ZOOLOGY

SUPER-CLASS AGNATHA
Classes Euphanerida (Silurian)
Heterostraci (Fteraspida) (Ordovician-Devonian)
Anaspida (Silurian-Devonian)
Osteostraci (Cephalaspida) (Silurian-Devonian)
Petromyzontia (?-Recent)
Myxinoidea (?-Recent)
It is almost certain that we know only a small proportion of the Agnatha
that existed during the Palreozoic. Further, next to nothing is known of their
ancestry. The earliest known vertebrate relics come from the Ordovician,
and the nature of these makes it possible that the first vertebrate animals
appeared in the Cambrian (Table I, p. 3). Opinion is sharply divided as to
whether the first true vertebrates were fresh-water or marine in habitat. It
is possible that the relatively strongly swimming vertebrates may have arisen
in fresh water, where such powers would enable them to withstand river
currents sweeping seawards. The vertebrate kidney, so different from the
excretory apparatus of Amphioxus (p. 47), possibly arose during the slow
colonisation of coastal and inland waters (p. 156). The earliest fresh-water
chordates no doubt took in surrounding fluid by osmosis through the permeable
regions of the body. The water had to be discharged. At the same time it
was necessary for vital solutes to be retained, and waste metabolites to be
discarded, if the integrity of the internal environment were to be maintained.
Pioneer movement into brackish and fresh water would impose the necessity
for considerable elaboration of the simple mesodermal funnels of excretion
which carried fluid from crelom to the exterior. In any case there arose a
device which made it possible for the power of the heart-pump to be used to
eliminate unwanted fluid, and contained substances, through a filter (Fig. roo).
This seems to have been at first a relatively simple knot of capillaries near each
tubule mouth. Upon this basic regulatory mechanism there have been
imposed the many modifications made necessary by the re-colonisation of
sea-water, and the exploitation of land, including deserts and other environ-
mental niches. A good case can be made out that all vertebrates, what-
ever their habitat to-day, are of fresh-water ancestry (see Romer, Homer
Smith).
But even stronger evidence, perhaps, can be adduced for an opposite view
(see Krogh, Watson, Robertson). While it is undeniable that most fossils of
very early vertebrates so far discovered have come from fresh-water sediments
(e.g. see p. 174), it is equally true that the earlt"est known remains (plates and
scales of Middle Ordovician ostracoderms) were unearthed from sandstone
that, judged by an associated, and indubitably marine, invertebrate fauna,
was not of fresh-water origin. It has been suggested, but not established, that

A. J. Marshall et al., Textbook of Zoology

© Macmillan Publishers Limited 1962
 PHYLUM CHORDATA r6s
these fragments are the remnants of fresh-water animals that were washed
downstream and incorporated in littoral deposits. As indicated above, there
have been suggestions that the powerful lateral body flexure of vertebrates
would help the earliest fresh-water craniates to withstand currents tending to
sweep them seawards. But it is perhaps equally easy to believe that the
development of such enhanced locomotion and stability could occur in the
relative turmoil of the ocean, and that such pre-adapted (p. 385) ancestral
forms could then swim against estuarine currents to colonise the brackish and
fresh-water streams and lakes in which so many have left their bones.
It should be remembered too that all three protochordate groups, the
Adelochordata (p. 5), Tunicata (p. 20), and Acrania (p. 47), are, and perhaps
always have been, restricted to the sea. Again, it is possible that the glomer-
ular kidney (p. ror) existed in marine protovertebrates, thus additionally
pre-adapting them for life in fresh water. Finally, the blood of some extant
agnathans (myxinoids, p. 197) has an ionic constitution strikingly similar to
that of the surrounding sea: ' this may well be a primary feature, derived
directly from marine chordate ancestors' (Robertson).
The first vertebrate group of which numerous specimens are available is
the Agnatha. The first common representatives of these were fresh-water,
bottom-living animals of the Silurian (p. 3). Groups continued to expand
during the Devonian, when they attained their maximum development.
Between that time (some 270 to 300 million years ago), and the present day,
little is known of them. Extant agnathans, unlike most of the early forms,
lack bones, true teeth, and other durable parts, and that is probably why
fossils are unknown.
It has been sometimes claimed that the not uncommonly found conodonts
are the teeth of unknown armoured agnathans. This is unlikely. Conodonts,
however, should perhaps for the time being find a place in a volume of the
present character, and this is probably the best place to discuss them. They
are small (0·2-3 mm. long), cone- or tooth-shaped, widely distributed, fossils
which show a great deal of variation but fall possibly into two groups, laminated
and fibrous. Their chemical composition is essentially bone-like and not very
different from that of the armour of Devonian fishes (p. 285). Laminated
conodonts occur in sedimentary strata from the Ordovician to the Triassic.
Fibrous conodonts occur possibly only in Ordovician deposits and may re-
present a distinct group. A highly suspect binomial classification has been
erected, and subscribed to, in many of the several hundred papers dealing with
conodonts.
Conodonts seem to be formed by the accretion of material around a basal
(but not pulp-) cavity. Radial canals, extending from cavity to surface,
occur. The fibrous specimens often have very bone-like material attached to
them. A few lamellar specimens have a similar attachment. There is some
r66 ZOOLOGY
evidence that conodonts were regenerated when broken, but they show little
evidence of wear.
These enigmatic objects have been variously held to be the branchial filters,
or skeletal supports, of primitive vertebrates or, sometimes, the teeth of
Agnatha or of primitive elasmobranch fishes. They have also been claimed
to be the teeth, or copulatory organs, of worms. Some have believed them to
be part of the armour of crustaceans or to be allied to the radular teeth of
gastropods. Rhodes, the latest reviewer, concludes that : (r) 'if of vertebrate
origin they may represent some group . . . other than fishes, now extinct,
and, apart from the conodonts, entirely unknown', or (2) they may be mastica-
tory organs of annelid affinity. Apart from their anatomy and chemistry,
the one thing that is perfectly clear is that nobody at present has any real idea
of their function or identity.
The Agnatha, apart from their antiquity per se, are of great interest in
possessing tru-e bone in the oldest groups. It was for many years confidently
held that cartilage was the forerunner of bone. To-day, however, it is now
widely agreed, with de Beer, that bone is the more primitive material and that
cartilage in the adult is probably a neotenous phenomenon. On palceonto-
logical evidence Romer has reached the same conclusion concerning its anti-
quity. It has not yet been proved, but there are excellent reasons for suppos-
ing, that cartilage may be essentially an adaptation of embryonic tissue in the
adult form. Although it could be held that Silurian and Devonian dermal
bone (as well as skeletal bone in the Cephalaspida) might be derived from car-
tilage of an even earlier date, there is no doubt that the earliest known verte-
brate fragments (from the Ordovician, about 400 million years ago) were
ossified. Stensi6 believes it possible that bone arose as early as the Lower
Ordovician, or perhaps even in the Upper Cambrian. There is no evidence of
a progressive increase in ossification in any craniate group.
The early agnathans were generally heavily armoured (Figs. 104, ros).
There is no certainty why this was so. Their buccal arrangements reveal that
they did not devour each other. The fresh-water forms were isolated from
attacks by large predacious marine invertebrates. There is evidence, however,
that large water-scorpions (Eurypterida, Vol. I) with formidable claws and
mouth-parts existed in the same aquatic environment. These animals were
allied to to-day's arachnids, and reached a length of about 9 feet-perhaps
the biggest size attained by arthropods. It is an interesting fact that when
faster fishes came to replace the early slow-moving armoured agnathan types
in the succeeding Devonian period, the eurypterids also faded and disappeared.
Homer Smith, on the other hand, believes that the armour of Silurian and
Devonian forms was a defence against ' the osmotic invasion of fresh-water
rather than the claws and tail-spines of eurypterids '. There is little doubt that
dermal ossification would in fact provide such defence; but it would seem that,
 PHYLUM CHORDATA

if protection against osmotic inflow was the primary factor involved, this may
well have been obtained by more economical means. Further, many con-
temporary animals were not protected in this way (e.g. see Fig. 103).

CLASS EUPHANERIDA
Perhaps the most primitive vertebrate yet described was the totally un-
armoured Jamoytius kerwoodi, described by White from the Scottish Silurian
(Fig. 103). First thought to be allied to Lasanius (p. 171), jamoytius was only
about 7 inches long. It was blunt headed, with a rounded, somewhat elon-
gated, body. With the ' possible exception of the supports of the median

Myoc. D.F.

FIG. IOJ.- Ciass Euphanerida, Family Jamoytiidm. Jamoytius ke-rwoodi. Specimen (lacking
tail) shown in three.quarter view, at about !th of natural size. A. F. anal fin supports; D . F .
dorsal fin supports; D. S. displaced skin; E. eye; Int. contents of intestine; L. F. F. lateral fin.
fold; Myoc. septal spaces; Myom. myomere; No. notochord. The above is one of the illustra-
tions used in White's description of the new order, family and species.

fins', there is no indication of internal calcification. It had relatively enormous

and widely separated eye-regions, a brief anal fin, and a dorsal fin continuous
along the posterior two-thirds of the body. The intestine was short and
straight. There was a persistent notochord and myotomes that were simple
in arrangement. No evidence of gill pouches opening to the exterior has been
found.
White believes that the nakedness of J amoytius is a primary character and
that earlier members of this apparently unspecialised group may have con-
stituted the stock from which gnathostomes as well as agnathans were derived.
There is a further suggestion that the Acrania (e.g. Amphioxus) (p. 68) arose
as degenerate forms from animals such as the present one. Gregory has
suggested that ' the boneless J amoytius may well have been on the way towards
Amphioxus, while the bony Cephalaspis (p. 172) may have been on or near the
r68 ZOOLOGY
line leading to the lampreys'. This view, however, implies that jamoytius
was secondarily naked.
The unusual name J amoytius is derived from that of the late J. A. Moy-
Thomas, a distinguished worker in the present field who tragically died at the
height of his powers.

CLASS HETEROSTRACI (PTERASPIDA)

A great deal is now known of the three armoured agnathan groups of the
Pal;eozoic. These, the Heterostraci, Anaspida, and Osteostraci, and probably

FIG. 104.-Class Hetero-

straci, Family Pteraspidre.
Pteraspis. Restoration of
P . rostrata from dorsal (left-
hand figure) lateral and
ventral views. a. l. p.
anterior lateral plate ; b. p.
branchial plate; c. p. corn-
ual plate; d. d. dorsal disc;
d. sp. dorsal spine; o. p. oral
plate; or. p. orbital plate;
pi. pineal plate; p. l . p.
posterior lateral plate; r .
rostrum; v. d. ventral disc.
(After E . I. White.)

the naked lampreys and hags as well, are clearly related. None, however,
appears to have been directly derived from another. Each stock probably
branched successively from the ancestral stem in the order used below.
The Heterostraci, although probably more advanced than the Euphanerida
(see above), are known to have existed in the Middle Ordovician, from which
plates of Astraspis have been described. Such animals (e.g. Pteraspis, Fig.
104) were apparently seldom more than a foot long, although some grew to a
 PHYLUM CHORD/\.TA r6g
length of 5 feet. They became more plentiful in the Upper Silurian and Lower
Devonian and persisted into the Upper Devonian. They possessed a broad,
depressed head protected by a carapace of exoskeletal plates. The eyes were
widely separated, and lateral in position. There was a sub-ventral mouth at
the extreme anterior end of the body which was equipped with probably mobile
plates. White has suggested that this buccal apparatus was protrusible and
able to scoop up bottom-mud and the contained organic material that served
as food.
There were no fins on the body. The tail behind the carapace was laterally
compressed, downwardly turned (hypocercal), and protectively covered with over-
lapping rhomboid scales of the same composition as the plates of the carapace.
The Heterostraci differed widely from the Osteostraci (see below) in several
essential points. There are no traces of a median nostril and there are strong
indications that the olfactory organ was a paired structure opening into the
roof of the mouth. The plates and scales lacked bone-cells. The plates
were formed of three layers, of which the outer was composed of a substance
allied to dentine. The middle layer was formed of vascular 'bone'; the
lower was laminated. The gill-pouches, apparently of the adult cyclo-
stome pattern, united as a single exhalent pore placed rather far back on the
sides of the carapace. There was no sign of the complicated and prominently
innervated sensory organ formerly thought to be one of electrical discharge
(Fig. ro8, p. 173).
The four groups which form the Heterostraci (Paleaspidce, Cyathaspidce,
Pteraspidce (Fig. 104), and Drepanaspidce (Fig. 105)) were distinguished from
one another by the number of plates in the carapace. The Paleaspidce had
an undivided upper shield ; the Cyathaspidce had it divided into four plates ;
the Pteraspidce nine and the Drepanaspidce twelve major and many smaller
plates.
A description of Pteraspis rostrata, a species of the best-known genus, will
serve for the remainder. The carapace was somewhat elongated and rect-
angular in section. The dorsal surface was composed of nine plates, of which
three, the rostral, pineal, and large dorsal, were unpaired. The remaining six
plates formed three pairs. These are the orbital (in which lay the eye-sockets),
the branchial (with the single gill opening at the posterior border), and just
behind these, the cornual plates. In the adult all these plates were bound
together by a fusion of their inner lamince. There was a prominent median
spine inserted at the base of the dorsal disc. On the ventral surface, a large
unpaired plate covered most of the area except for a number of small oral
plates round the lower border of the mouth. Immediately behind these were
three pairs of small plates, the post-oral and the anterior and posterior laterals.
The trunk was covered with thick rectangular scales and there was a series of
thorn-like scales along the upper and lower surface. From impressions of the
IJO ZOOLOGY
internal surface of the upper carapace it seems clear that there were seven
pairs of gill pouches. An X-like impression just behind the pineal opening has
been interpreted as evidence of two semicircular canals (seep. 145) on each side.
There was a prominent lateral line system (seep. 135).

FIG. 105.-Class Heterostraci, Family Drepanaspidm. DTepanaspis. Dorsal and ventral

views. D. gemundenensis. C. cornual plate; D, median dorsal plate; M.-M3 , ventral oral
pla tes; 0 1 -03 , ocular plates; R, rostral plate; V, ventral plate; B , branchial plate. (From
Stensi6, a fter Ki;er.)

CLASS ANASPIDA
Anaspids share some characters with the pteraspids but show a greater
resemblance to the cephalaspids (Osteostraci) although, it should be emphasised
again, they sprang from a common ancestry and were derived from neither.
They were fish-shaped animals, a few inches long, with a downwardly turned
tail (hypocercal) (Fig. 106). The internal, presumably cartilaginous, skeleton
is unknown, but a highly developed dermal skeleton was present. The scales
on the head were small and showed a complicated arrangement which differed
in pattern in different genera. The scales on the trunk were arranged in lateral
and ventral series, which, with a set of tall lateral scales, showed great resem-
blance to the condition found in cephalaspids, as does also the mode of their
articulation. There are no paired appendages, unless a pair of pectoral spines
represented them. There was-an anal spine, and a series of ridge spines along
the dorsal surface. The eyes were lateral, and the pineal and nasa-hypophysial
openings were placed on the top of the head in relatively the same position as
 PHYLUM CHORDATA IJI

in the cephalaspids. A series of gill openings, varying in number in different

genera, were placed in front of the pectoral spine. The mouth had a shape
that resembled that of jawed animals, but there is no evidence that jaws

FIG. 106.-0rder Anaspida, Family Birkeniidre: Rllyncholepis parvulus. (After Ki.er.

existed. Lasanius, from the Upper Silurian, was almost unarmoured except
for dorsal and pectoral spines. There were several groups. Birkenia, Rhyncho-
lepis (Fig. 106), Saarolepis ( = Anaspis) are typical genera.

CLASS OSTEOSTRACI (CEPHALASPIDA)

The cephalaspids (Fig. 107) had developed prominent ' pectoral fins ' and
a heterocercal tail. Much splendidly preserved material has been discovered,
allowing a relatively clear picture of their essential features. They were
bottom-living animals with a single nasa-hypophysial opening placed far back

FIG. I O].- Order Osteostraci, Family Cephalaspidre. Hernicyclaspis. H . murchisoni . D 2

dorsal fin; d. cr. dorsal crest of trunk; sd. cr. do rsal scutes; Ppl. pineal plates ; sclr. sclerotic ring ;
lsf. lateral 'sensory field ' ; P ee. pectoral fin; vhp. caudal fin. (After Stensii:i.)

on the head. The head region was expanded and flattened. It was protected
by a hard, bony carapace or head-shield which was often ornamented with
tubercles, and usually produced on each side into a backwardly directed ' horn'
which left a bay, or pectoral sinus, between itself and the trunk. Beneath the
true dermal bone in at least some cephalaspids there occurred a fragile and
continuously ossified endocranium, equally curious both in shape and constitu-
tion. This was usually cartilaginous in later forms. The body was protected
by rows of scales in which true bone-cells were present. A dorsal fin occurred.
172 ZOOLOGY
Two Orders, Cephalaspidiformes (Fig. 107) and Tremataspidiformes, are
generally recognised.
In Cephalaspis, the most completely known genus, there was, as mentioned
above, a pair of lobed appendages arising from the pectoral sinus on each side.
These were neomorphs, and therefore not homologous with the pectoral fins of
vertebrates. Of the internal anatomy, too, a good deal is known. The brain,
with its ten pairs of cranial nerves, was closely comparable with that of the
living cyclostomes, but in addition there was a well-developed system arising
in the acoustico-lateralis region. From here five pairs of stout vessels ran to
depressions on the upper surface of the head-shield. By analogy with such
modem forms as torpedoes (p. 271), Stensio suggested that these nerves and
depressions formed a kind of electric organ (Fig. 108).
There are good reasons for believing this theory to be invalid. Westall
has reinterpreted the so-called ' electric fields ' as a sensory organ. He be-
lieves them probably to have been ' receptors of vibrationary stimuli ', con-
sisting of invaginated pockets, probably derived from placodes, and richly
supplied by neuromast organs innervated by the acoustico-lateralis system.
In short, they possibly compensated for the relatively reduced lateral line
system (p. 135), which was in them much less extensive than that of the
Heterostraci existing in the same habitat. Watson, too, is of the opinion
that they may constitute a sensory apparatus that might allow 'pressure
waves incident on the head to be conveyed along the fluid-filled canals to the
ear ' .
Cephalaspis was a bottom dweller. The eyes were placed close together on
the top of the cephalic shield, with the pineal aperture between them. It is
possible that the pineal was a photoreceptor which helped the animal gauge the
depth of water and, by means of pituitary hormones, protectively control its
colour in a manner comparable with that of the ammocrete larva of Petromyzon
(p. 196). The nasa-hypophysial opening was a little anterior to the pineal
aperture. (See p. 345.)
The ventral mouth was followed on each side by ten pairs of gill-pouches,
of which the first two (anterior and spiracular gill-cleft) were innervated by the
ophthalmicus profundus and trigeminal nerves respectively. Each such nerve,
then, was in Cephalaspis a typical branchiomeric nerve, i.e. one associated with
a functional gill-slit. We will see that in jawed animals (pp. 205, 210) these
nerves supply apparently quite unrelated structures (i.e. the snout and jaws).
This remarkable change in situation and function has been dictated by the
enlargement of the mouth and buccal cavity in relation to the shift to macro-
phagous habits. The comers of the mouth migrated backwards, obliterating
the first, and absorbing the second, pairs of gills (Fig. 137). For many reasons
it now appears certain that in this way the mouth came into proximity with
the branchial arch (posterior to the second pair of previously functional gills),
 PHYLUM CHORDATA 173

F rG. IoS.-Osteostraci: Cranial anatomy of Kireraspis.

Up per left: top v iew of hea d and anterior pa rt of trunk region: deo. d orsal sensory (supposed
' electric') organ; leo. lateral organ ; n . opening of single dorsal nost ril and h ypoph ysis ; o. orbit ;
p. p ineal opening.
Lower left: diagra mma tic dissect ion of top of h ead, t o show b ra in, nerves, a nd sense organs.
N erves to sensory organ shown on the left ; other major nerves (numbered) on right, a rranged
in a series corresponding to the gilt slits. c. Cerebellum; d. nerve to dorsal sensory field; e.
orbit ; f. forebra in ; m . m idbrain ; mo. medulla oblongat a; n . opening for nostril a nd h ypophysis ;
o. ear region (two canals) . I n this a nd t he next figure Roman numera ls indicat e cra nia l nerves.
Upper right: restoration of ventral surface of head, with the plates covering the t h roat re.
m oved, showing the ten gill sacs a nd t he small, a nteriorly placed m out h. dpr. mou t h; ibs.
par t it ions between gill sacs ; kebr. d uct s from gill pouches t o surface; res. cesophagus ; psg. a
p respiracula r gill pouch lost in higher vertebrates; rpm. rostral region in front of gills ; sg. gill
pouch corresponding to t he spira cle of higher fishes ; vm. muscles of gill pouches.
L ower right: restoration of ventral surface of head, covered by sma ll plates: ebrc. opening of
the gills; 111. mouth. (From R omer , after St ens io .)
174 ZOOLOGY

and the skeleton of this arch and its musculature came to be modified into a
powerful masticating apparatus, i.e. the modern jaw.
In Cephalaspis the whole of the ventral mouth and gill region was supported by
a system of small, tessellated, bony plates, constituting the oro-branchial area.
The considerably expanded gills had nothing in common with those of fishes.
It has been suggested that their structure resembled that found in lampreys of
to-day (p. 175) or, alternatively, perhaps that of the ammocrete larva (p. 196).
It is generally held that ostracoderms strained organic particles from debris
sucked in from the floor of fresh-water lakes and streams : the small ventral
mouth, dorsal eyes, and generally flattened form point to this conclusion.
The Osteostraci ranged from the Upper Silurian to
the Upper Devonian, and a considerable number of
genera and species is known. They showed great
variation in the proportions of the cephalic shield and
in its ornamentation. Representative genera are
Cephalaspis, H emicyclaspis, A teleaspis, Benneviaspis,
Kia:raspis (Fig. ro8), Didymaspis, Thyestes, and others.
The tremataspids differ from other cephalaspids

I
chiefly in the absence of the pectoral sinus, and, in con-
sequence, of the paired appendages. The cephalic shield
was well developed, and extended backwards on to the
trunk. The lateral sensory fields were divided into
anterior and posterior portions. The members of some
genera of the Cephalaspidiformes were transitional in
shape; the two groups were clearly allied.

THE ' C<ELOLEPIDA '

FIG. 109.-Lanarkia
spinosa, possibly related Several genera of small fish-like animals which
to the Heterostraci and usually appear as flattened impressions with a broad
formerly often placed with
head-region, a somewhat fusiform body, and a hetero-
'Order' Ccelolepida. R e·
stored outline in the posi-
cercal or hypocercal (e.g. Ccelolepis) tail have been de-
tion in which it occurs as
a fossil, the head beingscribed. They occurred from the Upper Ordovician to
flattened and the t ail
the Devonian. Their armour appeared to be restricted
twisted round so as to a p-
pear in profile. On eachto minute denticles (Fig. rog) that produced, in some
side a much-enlarged der-
mal denticle is shown. forms at least, a shagreen surface perhaps not unlike
(From the Cambridge Natu-
that of selachians (p. 267). It has often been considered
ral History, afterTraquair.)
that the ' Crelolepida ' forms a single well-defined
assembly. Westoll, however, has made it very probable that two or more
structurally distinct groups exist. One such is represented by Thelodus and
Lanarkia (Fig. rog) and another by Cephalopterus.
The first group were related to the Heterostraci and the second to the
 PHYLUM CHORDATA IJ5
Osteostraci. Other forms (e.g. Pldebolepis, Co::lolepis) showed certain resem-
blances to the Anaspida. It is still uncertain whether the ccelolepids represent,
or include (r) forms that belonged to each of the above groups and never
became ossified, (2) secondarily unossified types that descended from armoured
ancestors, and (3) the ' larval ' stages of animals that might have developed
dermal ossifications at a later stage of their development.

EXTANT AGNATHA (CYCLOSTOMATA)

Lampreys and hags, the only surviving agnathans, are often grouped to-
gether in the Class or Order Cyclostomata, distinguished from all other living
craniates by the possession of pouched gills and a single nostril, and the
primitive absence of jaws or paired fins. However, such features, which are
shared with one or other of the fossil groups, do not provide evidence that hags
and lampreys are more closely related to each other than they are to the
earliest agnathans. It seems probable that the lampreys are related to a
cephalaspid-anaspid stock, but the affinities of the hags are very doubtful. In
view of this it is advisable to regard the Cyclostomata as a 'group of con-
venience' and to treat the lampreys and hags as separate Classes of the Agnatha.

CLASS PETROMYZONTIA
This group comprises the lampreys, Agnatha with a round sucking mouth,
a pharynx which ends blindly, and (like the cephalaspids) a nasa-hypophysial
sac opening by a single nostril on the dorsal surface of the head. There are
about half a dozen genera, differing from one another only in minor anatomical
fea tures. Petromyzon (Fig. rro) occurs on both coasts of the North Atlantic,
and Entosphenus on both coasts of the North Pacific. Geotria occurs along the
southern coast of Australia, and in New Zealand, Chile, and Argentina.

d. f.,
g.
1.1.

·-e~~_.{l~~~
.- l ·~·-~.,- _:.:....:,............:.....,.~~~~
b. f.
FIG. 1 10 .- Class Petromyzontia, Family Petromyzonidre. Petromyzon. Adult Lamprey
(P. marin us). b. f. buccal funnel; c. f. cauda l fin; cl. cloaca; d. f. 1 , 2 1st and znd dorsal fins ; g. ex-
terna l gill aperture; /.I. la tera l line organs; n. nasal aperture. (Drawn by R. Strahan.)

M ordacia is restricted to the Southern Pacific. The remaining genera, all

Holarctic, appear to be degenerate descendants of the Northern genera already
mentioned.
 ZOOLOGY

EXAMPLE OF THE CLASS:-

THE LAMPREY (PETROMYZON)
Three species of lamprey are common in European river systems. The
Sea-lamprey (P. marinus) attains a length of a metre, the Lampem, or common
fresh-water lamprey (Lampetra fluviatilis), reaches a length of about go em.,
and the Sandpride, or lesser Fresh-water Lamprey (L. planeri), does not exceed
45 em. in length.
Two quite different phases occur in the life-cycle of Petromyzon. The larval
phase or ammoccete feeds on microscopic particles by means of an endostyle
not unlike that of Amphioxus. The adult has a suctorial mouth, and a rasping
apparatus that enables it to cling to fishes and feed on their tissues. Com-
paratively little is known about the habits and life-cycle of even the common
species. The adult lives in the sea for most of its life. It attaches itself to a
fish, rasps away some of the host tissue, and then feeds on its blood. It is often
found in this situation when the host is caught. At the onset of its single
breeding season it migrates coastwards and ascends a river into fresh water.
In Europe this occurs in the autumn, when great numbers of L. fluviatilis
invade favoured streams. They are not infrequently used as food, though
more particularly in past eras, as the sad fate of Henry I of England will recall.
In some instances they ascend rapids in their urge to reach the breeding-
ground, and cling to the rocks with their suckers between leaps forward and
upward. During this migratory phase the gut atrophies and they cannot feed.
They are sustained by fat reserves accumulated sub-cutaneously and in the
body-muscles. Towards the end of winter the gonads become mature. Secon-
dary sexual characters develop-a prominent anal fin in the female, and a
penial-tube and a thickening at the base of the dorsal fin in the male.
In the spring, lampreys aggregate in pairs in shallow spawning-grounds
and build their nests. They move stones from the stream-bed with their
suckers and construct a sandy floor with a crescentic ridge on the downstream
border. Although copulation occurs (the male clinging to the female by means
of its buccal funnel), fertilisation is external. Both eggs and spermatozoa
(which are ejected through the penial tube and are very short-lived) are ex-
truded into the surrounding water, whence they are carried by the current into
the interstices of the stony rim of the nest. After each of the numerous short
matings, sand is stirred up from the floor of the nest by vigorous tail move-
ments, and this serves to embed the eggs more firmly among the stones. It is
said that the seasonally developed anal fin of the female assists in this operation.
When spawning is completed, the adults drop away from the nest and die within
several days.
It is of great interest that lampreys that inhabit lakes in North America
(where their ancestors were isolated) have successfully adapted themselves to
 PHYLUM CHORDATA 1 77
live their whole lives in fresh water, but nevertheless move a few miles up
lake-side streams at the onset of the breeding season.
External Characters.-The head and trunk (Figs. no, rn) are nearly
cylindrical, with the tail-region compressed or flattened from side to side. At
the anterior end, and directed downwards, is a large, basin-like depression, the
buccal funnel (b. f.), surrounded by a marginal membrane (m. m.) outside of
which arise the overlapping oral fimbrice (o. f.) and the longer, sensory cirri (c.).
The inside of the buccal funnel is beset with radiating rows of yellow horny
teeth (i. o., l. t., s. o.). The teeth rest on cartilaginous pads and are principally
mesodermal in origin, and therefore not homologous with true vertebrate
teeth, which first appear in fishes.
From the bottom of the buccal o. f. s.o.
funnel there projects a prominence,
the so-called ' tongue ', bearing even
larger horny teeth (t. t.), and having t.
immediately above it the narrow
mouth (m.). On the dorsal surface of
the head is a single median nostril
(n.), and immediately behind it a
transparent area of skin indicates the
position of the pineal organ. The
paired eyes have no eyelids, but are
covered by a transparent area of skin.
The gill-slits (g.) are seven pairs of
small apertures on the sides of the FIG. I I I . -Petromyz on: Buccal funnel.
cirrus; i. o. infra-oral tooth p lates; l . t.
head, the first a little behind the eyes. c. lateral bicuspid tooth; m . mouth; m . rn.
On the ventral surface, marking the marginal membrane ; o. f. oral fimbri;e ; s. o.
supra-oral tooth; t. t. tongue tooth. (Redrawn
junction between trunk and tail, is the after Bigelow and Schroeder.)
very small anus lying in a slight de-
pression. Immediately behind it is a small papilla pierced at its extremity by
the urinogenital aperture (cl.). There is no trace of paired appendages. Two
dorsal fins of approximately equal dimensions (separated by a notch) and a
caudal fin are present, the second dorsal being continuous with the caudal. All
these fins are median and unpaired. As previously mentioned, the sexes during
the period of gonad maturation can be distinguished by the number and structure
of the fins and, in addition, by the appearance of a penial tube in the male.
This is formed by the junction of the cloacal margins. At this period, too, the
cloacal margins of the female become distended and sometimes reddened.
The skin is soft and slimy and, as in Myxine, lacking in exoskeletal structures
(Fig. II2). Among extant forms scales, like true teeth, do not appear until
we ascend further the chordate stem. Keratin (p. 85) does not occur in the
epidermis although it possibly forms the caps on the 'teeth' (p. 184). As the
VOL. II. M
 ZOOLOGY
outermost layers of the epidermis become worn away by abrasion, new cells are
added by division of those at the base of the layer. Thus, these forms differ
from Amphioxus and resemble the higher vertebrates in their possession of a
several-layered, more durable, epidermis. Underlying the epidermis is a layer
of collagen and elastic fibres which forms the cutis. This layer contains also
pigment-cells or chromatophores. The pigment in these star-shaped cells changes

FIG. II 2.-Myxine: Integument. Vertical section showing structure and cell-types in M .

glutinosa. a. t. adipose tissue; b. v. blood vessels; c. t. f. connective t issue fibres; d. dermis; e.
epidermis ; m. c. mucous cell ; p. c. pigment cell; t. c. thread cell. (Thread cells are peculiar to t he
Myxinid<e. Their mucous cells are atypical in that t he nucleus lies in the centre of the cell
(cf. Fig. 113, r) . (Dra wn by G. H. 0. Burgess. )

its position under various conditions of illumination. Thus the ' colour' of the
lamprey changes from pallid to dusky and vice versa. This faculty is particularly
well shown in the larva (p. 196).
The epidermis is equipped also with numerous unicellular glands which
secrete a slime which makes Petromyzon difficult, or impossible to hold with the
bare hand. In addition, two other types of secretory cell are formed (Fig. II3).
One, the granular cell, is of yet unknown function. The second, the club cell
(Kolbenzelle) , is an elongated cell with hyaline cytoplasm. These are found also
in certain t eleosts, notably in Anguillid<e (eels). There have been suggestions
 PHYLUM CHORDATA 179
that club cells are neural (but fibres have not been demonstrated to end in
them), or scab-forming (but at least in Anguilla and Tinea such is not the
case). On the other hand, there is evidence that club cells contribute to the

t.
n.

.
_

n.

Frc. 1 13.-Integument: Epidermal cells of agnathans and fishes. 1. Mucous cell of Salmo lrttlta;
2. club cell of Conger conger; 3- thread cell of Myxine glutinosa; 4- granular cell of Petro-
myzon marinus.
c. cytoplasm; g. granules; n. nucleus; m. mucus cell; I. ' thread' in thread cell. (Drawn by
G. H . 0 . Burgess.)

sliminess of eels in which mucous cells are present only in relatively small
numbers.
The segmental sense-organs take the form of a lateral line (p. 136). These
organs are restricted to fishes and fish-like vertebrates (including tadpoles of
Amphibia) . In lampreys, lateral line organs are not enclosed in a canal, but
are exposed to the exterior.
r8o ZOOLOGY
Endoskeleton.-The axial skeleton of the trunk is simple. There is a per-
sistent notochord (Fig. II4, nc.) with a tough sheath composed of an inner
fibrous and an outer elastic layer. This longitudinal central rod probably pre-
vents the body from shortening when the muscle segments contract and drive
the animal forward. Attached to the sides of the notochord are small, vertical,
rods of segmentally arranged
cartilage, bounding the spinal
canal on each side, and which
correspond to the rudimentary
neural and interneural arches.
In the caudal region these fuse
into a single plate perforated
by foramina for the spinal
nerves and send off processes
to the base of the fin. For
the rest of its length the spinal
canal is enclosed only by
nc.
tough, pigmented connective-
c.t. tissue. Slender rods of carti-
lage support the median fins.
cd. d. ao. The cranium also is primi-
tive in structure. Its floor is
formed by a basal plate (Fig.
ns, b. pl.) , made by the union
S·C. S.
mes.

of the parachordals and trabe-

culre, surrounding posteriorly
the anterior end of the noto-
chord. Immediately anterior
to the notochord is a large
l.v. aperture, the basi-cranial fon-
F IG. 114.- Petromyzon: Trunk region. Transverse
section. cd. posterior cardina l vein; c. t. connective
tanelle (b. cr. f.), due to the
tissue ; c. r. cartilaginous rods ('fin rays'); d. ao. dorsal non-union of the posterior ends
aorta ; f. c. fat column; int. intestine ; i. v. intra-intestinal
vein; mes. mesonephros ; mes. d. mesonephric duct ; my. of the trabecul<e. Through
myotome; n . ca. spina l canal; nc. notochord ; s-c. s. sub- this passes the nasa-hypophy-
cardinal venous sinus; sp. c. spinal cord; sp. v. spira l
valve ; ts. test is. (Drawn by R. Strahan .) sial sac (Fig. n 8, na. hy p.;
Fig. 121 , pty. p.) extending
from the olfactory sac t o the ventral surface of the notochord. Lateral
walls extend upwards from each side of the basal plate, but the roof of the
cranium is formed by membrane except at one point. H ere a narrow transverse
bar (cr. r.) extends across between the side-walls and furnishes a rudimentary
roof. United with the posterior end of t he basal plate and forming the end
of the neurocranium are the otic capsules (Fig. n s. au. c). The side-walls are
 PHYLUM CHORDATA r8r

pierced by apertures for the cranial nerves. So far the skull is typical, though
extremely simple; its remaining parts, however, are in many cases very
difficult of interpretation.

cn.c
FIG. IIj.- Petromyzon : Skull. Dorsal (A). ventral (B). and sectional (C) views of P. marinus.
The cartilaginous parts are d otted. a. d. c. anterior dorsal cartilage; an. c. annular cartilage;
au. c. auditory capsule; b. cr. f. basi-cranial fontanelle; b. pl. basal plate; en. c. cornual cartilage ;
cr. r. cranial roof; n. a. neural arch; na. ap. nasal aperture; nclt. notochord; Nv. I, olfactory
nerve; Nu. z, 5, and 8, foramina for the optic, trigeminal, and auditory nerves; Nv s'. trigeminal
nerve; olf. c. olfactory capsule; p. d. c. posterior dorsal cartilage; p. lat. c. posterior lateral
cartilage; sb. oc. a. sub-ocular arch; st. p. styloid process. (After \'V. K. Parker.)

The olfactory capsule (olf. c.) is an unpaired concavo-convex plate which

supports the posterior wall of the olfactory sac and is pierced by paired apertures
for the olfactory nerves. It is unique in being united to the cranium by fibrous
tissue only. Extending outwards and downwards from each side of the basal
182 ZOOLOGY
plate is an inverted arch of cartilage, called the subocular arch (Figs. n6 and
IIJ, s. o. a.) because it supports the eye. From its posterior end a slender
styloid process (st. p.) passes directly downwards and is connected at its lower end

ling . c. corn. c.

FrG. n6.-Peh·omyzon: Skull and branchial basket in adult. P . maJ•inus. a. d . p. anterior

dorsal plate; a.l. c. anterior lateral cartilage; an. c. annular cartilage; br. 1 rst branchial bar ; corn. c.
cornual cartilage; cr. w. cranial wall; ep. b. epitrematic bar; ex. hy. extrahyal bar; g . ap. gill
aperture; hy. b. hypotrematic bar; ling. c. lingual cartilage; m. v. b. m edian ventral bar; m. v. c.
median ventral cartilage; n. a. neural arch; nc. notochord; olf. olfactory capsule; at. otic capsule;
p. d. p . posterior dorsal plate; per. c. pericardia! cartilage; p. l. c. posterior lateral cartilage;
s. c. b. subchordal bar; s. o. a. subocular arch ; st. c. styloid cartilage; sty. c. stylet cartilage ; trab.
trabecula; v. p. vertical pillar of subocular arch. (Modified after Parker.)

cr. w.
o/1. st. c.

corn. c.
ex. hy.
FrG. II].-Lampetra: Skull during metamorphosis. (Tongue skeleton omitted.) L. plane~·i.
a. d. p. anterior dorsal plate; an. c. annular cartilage; br. 1 rst branchial bar; corn. c. cornual cartilage;
cr. w. cranial wall; ex. hy. extrahyal bar; nc. notochord; olf. olfactory capsule; ot. otic capsule;
pch. parachordal; p. d. p . posterior dorsal plate; p . l. c. posterior lateral cartilage; s. o. a. sub-
ocular arch; st. c. styloid cartilage; sty. c. stylet cartilage ; trab. trabecula; trab. c. trabecular com.
missure; v. p. vertical pillar of subocular arch. (Drawn by R . Strahan, reconstructed after Damas.)

with a small cornual cartilage (corn. c.). These two cartilages arise in a similar
manner to the skeleton of the gills and probably represent the reduced arch
of the mandibular somite. The styloid process has been identified with the
extra-mandibular cartilage of gnathostomes (p. 203), and the cornual cartilage
 PHYLUM CHORDATA

may possibly represent the mandibular cartilage. In close relation with the
angle of the subocular arch is an upwardly directed plate, the posterior lateral
cartilage (p. l. c.).
Connected with the anterior end of the basal plate is the large bilobed
posterior dorsal plate (p. d. p.). Below and projecting in front of it is the
anterior dorsal plate (a. d. p.). Also supporting the upper lip are the paired
anterior lateral cartilages (a. l. c.) which arise in the premandibular somite and
possibly represent the internal visceral arch of that segment. The great ring-
shaped annular cartilage (an. c.) supports the edge of the buccal funnel and
carries at its sides two backwardly directed rods, the stylet cartilages (sty. c.).

ap.

vent. atr. per. c.

FIG. 1 r8.-Petromyzon: Head region of adult. (Diagrammatic half-section.) a. d. p. anterior
dorsal plate; an. annular cartilage; ap. apical cartilage; atr. atrium; b. f . .buccal funnel; b1·.
brain; ccel. ccelom; gon. gonad; g. op. opening to gill; hs . hydrosinus; int. intestine;
l. liver; ling. c. lingual cartilage; m. v. b. median ventral bar of branchial skeleton;
m. v. c. median ventral cartilage (copula); na. nostril; na. hyp. nasa-hypophysial duct; nc. noto-
chord; ces. cesophagus; o. f. oral fimbri;e; olf. olfactory capsule; p. d . p. posterior dorsal plate;
per. c. pericardia! cartilage; ph. pharynx; s.c. spinal cord; t. t. tongue teeth; vel. velum; vel. t.
velar tentacles; vent. ventricle. (Drawn by R. Strahan.)

The' tongue' is supported by a long unpaired lingual cartilage (Fig. n8, ling.
c.). This is tipped in front by a small median, and a pair of still smaller lateral,
cartilages. Below it occurs a slender T-shaped median ventral cartilage (m. v. c.).
The visceral skeleton differs remarkably from the ordinary craniate type and
has many elements which cannot be homologised with it. It is most probably
represented in the extra-branchial cartilages of the lower gnathostomes, the
inner series, which forms the bulk of the branchial skeleton, being absent from
the post-mandibular segments in the lamprey.
The visceral skeleton consists of a branchial basket formed, on each side,
of nine irregularly curved vertical bars of cartilage (Fig. n6, br. 1 ). The
first is placed almost immediately posterior to the styloid cartilage and is
homologous with the extra-hyal cartilage of elasmobranchs (p. 219). The
second lies immediately in front of the first gill-cleft, and the remaining seven
just behind the seven gill-clefts. These bars are united together by four
 ZOOLOGY
longitudinal rods. One, the subchordal bar (s. c. b.), lies alongside the noto-
chord and is connected in front with the cranium. Two others, the epitre-
matic bar (ep. b.) and hypotrematic bar (hy. b.), are placed respectively above and
below the gill-clefts. The fourth, the median ventral bar (m. v. b.), is situated
close to the middle ventral line and is partly fused with its fellow of the opposite
side. The posterior vertical bar is connected with a cup-like pericardial
cartilage (per. c.), which supports the posterior and lateral walls of the peri-
cardium. The whole branchial basket lies external to the gill-pouches and
branchial arteries, not, like typical visceral arches, in the walls of the pharynx.
The median fins are supported by delicate cartilaginous rods (Fig. II4, c. r.,
Fig. 125, cr.). These are more numerous than the myomeres, and lie parallel to
one another in the substance of the fin, extending downwards to the fibrous
neural tube. The structure of the cartilage is peculiar and varies in different
parts. It has very little matrix. Such rods are probably not homologous
with the fin-rays of true fishes (p. 307).
The muscles of the trunk and tail are arranged in myomeres which take a
zigzag course. In the branchial region they are divided into dorsal and
ventral bands, which pass respectively above and below the gill-slits; but in
the trunk there is no division into dorsal and ventral parts. A great mass
of radiating muscle is inserted into the buccal funnel, and the 'tongue' has
an extremely complex musculature which derives its nerve-supply from the
trigeminal nerve.
Alimentary Canal and Associated Structures.-The teeth are laminated horny
cones. Beneath them lie mesodermal papillre (covered with ectoderm) which
bear a superficial resemblance to the germs of true calcified teeth. When
worn out they are succeeded by others developed at their bases. Opening
into the mouth below the tongue are paired ducts which lead from the two
buccal glands. These are embedded in the basilaris muscles and appear to
secrete a substance which prevents the coagulation of the blood of the host.
The mouth leads into a buccal cavity (Fig. n8, b. f.) formed from the stomo-
dreum of the embryo, and communicating behind with two tubes placed one
above the other. The dorsal of these is the cesophagus (as.); the ventral canal
is the respiratory tube (ph.), or pharynx. Guarding the entrance to the latter
is a curtain-like fold, the velum (vel.). The <:esophagus extends above the
pericardium and enters the intestine (int.) through a valvular aperture. The
intestine is unconvoluted. Its anterior end is slightly dilated, and is the only
representative of a stomach. Its posterior end is widened to form the rectum
(Figs. no, 125), which ends at a cloaca. As Barrington has shown, true gastric
digestion involving the action of pepsin does not seem to occur until we reach
the true fishes (as far as the vertebrates of to-day are concerned). The develop-
ment of the capacious stomach is correlated with the ancient change from
micro- to macro-phagy (p. 206). Although a stomach is lacking in some of the
 PHYLUM CHORDATA r85

higher fishes (p. 336), it is probable that in the Agnatha its absence is a primitive,
rather than a secondary, feature.
The whole of the intestine of Petromyzon is formed from the mesenteron of
the embryo, and the blastopore becomes the anus, there being no proctod<eum.
The lumen of the intestine is crescentic owing to the presence of a typhlosole
in which lie the anterior mesenteric artery and hepatic portal vein. Between
these vessels and the wall of the typhlosole is a reticular tissue whose cavities
communicate with capillaries. Blood cells of all types are formed in this spleen-
like tissue and passed from there to the blood. The typhlosole takes a slightly
spiral course along the intestine and is hence known as the spiral valve (Fig. II4,
sp. v.). There is no continuous mesentery, but only a number of narrow
supporting bands.
The liver (Fig. u8, l.) is a large bilobed organ. Gall-bladder and bile-duct
occur in the larva, but are at least sometimes absent in the adult. There is
no pancreas as such. In both larval and adult lampreys, however, there occur
in the gut epithelium aggregations of seemingly secretory cells of two different
kinds. There is suggestive experimental evidence that one kind (zymogen
cells) produce a proteolytic enzyme, and that the other sort is involved in
carbohydrate metabolism. This second, probably endocrine, group has no
communication with the lumen of the intestine. It arises in the larva as' cords
of cells' of one type (Barrington), and forms what have been termed follicles of
Langerhans. These may be homologous with the islets of Langerhans (p. 153)
of higher forms. Both secretory elements-hypothetical endocrine and zymo-
gen-occur in the sub-mucosa at the anterior end of the broader mid-gut.
The follicles of Langerhans are aggregated near the entrance of the bile-duct.
It must be emphasised, however, that the hvo elements, though occurring in
the same region, are not as intimately associated as are the acinous cells and
insulin-producing elements in the compound pancreas of the Gnathostomata
(p. 203).
Respiratory Organs.-Adult lampreys differ from all other Vertebrata in
that the pharynx ends blindly. By virtue of this difference it is referred to,
perhaps unnecessarily, as the ' respiratory tube ' (Fig. u8, ph.). In the
ammoccete (p. 196) the pharynx communicates posteriorly with a short,
narrow <:esophagus but, during metamorphosis, this connection is lost and the
cesophagus becomes extended forward dorsal to the respiratory tube and opens
separately into the buccal cavity.
The respiratory organs are typical gill-pouches. They have the form of
biconvex lenses, with numerous gill-lamell<e developed on the inner surfaces,
and are separated from one another by wide interbranchial septa. The gills
are much less pouch-like in the ammoccete, where they open widely into the
pharynx (Fig. ug). In the embryo an additional gill pouch rudiment occurs
in front of the first of the adult series.
r86 ZOOLOGY
Circulatory System.-The blood-vascular system shows a radical advance-
ment from the primitive Amphioxus plan in the possession of a distinct heart.
The atrium (Fig. II8, atr.) is situated to the left of the ventricle (v.) and receives
blood from a small sinus venostts (s. v.). There is no conus arteriosus, but the
proximal end of the ventral aorta presents a slight dilatation or bulbus arteriosus.
Both afferent and efferent branchial arteries supply each the posterior hemi-
branch of one gill-pouch and the anterior hemibranch of the next. They are
thus related to the gills, not to the gill-pouches. In addition to the paired
jugulars there is a median ventral inferior jugttlar vein returning blood from
the lower parts of the head. There is no renal portal system (p. 244), the two
trab. c. nc. sp.c.
br.

u.l.

1.1. an. r. ling. r. vel. g. end.

FIG. I 19.- Geot-ria: Ammoc<ete head region. (Diagrammatic half-section of G. australis.) an. r.
rudiment of annular cartilage (mucocartilage); ap. r. rudiment of apical cartilage (mucocartilage) ;
br. brain; b. tent. buccal tentacles ; end. endostyle; g. gill; ling. r. rudiment of lingual cart ilage
(mucocartilage); l. l. lowe r lip; na. nasal aperture; nc. notochord; olf. c. r . rudiment of olfactory
capsule (cartilage); r. d. p. rostra-dorsal plate (mucocartilage); sp. c. spinal cord; lrab. c. trabe-
cular commissure (cartilage); u. l. upper lip; vel. velum. (Drawn by R. Strahan.)

branches of the caudal vein being continued directly into the cardinals.
Blood from the kidneys drains into two sub-cardinal sinuses (Fig. II4, s.-c.s.)
which connect by numerous small passages with the cardinal veins above.
The left precaval disappears in the adult, so that the jugulars and cardinals
of both sides open into the right precaval. On the sides of the segmental
arteries and veins occurs diffuse chromaffin tissue (p. 152). There is evidence
that an extract of this tissue has properties strongly resembling that of mam-
malian adrenalin. Such tissue may represent a diffuse adrenal medulla. Islets
of cells around the lateral and ventral walls of the cardinal veins have been
thought possibly to represent adrenal cortical tissue (p. 152).
The erythrocytes of the blood contain a respiratory pigment, hcemoglobin,
which is intermediate in many of its properties between the hcemoglobins of
invertebrates and those of gnathostomes.
 PHYLUM CHORDATA

As previously mentioned, the formation of blood cells, hcematopoiesis,

occurs primarily in the spiral valve. In the adult, this function is shared
with the reticular tissue of the kidney. The erythrocytes, in common with
those of true fishes, amphibians, reptiles, and birds, retain their nucleus through-
out the life of the corpuscle. The presence of hcemoglobin and a force-pump,
the heart, enormously increases the capacity of the circulatory system to trans-
port speedily the oxygen (taken in at the gills) to the tissues throughout the
body. An extensive system of lymphatic sinuses (p. n6) also occurs.
Nervous System.-This, too, shows a great advance on that of Amphioxus.
An elaborate brain is developed along with a system of cranial nerves. It is
of great interest, however, that the dorsal and ventral roots of the spinal nerves
are unjoined, and therefore retain the Amphioxus plan. Similar to those of
Amphioxus, and differing from those of other vertebrates, the nerves of the
Lamprey and other cyclostomes are unmyelinated. The spinal cord is wholly
composed of transparent grey matter. It contains no blood-vessels, but is
dorso-ventrally flattened in a manner that may possibly allow metabolic
exchanges to take place without them. As in the fishes and other vertebrates,
the nerve cell-bodies are arranged around the narrow central canal, but synaptic
contacts occur at the periphery of the cord in what would be the 'white matter'
in other vertebrates. This arrangement gives rise to a neuropil (nerve felt-
work). In this, afferent sensory fibres terminate in close proximity to the
dendrites of motor-cells whose axons become associated in the ventral nerve-
roots. It is thus probable that the motor-fibres can be stimulated directly as
a result of sensory impulses initiated by contact with environmental stimuli.
The connection between brain and motor-fibres appears to be by means of the
large and prominent Muller's fibres which originate from giant cells in the
mid- and hind-brain and probably stimulate motor-cell dendrites in the ventral
region of the cord.
The brain (Figs. 120, 121) exhibits the typical pattern that is found through-
out the vertebrate series. Some of the structures that are well-developed in
the higher forms occur in a relatively primitive condition. This of course is
essentially a reflection of the life of the animal. The optic lobes (Figs. 120, 121,
opt. l.) are very imperfectly differentiated in the tectum of the mid-brain.
In the hind-brain the small size of the cerebellum is remarkable. This organ
occurs as a mere transverse band roofing over the anterior end of the 4th
ventricle, and its lack of complexity is not surprising when we observe the
Lamprey's mode of locomotion even when it is not being transported by its
host. The rest of the hind-brain is highly developed ; among other functions
it controls the powerful sucking structures by means of the trigeminal nerve.
The central region of the tectum of the mid-brain is epithelial and is related to
a vascular thickening of the overlying pia-mater which forms a choroid plexus,
so that when the membranes are removed an aperture is left. No choroid
r88 ZOOLOGY

plexus is found in the mid-brain of higher vertebrates, where such structures

are limited to the fore- and hind-brain.
On the dorsal border of the lateral wall of the diencephalon are two ganglia
habenulce, the right (r. gn. hb.) being much larger than the left (l. gn. hb.).
These constitute an epithalamus and are connected to the pineal apparatus.

Nut
.Nv.f
o(/l - .._
Pn- - -- -+-

opt.l - ~ --. - cr. c rh

I ~ ~ - Nu .S
,•
'
- Nu.S
m.ed.ohl
~- m.ed . ohl

FIG. I2o. -Petromy~on: Brain. Dorsal (left) and ventral (right) views of P . marinus. ch. pl. I ,
anterior choroid plexus forming roof of prosencephalon and diencephalon; ch. pl. 2, aperture in
roof of mid-brain exposed by removal of middle choroid plexus ; ch. pl. 3, m etacrele exposed by
removal of posterior choroid plexus; crb. cerebellum; cr. crb. crura cerebri ; crb. h. olfactory
lobes ; dien. diencephalon; inf. infundibulum ; l. gn. hb. left habenular ganglion; med. obl. medulla
oblongata ; Nv. I, olfactory, Nv. 2 , optic, Nv. 3, oculo-motor, Nv. 5, trigeminal, and Nv . 8, auditory
n erves; olf. l. olfactory bulbs; opt. l. optic lobes; pn. pineal eye; r. gn. hb. right ganglion
habenul<e. (After Ahlborn.)

This consists of the pineal eye which lies against a transparent area of the cranial
roof and the smaller parapineal body lying below and somewhat behind it. A
stalk containing nerve-fibres leads from the pineal eye to the right habenular
ganglion, and a similar one from the parapineal body leads to the left habenular
ganglion. Since the habenular ganglia are paired structures, it would seem
that pineal and parapineal eyes were originally paired and have only secon-
darily come to lie one on top of the other. The pineal eye contains a pigmented
 PHYLUM CHORDATA 189
retina with sensory cells, and a flattened imperfect lens. In the parapineal
body these structures are extremely reduced and hardly recognisable.
The pineal apparatus is better developed than in any of the higher forms
now living, and Young has shown that it is of vital importance in the everyday
life of the Lamprey. As light conditions in the environment change, so does
the colour of the larval form. The pineal apparatus is photosensitive, and it
seems that impulses are conveyed via the well-developed hypothalamus (in the
ventral region of the diencephalon) to the endocrine pituitary gland. The
pituitary or hypophysis is closely applied to the hypothalamus : there is no
hypophysial stalk, such as is developed in the higher Vertebrata. The gland
is a compound one, consisting of a comparatively large pars anterior and

FIG. J2z.-Petromyzm1: Brain with olfactory and nasa-hypophysial ducts. Cblm. cerebellum ;
crb. h. olfactory lobe; dien . diencephalon; f. fold in nasal duct; gl. accessory olfactory organ ;
inf. infundibulum; I. gn. hb. left habenular ganglion ; m ed. obl. m edu lla oblongata; na. ap . nostr il ;
nch. notochord; Nv. I, olfactory ner ve; Nv. :2, optic n. ; Nv. 3, oculomotor n.; Nv. 4, trochlea r
n.; Nv. 5, trigeminal n.; Nv. 6, adbucens n. ; Nu. 7, facial n. ; Nv . S, auditory n .; Nv. IO, vagus
n.; Nv. 1 2, hypoglossal n. ; olf. cp. olfactory capsule ; olf. I. olfactory bulb; olf. m . rn. ol·
factory mucosa; opt. l. optic lobe; pn. parapineal organ; pn. e. pineal eye; pty. b. pituitar y
gland; pty. p. hypophysial sac; sp. median septum of olfactor y capsule; sp. I, dorsal root of
Jlrst spinal nerve. (Combined from figures by Ahlborn and Ka:nsche.)

a flattened pars intermedia. These are separated from the third ventricle
by a thin tract of epithelial tissue which may represent the pars nervosa (of
the posterior lobe) of higher forms. There is experimental evidence that the
expansion of melanophores, causing the Lamprey's skin to darken, is caused
by a secretion from the intermedia when this is stimulated by nerve impulses
originating from the pineal.
In front of the diencephalon are paired bean-like masses, the cerebral
hemispheres, each consisting of a small posterior portion, the olfactory lobe (crb.
h.) and a larger olfactory bulb (olf. l.).
The cavity of the fore-brain extends laterally into these hemispheres
to form rudimentary lateral ventricles. The part of the lateral ventricle
which is contained in the olfactory bulb is sometimes referred to as the
rln'noca:le.
rgo ZOOLOGY
Organs of Special Sense.-The external nostril (Fig. no, n.; Fig. uS, na.)
leads by a short passage into a rounded olfactory sac (Fig. II8, olf.) placed
just in front of the brain and having its posterior wall raised into
ridges covered by the olfactory mucous membrane (Fig. r2r, olf. m. m.).
Here are located specialised olfactory receptor-cells, and from these travel axons
which become aggregated in the short olfactory nerve (Fig. r2o, nv. I). This
joins the highly organised olfactory bulb (olf. l.) close by. From the bottom of
hyp. ' ·

v.
FIG. 122.-Petrmnyzon: Development. Five stages in the development of the cavities of the
anterior part of the body. A , B, C- successive embryonic stages; D-ammoccete; E-adult.
b. t. buccal tentacles; end. endoderm; g. cesophagus ; h. m. hind-mouth ; hs. hydrosinus ; hyp. r.
hypophysial rudiment ; hyp. s. hypophysial sac ; inf. infundibulum ; I. I. lower lip; na. nasal
a perture ; na. r. nasal rudiment ; nc. notochord; olj. c. olfactory capsule ; p . ant. pars anterior of
pituita ry; ph. pharynx ; pin . pinea l eye : pin. r. pineal rudiment : p . int. pars intermedia of
pituitary; pit. p ituitary gla nd; st. stomod<eum; 11 . l. upper lip; v. v elum; v. t. velar tentacles.
(Drawn by R . Strahan, after various authors.)

the olfactory sac is given off a large nasa-hypophysial sac (Fig. u8, n. hyp., Fig.
121, pty. p.) which extends downwards and backwards between the brain and the
skull-floor, passes through the basicranial fontanelle, and ends blindly below the
anterior end of the notochord. This elongated, compressible pouch seems to
act in a manner analogous to the rubber bulb of a pipette. The expanded and
closed end of the pouch lies directly above the dorsal wall of the pharynx and
between the first pair of gill-pouches. Directly above the pouch is the un-
yielding anterior end of the notochord (Fig. 121, nch.). Respiratory movements
squeeze the water in the pouch in a forward direction out through the nostril
 PHYLUM CHORDATA 191

(na. ap.) situated on the dorsal surface of the head. When the pressure relaxes,
a fresh stream of water flows into the pouch, and thus comes in contact with
the olfactory receptor-organs. If the aperture is experimentally blocked, the
Lamprey no longer swims away from noxious substances liberated in its vicinity.
The relations between the olfactory sac, nasa-hypophysial sac, and the
pituitary body are extraordinary. In the embryo, before the stomodreum
(Fig. 122, A, st.) communicates with the gut, two unpaired ectodermal invagina-
tions appear in front of the mouth. The foremost of these is the nasal rudiment
(na. r.). The other (which is situated between the olfactory sac and the

FIG. 123.- Lampetra: Eye and adjacent structures. (L. jluviatilis, anterior end to the left .)
Accommodation is accomplished by a lteration of the shape of the eyeball by the extrinsic cornea its
muscle. The tendon of this muscle is inserted on the rim of the spectacle, a specialised window-
like area of transparent skin. Muscle contraction draws the spectacle taut and fl attens
the subjacent cornea. This in turn is in close contact with the lens which is thus pushed
closer to the retina. The elasticity of sclera a nd vitreous b ody, and the equality of the intra -
ocular pressure, returns the eyeball to the resting shape. av. anterior surface of vitreous; c.
cornea; cr. external rectus; io. inferior oblique; ir. internal rectus; s. spectacle; sk. skin; sp.
space between spectacle and cornea; sr. superior rectus; t. t endon of comealis; v. venous sinuses
(cushioning the eyeball). (After Walls.)

mouth) is the hypophysial rudiment (hyp . r.), which in this case opens just
outside the stomodreum instead of within it as in other Craniata. Its inner
or blind end extends to the ventral surface of the fore-brain and terminates just
below the infundibulum (inf.). As development proceeds, the olfactory and
hypophysial invaginations become sunk in a common pit (B), which, by the
growth of the immense upper lip (up. l.), is gradually shifted to the top of the
head (C, D). In the ammoccete (p. rg6), this common nasa-hypophysial cavity
extends only a short distance under the forebrain. The hypophysial rudiment
grows out as a thin plate of tissue. This buds off (r) the intermedia, which
interdigitates with the nervous tissue from the floor of the third ventricle, and
(z) the anterior body of the pituitary gland. Later, all connection with the
rgz ZOOLOGY
naso-hypophysial cavity is lost. At metamorphosis the last-named grows out
into its adult condition independently of the pituitary gland. The peculiar
monorhinal condition of the lamprey is due to the hypertrophy of the upper
lip. This, in turn, is related to the function of the upper lip, which is the bur-
rowing organ of the ammocrete. We may therefore assume, with Watson, that
the cephalaspids, anaspids, and hags similarly possessed ancestors in which the
upper lip performed this function.
The paired eyes of Petromyzon are essentially similar to those of other
vertebrates. Each is moved by six external muscles, as in higher forms, but
the arrangement of these is peculiar. The accommodation of the eye is brought
about by an extraordinary arrangement (Fig. 123). A remarkable cornealis
muscle, which arises from the special-
i.e.
a.c.
isation of two anterior myotomes, pulls
on the cornea from one side and flattens
it. This corneal movement thrusts the
lens in towards the retina. Thus, the
.. ::<..
?:iJtfi"'
·:;,~ , whole lens moves in a manner roughly
similar to the lens of a camera, being
cr. w. omp moved back and forth in relation to the
end. d. ~........_ sacc. photographic plate. It is said that the
mac.
cornea of the lamprey does not move
when the position of the rest of the eye
Frc. 124.-Petromyzon: Membranous
labyrinth (left). a. c. anterior semicircular is altered by the extrinsic muscles.
canal; amp. ampulla; cr. crista; end. d.
endolymphatic duct; i . c. invagination canal; The membranous labyrinth (Fig.
mac. m acula; p. c. posterior semicircular 124) is remarkable in having only two
canal; sacc. sacculus; utr. utriculus. (Re-
drawn from Neal and Rand, after Hesse.) semi-circular canals. These, the an-
terior (a. c.) and posterior semi-circular
canals (p. c.), curve at right angles to each other and are equipped with a
sacculus (sacc.) and utriculus (utr.). Little or nothing is known concerning the
actual hearing capacity of Petromyzon. Sensory apparatus within the above
compartments appreciates changes in the animal's orientation, and resultant
nerve-impulses lead to appropriate reflex movements and keep it on an even
keel.
In eye ostomes the lateral line sense-organs (p. 136) are relatively simple and
are exposed to the surrounding water. They occur in the head region and in
a single line extending along each side of the body. They are innervated by
cranial nerves; those on the body and tail are connected to the central nervous
system by a branch of the vagus.
There is experimental evidence that skin photoreceptor organs occur in the
tail of the ammocrete larva at least ; the work of Young suggests that the caudal
lateral line organs are involved. Finally, organs of taste occur in the wall of
the pharynx between the gill-sacs.
 PHYLUM CHORDATA 193
Urinogenital System.-This shows a great advance on that of Amphioxus.
Complicated organs of excretion, the kidneys (Figs. II4, 125, mes.), have
appeared. These enable the animal to eliminate excess fluid and waste
nitrogenous and other substances, and at the same time maintain an appro-
priate osmotic balance whether it is living in a fresh- or a salt-water medium.
The kidneys are long, strap-shaped bodies developed from the mesonephros
of the embryo. The essential renal units within the adult kidney are numerous
minute Malphigian corpuscles. Each contains a 'knot' of microscopic blood-

in!. mes. mes. d. g .d. cl. u. g .p. my.

FrG. I 25.-J>efrotnyzon: Cloacal region. P. marinus. cd. posterior cardinal vein; cl. cloaca
c. r. cartilaginous rods(' fin rays'); d. ao. dorsal aorta; d. j. 2 second dorsal fin; g. d. genital pore
int. intestine; m. muscles of fin; m es . mesonephros; mes. d. mesonephric duct; my. myotome
n. a. neural arch; nc. notochord; u. g. p. urinogenital papilla. (Redrawn after Goodrich .)

vessels, the glomerulus, and opens into a winding tubule (p. 155). The heart-
beat pumps blood through the glomeruli. Excess fluids and excretory products
are shed down the tubules towards the exterior.
Each kidney is attached along one edge to the dorsal wall of the body-
cavity by part of the peritoneum. Along the other (free) edge runs the meso-
nephric duct (Fig. II4, mes. d.), which opens posteriorly into a small urino-
genital sinus placed just behind the rectum. This opens by a urino-genital
papilla into a pit, the cloaca, in which the alimentary canal also ends. The
side-walls of the sinus are pierced by a pair of small apertures, the genital pores
(Fig. 125, g. d.), which place its cavity in communication with the crelom.
VOL. II. N
194 ZOOLOGY
The kidneys contain lymphoid tissue where, as has been mentioned, blood
cells may be formed. This tissue also performs a phagocytic function by
destroying old blood cells and removing suspended material from the circulation.
The sexes are separate in the adult. However, sexual differentiation occurs
at a comparatively late stage, and so both oocytes and spermatocytes may be
found in a single gonad in the young Lamprey. The gonad (Fig. n8, gon.,
Fig. II4, ts.) is unpaired. The ovary in the female ruptures and extrudes
eggs directly into the ccelom, as in higher vertebrates. The testis, however,
is unique among vertebrates in the manner in which it discharges its sperm-
atozoa. The sperm-containing follicles of the testis rupture and the sperm-
atozoa, like the ova, are shed into the ccelom. The gametes escape through
the pair of genital pores mentioned above. These, alike in each sex, lead
as short canals to the urinogenital sinus from which eggs or sperms escape
to the exterior where fertilisation takes place. In adult males there occur
prominent aggregations of interstitial Leydig-eells in the typical vertebrate
arrangement (Fig. 97, p. 152).
Development-The egg is telolecithal, having a considerable accumulation
of yolk in the v"egetal hemisphere. In correspondence with this, cleavage is
complete but unequal, a condition comparable with that in the frog (p. 418).
Early segmentation divides the animal hemisphere into a number of small,
relatively yolk-less micromeres (Fig. 126, A, micr.), and the vegetal hemisphere
into a smaller number of yolky macromeres (A, y. end.). A cavity, the blastocccle
(blc.), develops between these two cell Ia yers. The faster -dividing micro-
meres grow around to enclose the macromeres and also invaginate through a
crescentic groove, the dorsal lip (B, d. l.) of the blastopore (B, blp.) just below
the equator of the egg. As development proceeds, the blastopore moves to-
wards the vegetal pole of the embryo (C) and later passes it (D). In contrast
to the condition in Rana (p. 418), the position of the blastopore bears no relation
to the plane of the first cleavage. The process of invagination through the
blastopore gives rise to the cavity of the archenteron (C, arch.). The ventral
wall of this cavity is formed from the macromeres, while the dorsal and lateral
walls are formed from invaginated chorda-mesoderm (C, ch. m.) which later
develops into the notochord (D, nch.) and somites (F, sam.). Somewhat later,
a groove appears in the endodermal floor and the sides of the groove come
together to form an endodermal roof (F, end. r.) to the gut. Apart from this
contribution, the macromeres take no part in organ-building. At the time of
hatching they still retain considerable food stores. Owing to the mass of this
tissue, the lumen of the archenteron is much narrower than in Amphioxus.
By the completion of gastrulation, the blastopore has taken up a position
at the postero-ventral end. The development of the central nervous system
differs widely from the corresponding process in Amphioxus, and is only
approached among the Craniata by the bony fishes. The dorsal surface be-
 PHYLUM CHORDATA rgs
comes flattened along a narrow longitudinal area, and along this a groove
appears. This terminates immediately in front of the blastopore. The area
along which the groove runs soon becomes raised up above the general surface
and forms a narrow longitudinal elevation. Sections of this stage show that
the ectoderm has developed a thickening along the course of the longitudinal
groove, and this comes to grow downwards towards the archenteron as a

blc.

Frc. rzG.-Peh·mnyzon: Development. Successive stages in gastrulation shown in recon-

structions from which parts of the embryo have been removed to reveal internal structures. The
centre of gravity shifts during gastrulation and this affects the orientation so that the region
marked by the arrow eventually comes to lie dorsally. The shift in orientation is a gradual one,
but for convenience is shown in two stages. an. anus; anh. archenteron; blc. blastocrele; blp.
blastopore; ch. nz. chorda-mesoderm; d. l. dorsal lip of blastopore; ect. ectoderm; end. r. endo-
dermal roof of archenteron; mes. mesoderm; micr. micromere layer; n. c. nerve cord; nch. noto-
chord; n. k. neural keel; sam. somite; slam. stomod<eal invagination; y. end. yolky endoderm.
(Drawn by R. Strahan, reconstructed from Brachet after Weissenberg and de Sely-Longchamps.)

solid longitudinal neural keel. This is the rudiment of the central nervous
system. Subsequently the keel becomes separated off from the surface ecto-
derm, and lies below it as a solid cord. It is only at a considerably later period
that a lumen appears in this cord, and gives rise to the ventricles of the brain
and the central canal of the spinal cord. In contrast to the condition in
Amphioxus, somite formation is limited to the dorsal mesoderm on either side
of the mid-line, a feature which is also characteristic of the remainder of the
Craniata. The blastopore does not close; it is converted into an anal aper-
ture, so that there is no proctodceum. The dorsal lip of the blastopore, very
rg6 ZOOLOGY
prominent from the first , is produced into the rudiment of the tail region. The
mouth is developed later than the anal aperture by stomodreal invagination.
The membrane previously separating the fore-gut from the mid-gut becomes the
velum (Fig. 122, v.) of the larva.
The young is hatched as an ammoca:te larva (Fig. 127) some 10 mm. long.
This differs from the adult in several important respects. The median fin is
continuous. There is a semi-circular, hood-shaped ztpper lip (u. l.) instead of
the suctorial buccal funnel of the adult. Teeth are absent. A set of branched
buccal tentacles (Fig. 119, b. tent.) guards the entrance to the alimentary canal.
Behind the tentacles lies the velum (vel.), a pair of cup-shaped structures
attached to the anterior wall of the pharynx.
A B These act as paddles and maintain a stream
-na .ap of water through the pharynx. In this
e-Ye they are assisted by pumping movements
of the pharynx. Although the ammoccete
l .l / burrows in mud, it feeds by taking in
fairly clean water from the oral end of
its burrow. The endostyle (end.), homo-
logous with that of Amphioxus, passes a
stream of mucus forward a.long a groove
in the floor of the pharynx to a pair of
peripharyngeal grooves behind the velum.
Movements of the velum dislodge strands
of mucus which form a sort of net across
t the entrance to the pharynx. Further
FIG . 127.- Lcnnpetra: Development. down the pharynx, the strands come to-
Head of larval L. fluviatilis. A, from be- gether to form a cord which passes to the
n eath; B, from the side. br. 1, first
branchial aperture; eye, eye; l. l. lower cesophagus. Propulsion is now achieved
lip; na. ap. n ostril; u. l. upper lip.
(After W. K Parker.) by the action of a:sophageal cilia. Food is
in this manner filtered from the water
current and passed to the digestive region of the alimentary canal without being
diluted with a great amount of water. In Amphioxus the feeding current is
maintained by cilia; in the ammoccet e by muscles.
This evolution of a muscular, in place of a ciliary, ingestive apparatus was
an immense step forward in evolution. It enabled a far swifter and greater
intake and, consequently, the development and sustenance of far bigger
animals.
During its early life the larva spends most of its time buried in a U-shaped
burrow in the mud, emerging only at night to change its feeding ground. It
swims with its head downwards and then burrows again with great rapidity.
Two major instinctive responses combine to maintain the ammoccete in its
proper environment: photokinesis and thigmotaxis. The photokinetic response
 PHYLUM CHORDATA 197
is shown when ammoccetes are illuminated. Their reaction is to swim in a
random fashion until reaching a shaded area, when locomotion becomes much
more limited. The thigmotactic response is shown when ammoccetes are given
the alternative of a smooth bottom or tubes into which to swim. Spontaneous
movements are much fewer when touch receptors are stimulated over a large
part of the body-surface than when only a limited number are, so that animals
tend to stay in a tube after reaching it, even if the illumination is fairly high.
Together these responses lead the animals to seek crevices or burrow in dark
places. A batch of photoreceptors occur in the skin of the tail, as Young has
shown. These presumably act as an extra warning device to prevent the animal
from leaving its tail exposed above the burrow.
We have seen that larval lampreys exhibit a muscular feeding device, yet
retain an endostyle, a characteristic part of the feeding mechanism of ancestral
forms. At the time of metamorphosis, the endostyle becomes modified and
develops into an entirely different structure that occurs throughout the rest
of the vertebrate series-the endocrine thyroid gland. The endostyle loses
its mucus-secreting capacity and develops a series of iodine-containing
vesicles that are clearly of the same nature as the thyroid follicles of higher
animals.
At the same time the buccal region becomes elongated to accommodate the
tongue ; and its margin becomes circular. The tentacles recede and are re-
placed by teeth on the buccal funnel and tongue. The velum becomes reduced,
and skeletal rods develop from it to form a filter guarding the respiratory
pharynx: food material is thus diverted to the cesophagus, which has already
been cut off from the pharynx proper. The gills are modified and open into
the pharynx. The nasa-hypophysial sac becomes enlarged posteriorly. The
larval cranium, which consists of little more than the parachordals, trabeculre,
and olfactory and otic capsules, becomes filled out into the adult condition.
The skeleton of the upper lip and tongue, which is absent in the larva, also
develops during metamorphosis. The eyes complete their development and
move to the surface. The continuous dorsal fin is excavated to form the hvo
dorsal, and single caudal fins; and the colour of the skin changes from muddy
brown to metallic on the dorsal surface. In some species, brilliant-coloured
stripes also appear at this stage. The Lamprey now leaves its river habitat
for the open sea and begins the carnivorous phase of its life-history.

CLASS MYXINOIDEA
The exclusively marine hags or slime-eels are exemplified by the widespread
genera Myxine, Paramyxine, and Eptatretus (= Bdellostoma). These inhabit
muddy bottoms, where they rest in temporary burrows. They feed on a wide
range of dead animals and may perhaps prey upon living polychaetes and
rg8 ZOOLOGY
priapuloids. They eat dead or dying fish, but the old view that they are
parasitic upon fishes is incorrect. Their method of feeding is unique among
vertebrates in that they bite with protrusible horny teeth (Fig. 130, d. pl.)
which move against each other from side to side. Other unique features
include the direct communication between the nostril and the pharynx, the
existence of auxiliary hearts in the caudal and visceral circulation, and the
fusion of the two semicircular canals form a ring-shaped structure.

COMPARISON OF THE MYXINOIDEA WITH THE LAMPREY

Myxinoids show a much greater range of variation than lampreys. Here
we shall only indicate the points in which they differ from lampreys and from
each other. The body of myxinoids is
n.r.
cylindrical and from 40 to So em. long.
n n

They are remarkable for the immense

amount of slime they can produce when
disturbed. The slime comes from thread
cells, peculiar to m yxinoids, which occur
singly in the epidermis (Fig. II3) or
massed in the segmental slime-glands
(Fig. rz8). During cell development a
.
. .. . . continuous thread of fibrous protein is
. ·:. !~ laid down within the membrane. When
. .. ·.::·: the hag is stimulated by agitation or local
: :· ~ . .:.. :
---:--~ . br. op. pressure, large numbers of these cells
...
I
-·~ .
burst, releasing their threads, which are
·:.:: -. -. ;.:~.
-':~.:-. : ·. from r to 3fL in diameter and several em.
in length. The threads absorb water
t':. . .
0 : • • .:

,. • • 0 • ~

·. };~ c. op almost instantaneously and form around

the body a tenacious slimy gel as much as
ro em. thick. One m edium-sized Myxine
can gelatinise as much as 500 cc. of sea
·::·~; water in less than a minute.
A B The buccal cavity is even longer than
FIG. I28. -Class Myxinoidea: Myxine in lampreys, but the mouth is not ex-
glutinosa (A } a nd Eptatretus (= Bdello-
stoma) stoutii (B) from below. br. ap ., , panded into a funnel. The mouth and
first branchia l aperture; c. ap . common h ·1 h d
branchial aperture; mt!t. mouth; n . nostril; t e nostn are eac surroun ed by
n. t.,, 2 nasal tentacles; o. t .1. 2 oral t entacles; tentacles (Fig. 128) supported by carti-
ph. cut. aperture of pharyngo-cutaneous
duct; st. gl. slime g lands. (Drawn by R. !ages (Fig. 130, tent. 1- 4). There is a
Strahan.) single median palatine tooth (Fig. 129,
pal. t.) above the oral aperture, and two rows of smaller teeth arising from the
dental plate (Fig. 130) on the 'tongue'. The papill~ beneath the cone-like horny
 PHYLUM CHORDATA rgg
teeth bear a closer superficial resem-
blance to rudiments (or vestiges) of true
calcified teeth than is the case in
lampreys; but it appears that no odon-
toblasts and no calcified substance of any
kind are formed in connection with them.
The terminal nostril opens into a tubular
nasal duct (Fig. rzg, na. d.) leading to
the olfactory organ and is continuous
with the naso-hypophysial duct (na.
hyp.) which opens into the buccal cavity
just anterior to the velum (vel.). This
is homologous with the velum of the
ammoccete and, although having a
much more complex structure, performs
the same function of maintaining the
respiratory current. The continuous
flow of \Vater over the olfactory mucosa
permitted by this arrangement is pro-
bably valuable to a blind scavenger.
The only fin is a narrow caudal fin
surrounding the end of the tail. The
respiratory organs present striking
differences in the major genera. In
Eptatretus there are in different species
six to fourteen very small external
branchial apertures (br. ap . I) on each
side. Each of these communicates by a
short efferent duct with one of the gill-
pouches which is again connected with
the pharynx by another tube. Behind
and close to the last gill-slit, on the left
side, is an aperture leading into a tube,
the pharyngo-cutaneous duct (ms. eztl.),
which opens directly into the pharynx.
In A1yxine the efferent ducts (Fig. rzg,
ef. d.) all unite together before opening
on the exterior, so that there is only a
single external branchial aperture (br. ap.)
on each side. The pharyngo-cutaneous
duct (ms . cut.) opens into the left ex-
ternal branchial aperture (br. ap.) and
200 ZOOLOGY
serves for the expulsion of inhaled particles too large to enter the afferent
ducts.
The spinal cord (sp. c.) is covered only by fibrous tissue. There is no
trace of neural arches in the trunk, but in the posterior part of the caudal
region both neural canal and notochord are enclosed in a continuous cartila-
ginous plate. Similarly, the roof of the skull is entirely membranous (Fig. 130) .
The skull has a fundamental similarity to that of lampreys, but this is obvious

a.v.n.b.
pal. b.

ex. br. 1

vel. sk.

corn. c.

,.....~ ·3

FIG. IJo.-1Yfy:rine: Skull of adult. il!I. glutinosa. Portion of the palatine bar is removed on
left side. a. v. n. b. anterior vertical nasal bar; bas. basal cartilage; br. 1 Jst (internal) bra nchial arch ;
corn. c. cornual cartilage; d. l . b. dorsal longitudinal bar ; d. pl. dental plate; ex. br. 1 , 2 1st and
2nd extrabra nchial arches; ex. pq. extrapalatoquadrate; hy. hyoid arch; hyp. c. hypophysial
cartilage ; n c. notochord ; ot. otic capsule ; pal. b. p a latine bar; pelt. parachordal; pq. palata.
quadrate; p. v. n. b. posterior vertical nasal bar; s. n . c. subnasal cartilage; tent.,, 2 , 3 , 4 tentacular
cartilages; trab. trabecula; v. br. b. ventral branchial bar ; vel. sk. velar skeleton. (Drawn by R.
Strahan, reconstructed from Cole.)

only in the embryo (Fig. 131). Later developments, associated with the growth
of the nasa-hypophysial sac, the oral tentacles, and the backward migration of
the gills, bring about a considerable distortion. The nasal duct (Fig. 129, na. d.)
is supported by rings of cartilage (na. c.), and the oral tentacles by
flexible rods of the same tissue. The cornual cartilage (Fig. 131, corn. c.) and
the subnasal cartilage (s. n. c.) represent the skeleton of the upper lip of lampreys
and the basal plates (Figs. 1 29 and 131, bas.) are, in part, the homologue of the
lamprey tongue skeleton. The sub-ocular arch (Fig. 130, ex. pq., pq., hy.) re-
presents anteriorly the visceral skeleton of the mandibular arch and posteriorly,
 PHYLUM CHORDATA 201

the hyoid arch. Of the remaining branchial arches, only the first two are
represented, these being the first extra-branchial cartilage (ex. br. I.), and the
first branchial cartilage (br. I), and the second extra-branchial cartilage (ex. br. 2).
In the region of the common branchial apertures a pair of asymmetrical
cartilages (Fig. 129, p. br. sl?.) may represent a posterior remnant of the
branchial skeleton.
The myotomes of one side alternate with those of the other.

tent.3

FIG. I3L-My:rine: Skull of embryo. 111. glutinosa. a. v. n. b. anterior vertical nasal bar; bas,
basal skeleton; b·r. 1 rst (internal) br anchia l arch; corn. c. corn ua l cartilage; d. l. b. dorsal longi-
tudinal bar; ex. br. 1 , 2 rst and 2nd extrabranchial arches; PX. hy. extrahyal arch; ex. pq. extra-
palatoquadrate; hyp. com. hypophysial commissure; lab. c. labial cartilage; n c. notochord; ot.
otic capsule ; pal. b. palatine b a r ; pch. p a rac hordal; pq. palatoquadrate ; p . v. n . b. posterior
vertical nasal bar; s. n . c. subnasal cartilage ; tent.1 , 2 . , tentacu lar cartilages ; trab. trabecula ;
trab. com. trabec ular commissure; v. br. b. ventral branchial bar. (Drawn by R. Strahan, recon-
structed from Holmgren.)

The intestine is wide. The liver consists of separate anterior and posterior
portions, the ducts of which open separately into the gall-bladder. A pancreas-
like gland occurs in both Myxine and Eptatretus in a position comparable with
that of the 'insular organ' of lampreys (p. 185). There is no definite spleen,
but blood-forming lymphoid tissue is found scattered through the submucosa
of the gut. Blood cells may also be formed in the pronephros.
The brain differs considerably from that of Petromyzon, especially in the
larger olfactory lobes, the reduced ventricles, and the smaller mid-brain. The
dorsal and ventral roots of the spinal nerves unite instead of remaining separate.
202 ZOOLOGY
It is important to realise, however, that the junction is different from that
typical of the Gnathostomata.
The eyes are vestigial and sunk beneath the skin, which, as in the ammo-
crete (p. rg6), is light-sensitive. When stimulated these provoke a photo-
kinetic response. The membranous labyrinth (Fig. 132) has only a single semi-
circular canal, which, having an ampulla at each end, probably represents
both anterior and posterior canals (p. 145). Hags lack a pineal organ and a
lateral line system. In comparison with lampreys, their sensory system is
extremely limited.
The pronephros is retained in adult myxinoids as a pair of small bodies
lying dorsal to the pericardia! cavity, into which the nephrostomes open. In
Eptatretus, the tubules open into an incomplete
a.p.c. pronephric duct which, however, has no open-
ing to the exterior. Myxine lacks a pronephric
duct and the tubules end blindly. The func-
tional kidney is the mesonephros, and is speci-
end. d. ally interesting from the fact that in myxinoids
it retains in the adult its primitive segmental
arrangement. The 'ureter' sends off in each
segment a coiled tubule with a single Bowman's
capsule, into which a branch from the aorta
utr. sacc. enters and forms a glomerulus.
FIG. IJZ.-My;rine: Membranous Despite many statements to the contrary,
labyrinth (left). a. p. c. fused anter-
ior and posterior canals ; cr. crista; the sexes are probably separate in Myxine but,
end . d . endolymphatic duct; utr . sacc. as in Lampreys, differentiation of the immature
urticulo-sacculus. (Redrawn from
Neal and Rand, after Hesse.) gonad does not occur until late in life. The
female may thus retain an undeveloped testis
and the male a rudimentary ovary. The large, cylindrical eggs are enclosed
in a horny shell bearing terminal hooked processes.
The early development of Myxine is unknown, but there is no reason to
assume that it differs markedly from Eptatretus, in which segmentation is
markedly meroblastic, being confined to one pole of the elongate egg. The
blastoderm thus formed extends gradually over the surface of the yolk, which
it completely encloses only at a late stage, when the gill-clefts are all formed.
Eptatretus differs from Petromyzon and resembles the majority of the Craniata
in the mode of development of the central nervous system. This is formed not
from a solid ectodermal keel, but from an open medullary groove. The lips
of the groove bend inwards and unite to form a medullary canal. There is no
metamorphosis in the life-history.