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between the leaves. These gillbearing appendages can be flapped to
and fro, and they seem to be at times held apart by the flabellum, a
spatulate process which Patten and Redenbaugh regard as a
development of the median sensory knob on the outer side of the
coxopodite of the last pair of walking limbs.
Limulus has no trace of
Malpighian tubules, structures
which seem often to develop only
when animals cease to live in
water and come to live in air. The
Xiphosura have retained as
organs of nitrogenous excretion
the more primitive nephridia, or
coxal glands as they are called, in
the Arachnida. They are redbrick
in colour, and consist of a
longitudinal portion on each side
of the body, which gives off a lobe
opposite the base of the pedipalps
and each of the first three walking
legs—in the embryo also of the
chelicerae and last walking legs,
but these latter disappear during
development. A duct leads from
Fig. 155.—Diagram of the first gill of
Limulus, from the posterior side,
the interior of the gland and
showing the distribution of the gill- opens upon the posterior face of
nerve to the gill-book (about natural the last pair of walking legs but
size). After Patten and Redenbaugh. 1, one.
Inner lobe of the appendage; 2, outer The nervous system has been
lobe of appendage; 3, median lobe of
appendage; 4, gill-book; 5, neural nerve
very fully described by Patten and
of the ninth neuromere; 6, internal Redenbaugh, and its complex
branchial nerve; 7, gill-nerve; 8, nature plays a large part in the
median branchial nerve; 9, external ingenious speculations of Dr.
branchial nerve. Gaskell as to the origin of
Vertebrates. It consists of a stout
ring surrounding the oesophagus
and a ventral nerve-cord, composed—if we omit the so-called fore-
brain—of sixteen neuromeres. The fore-brain supplies the median
and the lateral eyes, and gives off a median nerve which runs to an
organ, described as olfactory by Patten, situated in front of the
chelicerae on the ventral face of the carapace. Patten distinguishes
behind the fore-brain a mid-brain, which consists solely of the
cheliceral neuromere, a hind-brain which supplies the pedipalps and
four pair of walking legs, and an accessory brain which supplies the
chilaria and the genital operculum. This is continued backward into a
ventral nerve-cord which bears five paired ganglia supplying the five
pairs of gills and three pairs of post-branchial ganglia; the latter are
ill-defined and closely fused together. As was mentioned above, the
whole of the central nervous system is bathed in the blood of the
ventral sinus.
The sense-organs consist of the olfactory organ of Patten, the
median and lateral eyes, and possibly of certain gustatory hairs upon
the gnathobases. The lateral eyes in their histology are not so
differentiated as the median eyes, but both fall well within the limits
of Arachnid eye-structure, and their minute anatomy has been
advanced as one piece of evidence amongst many which tend to
demonstrate that Limulus is an Arachnid.
Both ovaries and testes take the form of a tubular network which is
almost inextricably entangled with the liver. From each side a duct
collects the reproductive cells which are formed from cells lining the
walls of the tubes, and discharges them by a pore one on each side of
the hinder surface of the genital operculum. As is frequently the case
in Arachnids the males are smaller than the females, and after their
last ecdysis the pedipalps and first two pairs of walking legs, or some
of these appendages, end in slightly bent claws and not in chelae. Off
the New Jersey coast the king-crabs (L. polyphemus) spawn during
the months of May, June, and July, Lockwood states at the periods of
highest tides, but Kingsley[217] was never “able to notice any
connexion between the hours when they frequent the shore and the
state of the tide.” “When first seen they come from the deeper water,
the male, which is almost always the smaller, grasping the hinder
half of the carapace of the female with the modified pincer of the
second pair of feet. Thus fastened together the male rides to shallow
water. The couples will stop at intervals and then move on. Usually a
nest of eggs can be found at each of the stopping-places, and as each
nest is usually buried from one to two inches beneath the surface of
the sand, it appears probable that the female thrusts the genital plate
into the sand, while at the same time the male discharges the milt
into the water. I have not been able to watch the process more closely
because the animals lie so close to the sand, and all the appendages
are concealed beneath the carapace. If touched during the
oviposition, they cease the operation and wander to another spot or
separate and return to deep water. I have never seen the couples
come entirely out of the water, although they frequently come so
close to the shore that portions of the carapace are uncovered.”[218]
Fig. 156.—A view of the nervous system of Limulus from below.
(About natural size.) After Patten and Redenbaugh.
The brain lies upon the neural side of the endosternite, and the
ventral cord (22) passes back through the occipital ring. The
neural nerves are cut off, but the left haemal nerves and those
from the fore-brain (12) are represented entire.
From the fore-brain a median olfactory nerve (9) and two lateral
ones (8) pass forward to the olfactory organ; a median eye-nerve
(2) passes anteriorly and haemally upon the right of the
proventriculus (3) to the median eyes; and a pair of lateral eye-
nerves pass to the lateral eyes (15).
The first haemal nerve, or lateral nerve, follows the general course
of the lateral eye-nerve, but continues posteriorly far back on to
the neural side of the abdomen.
The haemal nerves of the hind-brain radiate from the brain to the
margins of the carapace, and each one passes anterior to the
appendage of its own metamere. The integumentary portions
divide into haemal and neural branches, of which the haemal
branches (5) are cut off. Each haemal branch gives off a small
nerve which turns back toward the median line upon the haemal
side of the body.
Intestinal branches arise from all the haemal nerves from the sixth
to the sixteenth, and pass to the longitudinal abdominal muscles
and to the intestine.
Cardiac nerves arise from all the haemal nerves from the sixth to
the thirteenth. Six of the cardiac nerves communicate with the
lateral sympathetic nerve (24), which innervates the branchio-
thoracic muscles (16).
The developing ova and young larvae are very hardy, and in a little
sea-water, or still better packed in sea-weed, will survive long
journeys. In this way they have been transported from the Atlantic to
the Pacific coasts of the United States, and for a time at any rate
flourished in the western waters. Three barrels full of them
consigned from Woods Holl to Sir E. Ray Lankester arrived in
England with a large proportion of larvae alive and apparently well.
According to Kishinouye, L. longispina spawns chiefly in August
and between tide-marks. “The female excavates a hole about 15 cm.
deep, and deposits eggs in it while the male fertilises them. The
female afterwards buries them, and begins to excavate the next
hole.”[219] A line of nests (Fig. 157) is thus established which is always
at right angles to the shore-line. After a certain number of nests have
been formed the female tires, and the heaped up sand is not so
prominent. In each “nest” there are about a thousand eggs, placed
first to the left side of the nest and then to the right, from which
Kishinouye concludes that the left ovary deposits its ova first and
then the right. Limulus rotundicauda and L. moluccanus do not bury
their eggs, but carry them about attached to their swimmerets.
The egg is covered by a leathery egg-shell which bursts after a
certain time, and leaves the larva surrounded only by the
blastodermic cuticle; when ripe it emerges in the condition known as
the “Trilobite larva” (Fig. 158), so-called from a superficial and
misleading resemblance to a Trilobite. They are active little larvae,
burrowing in the sand like their parents, and swimming vigorously
about by aid of their leaf-like posterior limbs. Sometimes they are
taken in tow-nets. After the first moult the segments of the meso-
and metasoma, which at first had been free, showing affinities with
Prestwichia and Belinurus of Palaeozoic times, become more
solidified, while the post-anal tail-spine—absent in the Trilobite larva
—makes its first appearance. This increases in size with successive
moults. We have already noted the late appearance of the external
sexual characters, the chelate
walking appendages only being
replaced by hooks at the last
moult.
Fig. 157.—The markings on the sand
made by the female Limulus when
depositing eggs. Towards the lower end
the round “nests” cease to be apparent,
the king-crab being apparently
exhausted. (From Kishinouye.) About
natural size.
Fig. 158.—Dorsal and ventral view of the last larval stage (the so-
called Trilobite stage) of Limulus polyphemus before the
appearance of the telson. 1, Liver; 2, median eye; 3, lateral eye; 4,
last walking leg; 5, chilaria. (From Kingsley and Takano.)
Limulus casts its cuticle several times during the first year—
Lockwood estimates five or six times between hatching out in June
and the onset of the cold weather. The cuticle splits along a “thin
narrow rim” which “runs round the under side of the anterior
portion of the cephalic shield.”[220] This extends until it reaches that
level where the animal is widest. Through this slit the body of the
king-crab emerges, coming out, not as that of a beetle anteriorly and
dorsally, but anteriorly and ventrally, in such a way as to induce the
unobservant to exclaim “it is spewing itself out of its mouth.” In one
nearly full-sized animal the increase in the shorter diameter of the
cephalic shield after a moult was from 8 inches to 9½ inches, which
is an indication of very rapid growth. If after their first year they
moult annually Lockwood estimates it would take them eight years to
attain their full size.
The only economic use I know to which Limulus is put is that of
feeding both poultry and pigs. The females are preferred on account
of the eggs, of which half-a-pint may be crowded into the cephalic
shield. The king-crab is opened by running a knife round the thin
line mentioned on p. 275. There is a belief in New Jersey that this
diet makes the poultry lay; undoubtedly it fattens both fowls and
pigs, but it gives a “shocking” flavour to the flesh of both.
CLASSIFICATION.
But five species of existing King-crabs are known, and these are
grouped by Pocock into two sub-families: (i.) the Xiphosurinae, and
(ii.) the Tachypleinae. These together make up the single family
Xiphosuridae which is co-extensive with the Order. The following is
Pocock’s classification.[221] The names used in this article are printed
in italic capitals.
Order Xiphosura.
Family 1. Xiphosuridae.
Sub-Fam. 1. Xiphosurinae.
Sub-Fam. 2. Tachypleinae.
Fossil Xiphosura.[224]
BY