The eggs of Limulus are fertilized in the sea after they have been laid. Scorpio, being a terrestrial animal, fertilizes by copulation. The male possesses elaborate copulatory structures of a chitinous nature, and the eggs are fertilized in the female without even quitting the place where they are formed on the wall of the reticular gonocoel. The female scorpion is viviparous, and the young are produced in a highly developed condition as fully formed scorpions.
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Fig. 28.—The right coxal gland of Limulus polyphemus, Latr. a2 to a5, Posterior borders of the chitinous bases of the coxae of the second, third, fourth and fifth prosomatic limbs. b, Longitudinal lobe or stolon of the coxal gland. c. Its four transverse lobes or outgrowths corresponding to the four coxae. (From Lankester, loc. cit., after Packard.)
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Differences between Limulus and Scorpio.—We have now passed in review the principal structural features in which Limulus agrees with Scorpio and differs from other Arthropoda. There remains for consideration the one important structural difference between the two animals. Limulus agrees with the majority of the Crustacea in being destitute of renal excretory caeca or tubes opening into the hinder part of the gut. Scorpio, on the other hand, in common with all air-breathing Arthropoda except Peripatus, possesses these tubules, which are often called Malpighian tubes. A great deal has been made of this difference by some writers. It has been considered by them as proving that Limulus, in spite of all its special agreements with Scorpio (which, however, have scarcely been appreciated by the writers in question), really belongs to the Crustacean line of descent, whilst Scorpio, by possessing Malpighian tubes, is declared to be unmistakably tied together with the other Arachnida to the tracheate Arthropods, the Hexapods, Diplopods, and Chilopods, which all possess Malpighian tubes.
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Fig. 29.—Diagram of the arterial system of A, Scorpio, and B, Limulus. The Roman numerals indicate the body somites and the two figures are adjusted for comparison. ce, Cerebral arteries; sp, supra-spinal or medullary artery; c, caudal artery; l, lateral anastomotic artery of Limulus. The figure B also shows the peculiar neural investiture formed by the cerebral arteries in Limulus and the derivation from this of the arteries to the limbs, III, IV, VI, whereas in Scorpio the latter have a separate origin from the anterior aorta. (From Lankester, “Limulus an Arachnid.”) |
It must be pointed out that the presence or absence of such renal excretory tubes opening into the intestine appears to be a question of adaptation to the changed physiological conditions of respiration, and not of morphological significance, since a pair of renal excretory tubes of this nature is found in certain Amphipod Crustacea (Talorchestia, &c.) which have abandoned a purely aquatic life. This view has been accepted and supported by Professors Korschelt and Heider (16). An important fact in its favour was discovered by Laurie (17), who investigated the embryology of two species of Scorpio under Lankester’s direction. It appears that the Malpighian tubes of Scorpio are developed from the mesenteron, viz. that portion of the gut which is formed by the hypoblast, whereas in Hexapod insects the similar caecal tubes are developed from the proctodaeum or in-pushed portion of the gut which is formed from epiblast. In fact it is not possible to maintain that the renal excretory tubes of the gut are of one common origin in the Arthropoda. They have appeared independently in connexion with a change in the excretion of nitrogenous waste in Arachnids, Crustacea, and the other classes of Arthropoda when aerial, as opposed to aquatic, respiration has been established—and they have been formed in some cases from the mesenteron, in other cases from the proctodaeum. Their appearance in the air-breathing Arachnids does not separate those forms from the water-breathing Arachnids which are devoid of them, any more than does their appearance in certain Amphipoda separate those Crustaceans from the other members of the class.
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| Fig. 30.—View from below of a scorpion (Buthus occitanus) opened and dissected so as to show the pericardium with its muscles, the lateral arteries, and the tergo-sternal muscles. PRO, Prosoma. tsm1, Tergo-sternal muscle (labelled dv in fig. 31) of the second (pectiniferous) mesosomatic somite; this is the most anterior pair of the series of six, none are present in the genital somite.
tsm4, Tergo-sternal muscle of the fifth mesosomatic somite.
tsm6, Tergo-sternal muscle of the enlarged first metasomatic somite.
Per, Pericardium. VPM1 to VPM7, The series of seven pairs of veno-pericardiac muscles (labelled pv in fig. 31). There is some reason to admit the existence of another more anterior pair of these muscles in Scorpio; this would make the number exactly correspond with the number in Limulus. (After Lankester, Trans. Zool. Soc. vol. xi, 1883.) |
Further, it is pointed out by Korschelt and Heider that the hinder portion of the gut frequently acts in Arthropoda as an organ of nitrogenous excretion in the absence of any special excretory tubules, and that the production of such caeca from its surface in separate lines of descent does not involve any elaborate or unlikely process of growth. In other words, the Malpighian tubes of the terrestrial Arachnida are homoplastic with those of Hexapoda and Myriapoda, and not homogenetic with them. We are compelled to take a similar view of the agreement between the tracheal air-tubes of Arachnida and other tracheate Arthropods. They are homoplasts (see 18) one of another, and do not owe their existence in the various classes compared to a common inheritance of an ancestral tracheal system.
Conclusions arising from the Close Affinity of Limulus and Scorpio.—When we consider the relationships of the various classes of Arthropoda, having accepted and established the fact of the close genetic affinity of Limulus and Scorpio, we are led to important conclusions. In such a consideration we have to make use not only of the fact just mentioned, but of three important generalizations which serve as it were as implements for the proper estimation of the relationships of any series of organic forms. First of all there is the generalization that the relationships of the various forms of animals (or of plants) to one another is that of the ultimate twigs of a much-branching genealogical tree. Secondly, identity of structure in two organisms does not necessarily indicate that the identical structure has been inherited from an ancestor common to the two organisms compared (homogeny), but may be due to independent development of a like structure in two different lines of descent (homoplasy). Thirdly, those members of a group which, whilst exhibiting undoubted structural characters indicative of their proper assignment to that group, yet are simpler than and inferior in elaboration of their organization to other members of the group, are not necessarily representatives of the earlier and primitive phases in the development of the group—but are very often examples of retrogressive change or degeneration. The second and third implements of analysis above cited are of the nature of cautions or checks. Agreements are not necessarily due to common inheritance; simplicity is not necessarily primitive and ancestral.
On the other hand, we must not rashly set down agreements as due to “homoplasy” or “convergence of development” if we find two or three or more concurrent agreements. The probability is against agreement being due to homoplasy when the agreement involves a number of really separate (not correlated) coincidences. Whilst the chances are in favour of some one homoplastic coincidence or structural agreement occurring between some member or other of a large group a and some member or other of a large group b, the matter is very different
