophagus is
derived from the stomodaeal invagination.
ECHINODERMATA.
547
opening to the exterior becomes formed (fig. 248 B, /.). The
diverticulum becomes the madreporic canal, and the opening
the dorsal pore.
The vaso-peritoneal vesicle next divides into two, an anterior vesicle (fig. 248 B, w.v.), from which is derived the
epithelium of the water-vascular system, and a posterior (fig.
248 B, /.?;.), which gives rise to the epithelioid lining of the body
cavity. The anterior vesicle (fig. 248 C, w.v.) becomes fivelobed, takes a horseshoe-shaped form, and grows round the
oesophagus (fig. 256, w.v.r). The five lobes form the rudiments
of the water-vascular prolongations into the tentacles. The
remaining parts of the water-vascular system are also developed
as outgrowths of the original vesicle. Five of these, alternating
with the original diverticula, form the five ambulacral canals,
from which diverticula are produced into the ambulacral feet ; a
sixth gives rise to the Polian vesicle. The remaining parts of
the original vesicle form the water-vascular ring.
We must suppose that eventually the madreporic canal loses
its connection with the exterior so as to hang loosely in the
interior, though the steps of this process do not appear to
have been made out.
The original hinder peritoneal vesicle grows rapidly,
and divides into two (fig. 248 C,
pi. and pr.}, which encircle the
two sides of the alimentary
canal, and meet above and
below it. The outer wall of
each of them attaches itself to
the skin, and the inner one to
the alimentary canal and watervascular system ; in both cases
the walls remain separated
from the adjacent parts by a
layer of the amoeboid cells
already spoken of. The cavity
of the peritoneal vesicles becomes the permanent body
cavity. Where the walls of
-ME
FIG. 249. LONGITUDINAL SECTION
THROUGH AN EMBRYO OF CUCUMARIA
DOLIOLUM AT THE END OF THE FOURTH
DAY.
Vpv. vaso-peritoneal vesicle; ME.
mesenteron; Blp., Ptd. blastopore, proctodaeum.
352
548 HOLOTHUROIDEA.
the two vesicles meet on the dorsal side, a mesentery, suspending the alimentary canal and dividing the body cavity longitudinally, is often formed. In other parts the partition walls
between the two sacks appear to be absorbed.
The amoeboid cells, which were derived from the invaginated
cells, arrange themselves as a layer round all the organs (fig.
249). Some of them remain amoeboid, attach themselves to the
skin, and form part of the cutis; and in these cells the calcareous spicula of the larva and adult are formed. Others
form the musculature of the larval alimentary tract, while the
remainder give rise to the musculature and connective tissue of
the adult.
The development of the vascular system is not known, but the discovery
of Kowalevsky, confirmed by Selenka, that from the walls of the watervascular system corpuscles are developed, identical with those in the bloodvessels, indicates that it probably develops in connection with the watervascular system. The observations of Hoffmann and Perrier on the communication of the two systems in the Echinoidea point to the same conclusion.
Though nothing very definite is known with reference to the development of
the nervous system, Metschnikoff suggests that it develops in connection
with the thickened bands of epiblast which are formed by a metamorphosis
of the ciliated bands of the embryo, and accompany the five radial tubes
(vide p. 555). In any case its condition in the adult leaves no doubt of its
being a derivative of the epiblast.
From the above description the following general conclusions
may be drawn :
(1) The blastosphere stage is followed by a gastrula stage.
(2) The gastrula opening forms the permanent anus, and the
mouth is formed by a fresh invagination.
(3) The mesoblast arises entirely from the invaginated cells,
but in two ways :
(a) As scattered amoeboid cells, which give origin to the
muscles and connective tissue (including the cutis) of the body
wall and alimentary tract.
(&) As a portion separated off from the archenteron,
which gives rise both to the epithelioid lining of the body cavity,
and of the water-vascular system.
(4) The oesophagus is derived from an invagination of the
epiblast, and the remainder of the alimentary canal from the
archenteron.
ECHINODERMATA. 549
(5) The embryonic systems of organs pass directly into those
of the adult.
The development of Synapta diverges, as might be expected, to a very
small extent from that of Holothuria.
Asteroidea. In Asterias the early stages of development conform to
our type. There arise, however, two bilaterally symmetrical vaso-peritoneal
diverticula from the archenteron. These diverticula give rise both to the
lining of the body cavity and water-vascular system. With reference to
the exact changes they undergo there is, however, some difference of opinion.
Agassiz (543) maintains that both vesicles are concerned in the formation of
the water-vascular system, while Metschnikoff (560) holds that the watervascular system is entirely derived from the anterior part of the larger left
vesicle, while the right and remainder of the left vesicle form the body
cavity. MetschnikofFs statements appear to be the most probable. The
anterior part of the left vesicle, after separating from the posterior, grows
into a five-lobed rosette (fig. 260, /), and a madreporic canal (h] with a dorsal
pore opening to the exterior. The rosette appears not to grow round the
oesophagus, as in the cases hitherto described. But the latter is stated to
disappear, and a new oesophagus to be formed, which pierces the rosette,
and places the old mouth in communication with the stomach. Except
where the anus is absent in the adult, the larval anus probably persists.
Ophiuroidea. The early development of the Ophiuroidea is not so
fully known as that of other types. Most species have a free-swimming
larva, but some (Amphiura) are viviparous.
The early stages of the free-swimming larvae have not been described,
but I have myself observed in the case of Ophiothrix fragilis that the
segmentation is uniform, and is followed by the normal invagination. The
opening of this no doubt remains as the larval anus, and there are probably
two outgrowths from this to form the vaso-peritoneal vesicles. Each of these
divides into two parts, an anterior lying close to the oesophagus, and a
posterior close to the stomach. The anterior on the right side aborts ; that
on the left side becomes the water-vascular vesicle, early opens to the
exterior, and eventually grows round the oesophagus, which, as in Holothurians, becomes the oesophagus of the adult. The posterior vesicles give rise
to the lining of the body cavity, but are stated by Metschnikoff to be at first
solid, and only subsequently to acquire a cavity the permanent body cavity.
The anus naturally disappears, since it is absent in the adult. In the
viviparous type the first stages are imperfectly known, but it appears that
the blastopore vanishes before the appearance of the mouth. The development of the ^vaso-peritoneal bodies takes place as in the free-swimming
larvae.
Echinoidea. In the Echinoidea (Agassiz, No. 542, Selenka, No. 564)
there is a regular segmentation and the normal invagination (fig. 250 A).
The amoeboid mesoblast cells arise as two laterally placed masses, and give
rise to the usual parts. The archenteron grows forward and bends towards
550
CRINOIDEA.
the ventral side (fig. 250 B). It becomes (fig. 250 C) divided into three
chambers, of which the two hindermost (d and c) form the stomach and
intestine ; while the anterior forms the oesophagus, and gives rise to the
FIG. 250. THREE SIDE VIEWS OF EARLY STAGES IN THE DEVELOPMENT OF
STRONGYLOCENTRUS. (From Agassiz.)
a, anus (blastopore) ; d. stomach ; o. oesophagus ; c . rectum ; w. vaso-peritoneal
vesicle ; v. ciliated ridge ; r. calcareous rod.
vaso-peritoneal vesicles. These latter appear as a pair of outgrowths
(fig. 251), but become constricted off as a single two-horned vesicle, which
subsequently divides into two. The left of these
is eventually divided, as in Asteroids, into a
peritoneal and water-vascular sack, while the
right forms the right peritoneal sack. An oral
invagination on the flattened ventral side meets
the mesenteron after its separation from the
vaso-peritoneal vesicle. The larval anus persists, as also does the larval mouth, but owing
to the manner in which the water-vascular
rosette is established the larval oesophagus appears to be absorbed, and to be replaced by a
fresh oesophagus.
Crinoidea. Antedon, the only Crinoid
so far studied (Gotte, No. 549), presents some
not inconsiderable variations from the usual
Echinoderm type. The blastopore is placed on
the somewhat flattened side of the oval blastosphere, and not, as is usual, at the hinder end.
The blastopore completely closes, and is not converted into the permanent anus. The archenteron gives rise to the epithelioid lining of both body
cavity and water-vascular system. These parts do not, however, appear as
a single or paired outgrowth from the archenteron, but as three distinct
outgrowths which are not formed contemporaneously. Two of them are first
FIG. -251. DORSO-VENTRAL VIEW OF AN EARLY
LARVA OF STRONGYLOCENTRUS. (From Agassiz.)
a. anus ; d. stomach ; o.
oesophagus ; w. vaso-peritoneal vesicle; r. calcareous
rod.
ECHINODERMATA.
551
formed and become the future body cavity; but their lumens remain distinct.
Jngmally appearing as lateral outgrowths, the right one assumes a dorsal
position and sends a prolongation into the stalk (fig. 252 rp'\ and
the left one assumes first a ventral, and then an oral position (fur
252 lp\
The third outgrowth of the archenteron gives rise to the water-vascular
vesicle. It first grows round the region of the future oesophagus and so
forms the water-vascular ring.
The wall of the ring then
grows towards the body wall
so as to divide the oral (left)
peritoneal vesicle into two
distinct vesicles, an anterior
and a posterior, shewn in fig.
253, lp' and lp. Before this
division is completed, the
water-vascular ring is produced in front into five pro
FIG. 252. LONGITUDINAL SECTION THROUGH
AN ANTEDON LARVA. (From Carpenter: after
Gotte.)
al. mesenteron ; -wv. water- vascular ring ;
lp. left (oral) peritoneal vesicle; rp. right peritoneal vesicle ; rp'. continuation of right vesicle
into the stalk ; st. stalk.
cessesthe future tentacles
(fig. 252, wv) which project
into the cavity of the oral
vesicle (lp\ After the oral
peritoneal space has become
completely divided into two parts, the anterior dilates (fig. 253, //) greatly,
and forms a large vestibule at the anterior end of the body. This vestibule
(lp'} next acquires a communication with the mesenteron, shewn in fig. 253
at m. The anterior wall of this vestibule is finally broken through. By this
rupture the mesenteron is placed in communication with the exterior by the
opening at m, while at the same time the tentacles of the water-vascular ring
(/) project freely to the exterior. Such is Gotte's account of the prge-oral
body space, but, as he himself points out, it involves our believing that the
lining of the diverticulum derived from the primitive alimentary vesicle
becomes part of the external skin. This occurrence is so remarkable, that
more evidence appears to me requisite before accepting it.
The formation of the anus occurs late. Its position appears to be the
same as that of the blastopore, and is indicated by a papilla of the mesenteron attaching itself to the skin on the ventral side (fig. 253, an). It eventually becomes placed in an interradial space within the oral disc of the adult.
The water-vascular ring has no direct communication with the exterior, but
the place of the madreporic canal of other types appears to be taken in
the larva by a single tube leading from the exterior into the body cavity, the
external opening of which is placed on one of the oral plates (vide p. 571) in
the next interradial space to the right of the anus, and a corresponding
diverticulum of the water-vascular ring opening into the body cavity. The
line of junction between the left and right peritoneal vesicles forms in the
larva a ring-like mesentery dividing the oral from the aboral part of the body
552
CRINOIDEA.
cavity. In the adult 1 the oral section of the larval body cavity becomes the
ventral part of the circumvisceral division of the body cavity, and the
subtentacular canals of the arms and disc ; while the aboral section becomes
the dorsal part of the circumvisceral division of the body cavity, the cceliac
canals of the arms, and the cavity of the centro-dorsal piece. The primitive
,+wr
FIG. 253. LONGITUDINAL SECTION THROUGH THE CALYX OF AN ADVANCED
PENTRACRINOID ANTEDON LARVA WITH CLOSED VESTIBULE.
(From Carpenter ; after Gotte.)
ae. epithelium of oral vestibule; ;//. mouth; al. mesenteron; an. rudiment of
permanent anus; lp. posterior part of left (oral) peritoneal sack; lp' '. anterior part of
left (oral) peritoneal sack; wr. water-vascular ring; /. tentacle; mt. mesentery;
rp. right peritoneal sack; rp '. continuation of right peritoneal sack into the stalk;
r. roof of tentacular vestibule.
distinction between the sections of the larval body cavity becomes to a large
extent obliterated, while the axial and intervisceral sections of the bodycavity of the adult are late developments.
The more important points in the development indicated in
the preceding pages are as follows :
(i) The blastosphere is usually elongated in the direction
of the axis of invagination, but in Comatula it is elongated
transversely to this axis.
1 Vide P. H. Carpenter, "On the genus Actinometra." Linnean Trans., and
Series, Zoology, Vol. n., Part I., 1879.
ECHINODERMATA. 553
(2) The blastopore usually becomes the permanent anus,
but it closes at the end of larval life (there being no anus in the
adult) in Ophiuroids and some Asteroids, while in Comatula it
closes very early, and a fresh anus is formed at the point where
the blastopore was placed.
(3) The larval mouth always becomes the mouth of the
adult.
(4) The archenteron always gives rise to outgrowths which
form the peritoneal membrane and water-vascular systems. In
Comatula there are three such outgrowths, two paired, which
form the peritoneal vesicles, and one unpaired, which forms the
water-vascular vesicle. In Asteroids and Ophiuroids there are
two outgrowths. In Ophiuroids both of these are divided into a
peritoneal and a water-vascular vesicle, but the right watervascular vesicle atrophies. In Asteroids only one water-vascular
vesicle is formed, which is derived from the left peritoneal vesicle.
In Echinoids and Holothuroids there is a single vaso-peritoneal
vesicle.
(5) The water- vascular vesicle grows round the larval
oesophagus in Holothuroids, Ophiuroids, and Comatula ; in
these cases the larval oesophagus is carried on into the adult.
In other forms the water-vascular vesicle forms a ring which
does not enclose the cesophagus (Asteroids and Echinoids);
in such cases a new oesophagus is formed, which perforates this
ring.
Development of the larval appendages and metamorphosis.
Holothuroidea. The young larva of Synapta, to which J.
Muller gave the name Auricularia (fig. 255), is in many respects
the simplest form of Echinoderm larva. With a few exceptions
the Auricularia type of larva is common to the Holothuria.
It is (fig. 254 A and fig. 255) bilaterally symmetrical, presenting a flattened ventral surface, and a convex dorsal one.
The anus (an) is situated nearly at the hinder pole, and the
mouth (m) about the middle of the ventral surface. In front
of the mouth is a considerable process, the prae-oral lobe.
Between the mouth and anus is a space, more or less concave
according to the age of the embryo, interrupted by a ciliated
554
AURICULARIA.
A similar ciliated ridge is
A E
ridge a little in front of the anus,
present on the ventral surface
of the prae-oral lobe immediately in front of the mouth.
The anal and oral ridges are
connected by two lateral ciliated bands, the whole forming
a continuous band, which,
since the mouth lies in the
centre of it (fig. 255), may be
regarded as a ring completely
surrounding the body behind
the mouth, or more naturally
as a longitudinal ring.
The bilateral Auricularia
is developed from a slightly
elongated gastrula with an uniform covering of cilia. The
gastrula becomes flattened on the oral side. At the same time
the cilia become specially developed on the oral and anal ridges,
and then on the remainder of the ciliated ring, while they are
FIG. 254. A. THE LARVA OF A HOLOTHUROID. B. THE LARVA OF AN ASTEROID.
- //. mouth; st. stomach; a. anus; l.c>
primitive longitudinal ciliated band; pr.c.
prae-oral ciliated band.
FIG. 155. DIAGRAMMATIC FIGURES REPRESENTING THE EVOLUTION OF AN
AURICULARIA FROM THE SIMPLEST ECHINODERM LARVAL FORM. (Copied from
MUller.)
The black line represents the ciliated ridge. The shaded part is the oral side of
the ring, the clear part the aboral side.
/;;. mouth; an. anus.
simultaneously obliterated elsewhere ; and so a complete Auricularia is developed. The water-vascular ring in the fully-developed
larva has already considerably advanced in the growth round the
oesophagus (fig. 256 w.v.r).
Most Holothurian larvae, in their transformation from the
bilateral Auricularia form to the radial form of the adult, pass
through a stage in which the cilia form a number of transverse
ECHINODERMATA.
555
-2>.v
rings, usually five in number, surrounding the body. The
stages in this metamorphosis are shewn in figs. 256, 257, and
258.
The primitive ciliated band,
at a certain stage of the metamorphosis, breaks up into a
number of separate portions
(fig. 256), the whole of which are
placed on the ventral surface.
Four of these (fig. 257 A and B)
arrange themselves in the form
of an angular ring round the
mouth, which at this period projects considerably. The remaining portions of the primitive
band change their direction from
a longitudinal one to a transverse (fig. 257 B), and eventually
grow into complete rings (fig.
2570). Of these there are five.
The middle one (257 B) is the
first to develop, and is formed
from the dorsal parts of the
primitive ring. The two hinder
rings develop next, and last of
all the two anterior ones, one of
which appears to be in front of the mouth (fig. 257 C).
The later development of the mouth, and of the ciliated ridge
surrounding it, is involved in some obscurity. It appears from
Metschnikoff (No. 560) that an invagination of the oesophagus
takes place, carrying with it the ciliated ridge around the mouth.
This ridge becomes eventually converted into the covering for
the five tentacular outgrowths of the water- vascular ring (fig.
258), and possibly also forms the nervous system.
The opening of the cesophageal invagination is at first behind
the foremost ciliated ring, but eventually comes to lie in front of
it, and assumes a nearly terminal though slightly ventral position
(fig. 258). No account has been given of the process by which
this takes place, but the mouth is stated by Metschnikoff (though
FIG. 256. FULL-GROWN LARVA OF
SYNAPTA. (After Metschnikoff.)
m. mouth ; st. stomach ; a. anus ;
p.v. left division of perivisceral cavity,
which is still connected with the watervascular system ; w.v.r. water-vascular
ring which has not yet completely encircled the oesophagus; I.e. longitudinal
part of ciliated band ; pr.c. prae-oral part
of ciliated band.
556
BIPINNARIA.
Miiller differs from him on this point) to remain open throughout. The further changes in the metamorphosis are not considerable. The ciliated bands disappear, and a calcareous ring
of ten pieces, five ambulacral and five interambulacral, is formed
round the oesophagus. A provisional calcareous skeleton is also
developed.
All the embryonic systems of organs pass in this case
directly into those of the adult.
The metamorphosis of most Holothuroidea is similar to that just
described. In Cucumaria (Selenka) there is however no Auricularia stage,
and the uniformly ciliated stage is succeeded by one with five transverse
FIG. 257. THREE STAGES IN THE DEVELOPMENT OF SYNAPTA. A and B
are viewed from the ventral surface, and C from the side. (After Metschnikoff.)
m. mouth; oe. oesophagus; pv. walls of the perivisceral cavity; wv. longitudinal
vessel of the water- vascular system; p. dorsal pore of water-vascular system;
cr. ciliated ring formed round the mouth from parts of the primitive ciliated
band.
bands of cilia, and a prae-oral and an anal ciliated cap. The mouth is at
first situated ventrally behind the prse-oral cap of cilia, but the prae-oral
cap becomes gradually absorbed, and the mouth assumes a terminal
position.
In Psolinus (Kowalevsky) there is no embryonic ciliated stage, and the
adult condition is attained without even a metamorphosis. There appear to
ECHINODERMATA.
557
be five plates surrounding the
mouth, which are developed before
any other part of the skeleton, and
are regarded by P. H. Carpenter
(No. 548) as equivalent to the five
oral plates of the Crinoidea. The
larval condition with ciliated bands
is often spoken of as the pupa stage,
and during it the larvae of Holothurians proper use their embryonic
tube feet to creep about.
Asteroidea. The commonest and most thoroughly
investigated form of Asteroid
larva is a free swimming form
known as Bipinnaria.
This form in passing from
the spherical to the bilateral
condition passes through at
first almost identical changes
to the Auricularian larva.
The cilia become at an early
period confined to an oral
and anal ridge.
The anal ridge gradually extends dorsalwards, and finally
forms a complete longitudinal post-oral ring (fig. 259 A) ; the
oral ridge also extends dorsalwards, and forms a closed prae-oral
ring (fig. 259 A), the space within which is left unshaded in my
figure.
The presence of two rings instead of one distinguishes the
Bipinnaria from the Auricularia. The two larvae are shewn side
by side in fig. 254, and it is obvious that the two bands of the
Bipinnaria are (as pointed out by Gegenbaur) equivalent to the
single band of the Auricularia divided into two. Ontologically,
however, the two bands of Bipinnaria do not appear to arise
from the division of a single band.
As the Bipinnaria grows older, a series of arms grows out
along lines of the two ciliated bands (fig. 259 C), and, in many
cases, three special arms are formed, not connected with the
ciliated bands, and covered with warts. These latter arms are
FlG. 258. A LATE STAGE IN THE DEVELOPMENT OF SYNAPTA. (After Metschnikoff.)
The figure shews the vestibular cavity
with retracted tentacles ; the ciliated bands ;
the water-vascular system, etc.
p. dorsal pore of water-vascular system ;
pv. walls of perivisceral cavity; ms. amoeboid cells.
558
BIPINNARIA.
known as brachiolar arms, and the larvae provided with them
as Brachiolaria (fig. 259 D).
As a rule the following arms can be distinguished (fig. 259 C and D), on
the hinder ring (Agassiz' nomenclature) a median anal pair, a dorsal anal
pair, and a ventral anal pair, a dorsal oral pair, and an unpaired anterior
dorsal arm ; on the prae-oral ring a ventral oral pair, and sometimes (Miiller)
an unpaired anterior ventral arm.
The three brachiolar arms arise as processes from the base of the
unpaired dorsal arm, and the two ventral oral arms. The extent of the
development of the arms varies with the species.
FIG. 259. DIAGRAMMATIC REPRESENTATION OF VARIOUS FORMS OF ASTEROID
LARWE. A, B, C, BIPINNARIA; D, BRACHIOLARIA. (Copied from Muller.)
The black lines represent the ciliated bands ; and the shading the space between
the prae-oral and the post-oral bands.
m. mouth; an. anus.
The changes by which the Bipinnaria or Brachiolaria becomes
converted into the adult starfish are very much more complicated
than those which take place in Holothurians. For an accurate
knowledge of them we are largely indebted to Alex. Agassiz
(No. 543). The development of the starfish takes place entirely
at the posterior end of the larva close to the stomach.
On the right and dorsal side of the stomach, and externally
to the rig/it peritoneal space, are formed five radially situated
calcareous rods arranged in the form of a somewhat irregular
pentagon. The surface on which they are deposited has a
spiral form, and constitutes together with its calcareous rods, the
ECHINODERMATA. 559
abactinal or dorsal surface of the future starfish. Close to its
dorsal, i.e. embryonic dorsal, edge lies the dorsal pore of the
water-vascular system (madreporic canal), and close to its ventral
edge the anus. On the left and ventral side of the stomach is
placed the water-vascular rosette, the development of which was
described on p. 549. It is situated on the actinal or ventral surface
of the future starfish, and is related to the left peritoneal vesicle.
Metschnikoff (No. 560) and Agassiz (No. 543) differ slightly as to the
constitution of the water- vascular rosette. The former describes and figures
it as a completely closed rosette, the latter states that ' it does not form a
completely closed curve but is always open, forming a sort of twisted
crescent-shaped arc.'
The water-vascular rosette is provided with five lobes, corresponding to which are folds in the larval skin, and each lobe
corresponds to one of the calcareous plates developed on the
abactinal disc. The plane of the actinal surface at first meets
that of the abactinal at an acute or nearly right angle. The two
surfaces are separated by the whole width of the stomach. The
general appearance of the larva from the ventral surface after
the development of the water-vascular rosette (i) and abactinal
disc (A) is shewn in fig. 260.
As development proceeds the abactinal surface becomes a
firm and definite disc, owing to the growth of the original
calcareous spicules into more or less definite plates, and to the
development of five fresh plates nearer the centre of the disc and
interradial in position. Still later a central calcareous plate
appears on the abactinal surface, which is thus formed of a
central plate, surrounded by a ring of five interradial plates, and
then again by a ring of five radial plates. The abactinal disc
now also grows out into five short processes, separated by five
shallow notches. These processes are the rudiments of the five
arms, and each of them corresponds to one of the lobes of the
water-vascular rosette. A calcareous deposit is formed round
the opening of the water-vascular canal, which becomes the
madreporic tubercle 1 . At about this stage the absorption of the
larval appendages takes place. The whole anterior part of the
1 The exact position of the madreporic tubercle in relation to the abactinal plates
does not seem to have been made out. It might have been anticipated that it would
be placed in one of the primary interradial plates, but this does not seem to be the
case. The position of the anus is also obscure.
5 6o
BIPINNARIA.
larva with the great prae-oral lobe has hitherto remained
unchanged, but now it contracts and undergoes absorption, and
becomes completely withdrawn into the disc of the future starfish.
The larval mouth is transported into
the centre of the actinal disc. In the
larvae observed by Agassiz and Metschnikoff nothing was cast off, but the
whole absorbed.
According to M tiller and Koren and
Danielssen this is not the case in the larva
observed by them, but part of the larva is
thrown off, and lives for some time independently.
After the absorption of the larval
appendages the actinal and abactinal
surfaces of the young starfish approach
each other, owing to the flattening of
the stomach ; at the same time they
lose their spiral form, and become flat
discs, which fit each other. Each of
the lobes of the rosette of the watervascular system becomes one of the
radial water-vascular canals. It first
becomes five-lobed, each lobe forming
a rudimentary tube foot, and on each ^ d ctinal disc of youn Aste '
side of the middle lobe two fresh ones
next spring out, and so on in succession. The terminal median
lobe forms the tentacle at the end of the arm, and the eye is
developed at its base. The growth of the water-vascular canals
keeps pace with that of the arms, and the tube feet become
supported at their base by an ingrowth of calcareous matter.
The whole of the calcareous skeleton of the larva passes directly
into that of the adult, and spines are very soon formed on the
plates of the abactinal surface. The original radial plates,
together with the spines which they have, are gradually pushed
outwards with the growth of the arms by the continual addition
of fresh rows of spines between the terminal plate and the plate
next to it. It thus comes about that the original radial plates
persist at the end of the arms, in connection with the unpaired
FIG. 260. BIPINNARIA
LARVA OF AN ASTEROID. (From
Gegenbaur ; after Miiller.)
b. mouth ; a. anus ; h. madreporic canal ; t. ambulacral
rosette ; c . stomach ; d. g. e.
etc. arms of Bipinnaria ; A.
ECHINODERMATA. 561
tentacles which form the apex of the radial water-vascular
tubes.
It has already been mentioned that according to Metschnikoff (No. 560)
a new oesophagus is formed which perforates the water-vascular ring, and
connects the original stomach with the original mouth. Agassiz (No. 543)
maintains that the water-vascular ring grows round the primitive oesophagus.
He says " During the shrinking of the larva the long oesophagus becomes
" shortened and contracted, bringing the opening of the mouth of the larva
" to the level of the opening of the oesophagus, which eventually becomes
"the true mouth of the starfish." The primitive anus is believed by
Metschnikoff to disappear, but by Agassiz to remain. This discrepancy
very possibly depends upon these investigators having worked at different
species.
There is no doubt that the whole of the larval organs, with
the possible exception of the oesophagus, and anus (where absent
in the adult), pass directly into the corresponding organs of the
starfish and that the prae-oral part of the body and arms of the
larva are absorbed and not cast off.
In addition to the Bipinnarian type of Asteroid larva a series of other
forms has been described by Miiller (No. 561), Sars, Keren, and Danielssen
(No. 554) and other investigators, which are however very imperfectly
known. The best-known form is one first of all discovered by Sars in
Echinaster Sarsii, and the more or less similar larvae subsequently investigated by Agassiz, Busch, Miiller, Wyville Thomson, etc. of another species
of Echinaster and of Asteracanthion. These larvae on leaving the egg have
an oval form, and are uniformly covered by cilia. Four processes (or in
Agassiz' type one process) grow out from the body ; by these the larvae fix
themselves. In the case of Echinaster the larvae are fixed in the ventral
concavity of the disc of the mother, between the five arms, where a temporary brood-pouch is established. The main part of the body is converted
directly into the disc of the young starfish, while the four processes come to
spring from the ventral surface, and are attached to the water- vascular ring.
Eventually they atrophy completely. Of the internal structure but little is
known ; till the permanent mouth is formed, after the development of the
young starfish is pretty well advanced, the stomach has no communication
with the exterior.
A second abnormal type of development is presented by the embryo of
Pteraster miliaris, as described by Koren and Danielssen 1 . The larvae to
the number of eight to twenty develop in a peculiar pouch on the dorsal
surface of the body. The early stages are not known, but in the later ones
the whole body assumes a pentagonal appearance with a mouth at one edge
1 The following statements are taken from the abstract in Bronn's Thierreichs.
B. II. 36
5 62
OPHIUROID PLUTEUS.
of the disc. At a later stage the anus is formed on the dorsal side of an arm
opposite the mouth. The stomach is surrounded by a water-vascular ring,
from which the madreporic canal passes to the dorsal surface, but does not
open. At a later stage the embryonic mouth and anus vanish, to be replaced
by a permanent mouth and anus in the normal positions.
A third, and in some respects very curious, form is a worm like larva of
Miiller, which is without bands of cilia. The dorsal surface of the youngest
larva is divided by transverse constrictions into five segments. On the
under side of the first of these is a five-lobed disc, each lobe being provided
with a pair of tube feet.
At a later period only three segments are visible on the dorsal surface,
but the ventral surface has assumed a pentagonal aspect. The later stages
are not known.
Ophiuroidea. The full-grown larva of the Ophiuroids is
known as a Pluteus. It commences with the usual more or less
spherical form ; from this it passes to a form closely resembling
FIG. 261. DIAGRAMMATIC FIGURES SHEWING THE EVOLUTION OK AN OPHIUROID PLUTEUS FROM A SIMPLE ECHINODERM LARVA. (Copied from Miiller.) The
calcareous skeleton is not represented.
///. mouth; an. anus; d. anterior arms; d'. lateral arms; e'. posterior arms; tf.
anterolateral arms.
that of Auricularia with a rounded dorsal surface, and a flattened
ventral one. Soon however it becomes distinguished by the
growth of a post-anal lobe and the absence of a prae-oral lobe
(fig. 261 B). The post-anal lobe forms the somewhat rounded
apex of the body. In front of the mouth, and between the
mouth and anus, arise the anal and oral ciliated ridges, which
soon become continued into a single longitudinal ciliated ring.
At the same time the body becomes prolonged into a series of
ECHINODERMATA.
563
processes along the ciliated band, which is continued to the
extremity of each. The primitive ciliated ring never becomes
broken up into two or more rings. A ciliated crown is usually
developed at the extremity of the post-anal lobe. The arms are
arranged in the form of a ring surrounding the mouth, and are
all directed forwards.
The first arms to appear are two lateral ones, which usually remain the
most conspicuous (fig. 261 B and C, cf\ Next arises a pair on the sides of
the mouth, which may be called the mouth or anterior arms (C, d}. A pair
ventral to and behind the lateral arms is then formed, constituting the
posterior arms (D, e'\ and finally a pair between the lateral arms and the
anterior, constituting the anterolateral arms (D,^).
The concave area between the arms forms the greater part of
the ventral surface of the body. Even before the appearance of
any of the arms, and before the formation of the mouth, two
calcareous rods are formed, which meet behind at the apex of
the post-anal lobe, and are continued as a central support into
each of the arms as they are successively formed. These rods
are shewn at their full development in fig. 262. The important
points which distinguish a Pluteus
larva from the Auricularia or
Bipinnaria are the following :
(i) The presence of the postanal lobe at the hind end of the
body. (2) The slight development of a prae-oral lobe. (3) The
provisional calcareous skeleton in
the larval arms.
Great variations are presented
in the development of the arms
and provisional skeleton. The
presence of lateral arms is however
a distinctive characteristic of the
Ophiuroid Pluteus. The other
arms may be quite absent, but
the lateral arms never.
The formation of the permanent Ophiuroid takes place in
much the same way as in the Asteroidea.
36-2
FIG. 262.
OPHIUROID.
after Miiller.)
PLUTEUS LARVA OF AN
(From Gegenbaur ;
A. rudiment of young Ophiuroid ;
(?. lateral arms; d. anterior arms;
e . posterior arms.
564
OPHIUROID PLUTEUS.
There is formed (fig. 262) on the right and dorsal side of stomach the
abactinal disc supported by calcareous plates, at first only five in number
and radial in position 1 . The disc is at first not symmetrical, but becomes so
at the time of the resorption of the larval arms. It grows out into five
processes the five future rays. The original five radial plates remain as the
terminal segments of the adult rays, and new plates are always added
between the ultimate and penultimate plate (Mu'ller), though it is probable
that in the later stages fresh plates are added in the disc.
The ventral surface of the permanent Ophiuroid is formed by the concave
surface between the mouth and anus. Between this and the stomach is
FIG. 263. DIAGRAMMATIC FIGURES SHEWING THE EVOLUTION OF ECHINOID
PLUTEI. (Copied from Miiller.) The calcareous skeleton is not represented. E.
Pluteus of Spatangus.
m. mouth; an. anus; d. anterior arms; d' . point where lateral arms arise in the
Ophiuroid Pluteus; e. anterointernal arms; e. posterior arms; g'. anterolateral arms;
g. anteroexternal arms.
situated the water-vascular ring. It is at first not closed, but is horseshoeshaped, with five blind appendages (fig. 262). It eventually grows round
the cesophagus, which, together with the larval mouth, is retained in the
adult. The five blind appendages become themselves lobed in the same
way as in Asterias, and grow out along the five arms of the disc and become
the radial canals and tentacles. All these parts of the water-vascular system
are of course covered by skin, and probably also surrounded by mesoblast
cells, in which at a later period the calcareous plates which lie ventral to the
radial canal are formed. The larval anus disappears. As long as the larval
appendages are not absorbed the ventral and dorsal discs of the permanent
Ophiuroid fit as little as in the case of the Brachiolaria, but at a certain
period the appendages are absorbed. The calcareous rods of the larval arms
1 Whether interradial plates are developed as in Asterias is not clear. They seem
to be found in Ophiopholis bellis, Agassiz, but have not been recognised in other
forms (vide Carpenter, No. 548, p. 369).
ECHINODERMATA. 565
break up, the arms and anal lobe become absorbed, and the dorsal and
ventral discs, with the intervening stomach and other organs, are alone left.
After this the discs fit together, and there is thus formed a complete young
Ophiuroid.
The whole of the internal organs of the larva (except the anus), including
the mouth, cesophagus, the body cavity, etc. are carried on directly into the
adult.
The larval skeleton is, as above stated, absorbed.
The viviparous larva of Amphiura squamata does not differ very greatly
from the larvae with very imperfect arms. It does not develop a distinct
ciliated band, and the provisional skeleton is very imperfect. The absence
of these parts, as well as of the anus, mentioned on p. 549, may probably be
correlated with the viviparous habits of the larva. With reference to the
passage of this larva into the adult there is practically nothing to add to
what has just been stated. When the development of the adult is fairly
advanced the part of the body with the provisional skeleton forms an
elongated rod-like process attached to the developing disc. It becomes
eventually absorbed.
Echinoidea. The Echinus larva (fig. 263} has a Pluteus
form like that of the Ophiuroids, and in most points, such as the
FIG. 264. Two LARV/E OF STRONGYLOCENTRUS. (From Agassiz.)
m. mouth; a. anus; o. cesophagus; d. stomach; c. intestine; '. and v. ciliated
ridges; iv. water- vascular tube; r. calcareous rods.
presence of the anal lobe, the ciliated band, the provisional
skeleton, etc., develops in the same manner. The chief difference
between the two Pluteus forms concerns the development of the
lateral arms. These, which form the most prominent arms in
the Ophiuroid Pluteus, are entirely absent in the Echinoid
5 66
ECHINOID PLUTEUS.
Pluteus, which accordingly has, as a rule, a much narrower form
than the Ophiuroid Pluteus.
A pair of ciliated epaulettes on each side of and behind the
ciliated ring is very characteristic of some Echinoid larvae.
They are originally developed from the ciliated ring (fig. 266 A
FIG. 265. LATERAL AND VENTRAL VIEW OF A LARVA OF STRONGYLOCENTRUS.
(From Agassiz.) General references as in fig. 264.
b. dorsal opening of madreporic canal; e '. posterior arms ; e'". anterior arms;
f lV . anterointernal arms.
and B, z>"). The presence of three processes from the anal lobe
supported by calcareous rods is characteristic of the Spatangoid
Pluteus (fig. 263 E).
The first two pairs of arms to develop, employing the same names as in
Ophiuroids, are the anterior attached to the oral process (fig. 263 C, d] and
the posterior pair (*?') A pair of anterolateral arms next becomes developed
(j^). A fourth pair (not represented in Ophiuroids) appears on the inner
side of the anterior pair forming an anterointernal pair (e}, and in the
Spatangoid Pluteus a fifth pair may be added on the external side of the
anterior pair forming an anteroexternal pair (g).
Each of the first-formed paired calcareous rods is composed of three
processes, two of which extend into the anterior and posterior arms ; and the
third and strongest passes into the anal lobe, and there meets its fellow
(fig. 265). A transverse bar in front of the arms joins the rods of the two
sides meeting them at the point where the three processes diverge. The
process in the anterolateral arm (fig. 266 B) is at first independent of this
system of rods, but eventually unites with it. Although our knowledge of
ECHINODERMATA. 567
the Pluteus types in the different groups is not sufficient to generalise with
great confidence, a few points seem to have been fairly determined 1 . The
Plutei of Strongylocentrus (figs. 266 and 267) and Echinus have eight arms
and four ciliated epaulettes. The only Cidaris-like form, the Pluteus of
which is known, is Arbacia : it presents certain peculiarities. The anal lobe
develops a pair of posterior (auricular) appendages, and the ciliated ring,
besides growing out into the normal eight appendages, has a pair of short
blunt anterior and posterior lobes. An extra pair of non-ciliated accessory
mouth arms appears also to be developed. Ciliated epaulettes are not
present. So far as is known the Clypeastroid larva is chiefly characterized
by the round form of the anal lobe. The calcareous rods are latticed. In the
Pluteus of Spatangoids there are (fig. 263) five pairs of arms around the
mouth pointing forwards, and three arms developed from the anal lobe
pointing backwards. One of these is unpaired, and starts from the apex of
the anal lobe. All the arms have calcareous rods which, in the case of the
posterior pair, the anterolateral pair, and the unpaired arm of the anal lobe,
are latticed. Ciliated epaulettes are not developed.
Viviparous larvae of Echinoids have been described by Agassiz 2 .
The development of the permanent Echinus has been chiefly worked out
by Agassiz and Metschnikoff.
In the Pluteus of Echinus lividus the first indication of the adult arises,
when three pairs of arms are already developed, as an invagination of the
skin on the left side, between the posterior and anterolateral arms, the
bottom of which is placed close to the water-vascular vesicle (fig. 266 B, u/\
The base of this invagination becomes very thick, and forms the ventral disc
of the future Echinus. The parts connecting this disc with the external
skin become however thin, and, on the narrowing of the external aperture of
invagination and the growth of the thickened disc, constitute a covering for
the disc, called by Metschnikoff the amnion. The water- vascular vesicle
adjoining this disc grows out into five processes, forming as many tube feet,
which cause the surface of the involuted disc to be produced into the same
number of processes. The external opening of the invagination of the disc
never closes, and after the development of the tube feet begins to widen
again, and the amnion to atrophy. Through the opening of the invagination
the tube feet now project. The dorsal and right surface of the Pluteus,
which extends so as to embrace the opening of the madreporic canal and
the anus, forms the abactinal or dorsal surface of the future Echinus
(fig. 267, a). This disc fits on to the actinal invaginated surface which arises
on the left side of the Pluteus. On the right surface of the larva (dorsal of
permanent Echinus) two pedicellariae appear, and at a later period spines
are formed, which are at first arranged in a ring-like form round the edge of
the primitively flat test. While these changes are taking place, and the two
surfaces of the future Echinus are gradually moulding themselves so as to
1 Vide especially Muller, Agassiz, and Metschnikoff.
2 For viviparous Echini vide Agassiz, Proc. Amer. Acad. 1876.
5 68
ECHINOID PLUTEUS.
form what is obviously a young Echinus, the arms of the Pluteus with their
contained skeleton have been gradually undergoing atrophy. They become
irregular in form, their contained skeleton breaks up into small pieces, and
they are gradually absorbed.
The water-vascular ring is from the first complete, so that, as in
Asterias, it is perforated through the centre by a new oesophagus. According
FIG. 266. SIDE AND DORSAL VIEW OF A LARVA OF STRONGYLOCENTRUS.
(From Agassiz.) General reference letters as in figs. 264 and 265.
e" . anterolateral arms; v" '. ciliated epaulettes; ?&'. invagination to form the disc
of Echinus.
to Agassiz the first five tentacles or tube feet grow into the radial canals,
and form the odd terminal tentacles exactly as in Asterias 1 . Spatangus
only differs in development from Echinus in the fact that the opening of the
invagination to form the ventral disc becomes completely closed, and that
the tube feet have eventually to force their way through the larval epidermis
of the amnion, which is ruptured in the process and eventually thrown
off.
Crinoidea. The larva of Antedon, while still within the
egg-shell, assumes an oval form and uniform ciliation. Before it
1 Gotte (No. 549) supported by Muller's and Krohn's older, and in some points
extremely erroneous observations, has enunciated the view that the radial canals in
Echinoids and Holothuroids have a different nature from those in Asteroids and
Ophiuroids.
ECHINODERMATA.
569
becomes hatched the uniform layer of cilia is replaced by four
transverse bands of cilia, and a tuft of cilia at the posterior
extremity. In this condition it escapes from the egg-shell
FIG. 267. FULL-GROWN LARVA OF STRONGYLOCENTRUS. (From Agassiz.)
The figure shews the largely-developed abactinal disc of the young Echinus
enclosing the larval stomach. Reference letters as in previous figs.
(fig. 268 A), and becomes bilateral, owing to a flattening of the
ventral surface. On the flattened surface appears a ciliated
570
CRINOID LARVA.
depression corresponding in position with the now closed blastopore (vide p. 550). The third ciliated band bends forward
to pass in front of this (fig. 269). Behind the last ciliated band
there is present a small depression of unknown function, also
FIG. 768. THREB STAGES IN THE DEVELOPMENT OF ANTEDON (COMATULA.)
(From Lubbock; after Thomson.)
A. larva just hatched; B. larva with rudiment of the calcareous plates; C. Pentacrinoid larva.
ECHINODERMATA.
571
situated on the ventral surface. The posterior extremity of the
embryo elongates to form the rudiment of the future stem, and
a fresh depression, marking the position of the future mouth,
makes its appearance on the anterior and ventral part.
While the ciliated bands are still at their full development,
the calcareous skeleton of the future calyx makes its appearance
in the form of two rows, each of five plates, formed of a network
of spicula (figs. 268 B and 269). The plates of the anterior ring
are known as the orals, those of the posterior as the basals.
The former surround the left, i.e. anterior
peritoneal sack ; the latter the right, i.e.
posterior peritoneal sack. The two rows
of plates are at first not quite transverse,
but form two oblique circles, the dorsal
end being in advance of the ventral.
The rows soon become transverse, while
the originally somewhat ventral oral
surface is carried into the centre of the
area enclosed by the oral plates.
By the change in position of the
original ventral surface relatively to the
axis of the body, the bilateral symmetry
of the larva passes into a radial symmetry. While the first skeletal elements
of the calyx are being formed, the
skeleton of the stem is also established.
The terminal plate is first of all established, then the joints, eight at first, of
the stem. The centro-dorsal plate is
stated by Thomson to be formed as the
uppermost joint of the stem 1 . The larva, after the completion
of the above changes, is shewn in fig. 268 B, and somewhat more
diagrammatically in fig. 269.
After the above elements of the skeleton have become established the ciliated bands undergo atrophy, and shortly after
1 Gotte (No. 549) on the other hand holds that the centro-dorsal plate is developed
by the coalescence of a series of at first independent rods, which originate simultaneously with, and close to, the lower edges of the basals, and that it is therefore
similar in its origin to the basals.
FIG. 269. LARVA OF
ANTEDON WITH RUDIMENTS
OF CALCAREOUS SKELETON.
(From Carpenter; after
Thomson.)
i. Terminal plate at the
end of the stem ; 3. basals ;
or. orals ; bl. position of blastopore.
572
CRINOID LARVA.
wards the larva becomes attached by the terminal plate of its
stem. It then passes into the Pentacrinoid stage! The larva in
this stage is shewn in fig. 268 C and fig. 270. New joints are
added at the upper end of the stem next the calyx, and a new
element the radials makes its appearance as a ring of five
small plates, placed in the space between the basals and orals,
and in the intervals alternating with them
(fig. 270, 4). The roof of the oral vestibule (vide fig. 253 and p. 551) has in
the meantime become ruptured ; and
the external opening of the mouth thus
becomes established. Surrounding the
mouth are five petal-like lobes, each of
them supported by an oral plate (fig.
268 C). In the intervals between them
five branched and highly contractile tentacles, which were previously enclosed
within the vestibule, now sprout out :
they mark the position of the future
radial canals, and are outgrowths of the
water-vascular ring. At the base of each
of them a pair of additional tentacles is
soon formed. Each primary tentacle corresponds to one of the radials. These
latter are therefore, as their name implies,
radial in position; while the basals and
orals are interradial. In addition to the
contractile radial tentacles ten non-contractile tentacles, also diverticula of the
water- vascular ring, are soon formed, two
for each interradius.
In the course of the further development the equatorial space between the FlG - 2 7<>. YOUNG PEN
. TACRINOID LARVA OF AN
orals and the basals enlarges, and gives TEDON. (From Carpenter ;
rise to a wide oral disc, the sides of which after w >' ville Thom s"-)
- , , . ... . i. terminal plate of stem;
are formed by the radials resting on the c d. centro-donal plate; 3 .
basals; while in the centre of it are bftsals J 4- radials; or. orals.
placed the five orals, each with its special lobe.
The anus, which is formed on the ventral side in the position
ECHINODERMATA. 573
of the blastopore (p. 551), becomes surrounded by an anal plate,
which is interradial in position, and lies on the surface of the
oral disc between the orals and radials. On the oral plate in
the next interradius is placed the opening of a single funnel
leading into the body cavity, which Ludwig regards as equivalent to the opening of the madreporic canal (vide p. 55 1) 1 .
From the edge of the vestibule the arms grow out, carrying
with them the tentacular prolongation of the water-vascular ring.
Two additional rows of radials are soon added.
The stalked Pentacrinoid larva becomes converted, on the
absorption of the stalk, into the adult Antedon. The stalk is
functionally replaced by a number of short cirri springing from
the centro-dorsal plate. The five basals coalesce into a single
plate, known as the rosette, and the five orals disappear, though
the lobes on which they were placed persist. In some stalked
forms, e.g. Rhizocrinus Hyocrinus, the orals are permanently
retained. The arms bifurcate at the end of the third radial, and
the first radial becomes in Antedon rosacea (though not in all
species of Antedon) concealed from the surface by the growth of
the centro-dorsal plate. An immense number of funnels, leading
into the body cavity, are formed in addition to the single one
present in the young larva. These are regarded by Ludwig as
equivalent to so many openings of the madreporic canal ; and
there are developed, in correspondence with them, diverticula of
the water-vascular ring.
Comparison of Echinoderm Larvce and General Conclusions.
In any comparison of the various types of Echinoderm larvae
it is necessary to distinguish between the free-swimming forms,
and the viviparous or fixed forms. A very superficial examination suffices to shew that the free-swimming forms agree very
much more closely amongst themselves than the viviparous
1 I have made no attempt to discuss the homologies of the plates of the larval
Echinodermata because the criteria for such a discussion are still in dispute. The
suggestive memoirs of P. H. Carpenter (No. 548) on this subject may be consulted by
the reader. Carpenter attempts to found his homologies on the relation of the plates
to the primitive peritoneal vesicles, and I am inclined to believe that this method of
dealing with these homologies is the right one. Ludwig (No. 559) by regarding the
opening of the madreporic canal as a fixed point has arrived at very different results.
574
COMPARISON OF ECHINODERM LARV.-E.
forms. We are therefore justified in concluding that in the
viviparous forms the development is abbreviated and modified.
All the free forms are nearly alike in their earliest stage after
the formation of the archenteron. The surface between the
anus and the future mouth becomes flattened, and (except in
Antedon, Cucumaria, Psolinus, etc. which practically have an
abbreviated development like that of the viviparous forms) a
ridge of cilia becomes established in front of the mouth, and a
second ridge between the mouth and the anus. This larval
form, which is shewn in fig. 264 A, is the type from which the
various forms of Echinoderm larvae start.
In all cases, except in Bipinnaria, the two ciliated ridges
soon become united, and constitute a single longitudinal postoral ciliated ring.
The larvae in their further growth undergo various changes,
and in the later stages they may be divided into two groups :
(1) The Pluteus larva of Echinoids and Ophiuroids.
(2) The Auricularia (Holothuroids) and Bipinnaria (Asteroids) type.
The first group is characterized by the growth of a number
of arms more or less surrounding the mouth, and supported
by calcareous rods. The ciliated band retains its primitive
condition as a simple longitudinal band throughout larval life.
There is a very small prae-oral lobe, while an anal lobe is very
largely developed.
The Auricularia and Bi- A. B
pinnaria resemble each other
in shape, in the development
of a large prae-oral lobe, and
in the absence of provisional
calcareous rods ; but differ in
the fact that the ciliated band
is single in Auricularia (fig
271 A), and is double in Bipinnaria (fig. 271 B).
TheBipinnarialarvashews
THUROID. B. THE LARVA OF AN ASTEa great tendency to develop RIAS.
soft arms; while in the Auri- . ' mouth; st. stomach; a. anus; I.e.
, . ,_, , *_ 1-1- primitive longitudinal ciliated band; pr.c.
cularia the longitudinal ciliat- p r3 e-oral ciliated band.
FlG
THE LARVA OF A
ECHINODERMATA. 575
ed band breaks up into a number of transverse ciliated bands.
This condition is in .some instances reached directly, and such
larvae undoubtedly approximate to the larvae of Antedon, in
which the uniformly ciliated condition is succeeded by one with
four transverse bands, of which one is prae-oral.
All or nearly all Echinoderm larvae are bilaterally symmetrical,
and since all Echinodermata eventually attain a radial symmetry, a change necessarily takes place from the bilateral to the
radial type.
In the case of the Holothurians and Antedon, and generally
in the viviparous types, this change is more or less completely
effected in the embryonic condition ; but in the Bipinnaria and
Pluteus types a radial symmetry does not become apparent till
after the absorption of the larval appendages. It is a remarkable fact, which seems to hold for the Asteroids, Ophiuroids, Echinoids, and Crinoids, that the dorsal side of the larva is
not directly converted into the dorsal disc of the adult; but
the dorsal and right side becomes the adult dorsal or abactinal
surface, while the ventral and left becomes the actinal or ventral
surface.
It is interesting to note with reference to the larvae of the
Echinodermata that the various existing types of larvae must
have been formed after the differentiation of the existing groups
of the Echinodermata ; otherwise it would be necessary to adopt
the impossible position that the different groups of Echinodermata were severally descended from the different types of larvae.
The various special appendages, etc. of the different larvae have
therefore a purely secondary significance; and their atrophy
at the time of the passage of the larva into the adult, which
is nothing else but a complicated metamorphosis, is easily explained.
Originally, no doubt, the transition from the larva to the
adult was very simple, as it is at present in most Holothurians ;
but as the larvae developed various provisional appendages, it
became necessary that these should be absorbed in the passage
to the adult state.
It would obviously be advantageous that their absorption
should be as rapid as possible, since the larva in a state of
transition to the adult would be in a very disadvantageous
576 COMPARISON OF ECHINODERM
position. The rapid metamorphosis, which we find in Asteroids,
Ophiuroids, and Echinoids in the passage from the larval to the
adult state, has no doubt arisen for this reason.
In spite of the varying provisional appendages possessed by
Echinoderm larvae it is possible, as stated above (p. 574), to
recognise a type of larva, of which all the existing Echinoderm
larval forms are modifications. This type does not appear to
me to be closely related to that of the larvae of any group
described in the preceding pages. It has no doubt certain
resemblances to the trochosphere larva of Chaetopoda, Mollusca,
etc., but the differences between the two types are more striking
than the resemblances. It firstly differs from the trochosphere
larva in the character of the ciliation. Both larvae start from the
uniformly ciliated condition, but while the prae-oral ring is almost
invariable, and a peri-anal ring very common in the trochosphere;
in the Echinoderm larva such rings are rarely found ; and even
when present, i.e. the prae-oral ring of Bipinnaria and the terminal
though hardly peri-anal patch of Antedon, do not resemble
closely the more or less similar structures of the trochosphere.
The two ciliated ridges (fig. 264 A) common to all the Echinoderm larvae, and subsequently continued into a longitudinal ring,
have not yet been found in any trochosphere. The transverse
ciliated rings of the Holothurian and Crinoid larvae are of no
importance in the comparison between the trochosphere larvae
and the larvae of Echinodermata, since such rings are frequently
secondarily developed. Cf. Pneumodermon and Dentalium amongst Mollusca.
In the character of the prae-oral lobe the two types again
differ. Though the prae-oral lobe is often found in Echinoderm
larvae it is never the seat of an important (supra-oesophageal)
ganglion and organs of special sense, as it invariably is in the
trochosphere.
Nothing like the vaso-peritoneal vesicles of the Echinoderm
larvae has been found in the trochosphere ; nor have the characteristic trochosphere excretory organs been found in the Echinoderm larvae.
The larva which most nearly approaches those of the Echinodermata is the larva of Balanoglossus described in the next
chapter.
ECHINODERMATA. 577
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B. II. 37
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