retained as a rudimentary, separate, detached somite in front of the mesosoma, or disappears altogether (excalation). The atrophy and total disappearance of ancestrally well-marked somites frequently take place (as in all Arthropoda) at the posterior extremity of the body, whilst excalation of somites may occur at the constricted areas which often separate adjacent “regions,” though there are very few instances in which it has been recognized. Concentration of the organ-systems by fusion of neighbouring regions (prosoma, mesosoma, metasoma), previously distinct, has frequently occurred, together with obliteration of the muscular and chitinous structures indicative of distinct somites. This concentration and obliteration of somites, often accompanied by dislocation of important segmental structures (such as appendages and nerve-ganglia), may lead to highly developed specialization (individuation, H. Spencer), as in the Araneae and Opiliones, and, on the other hand, may terminate in simplification and degeneration, as in the Acari.}
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Fig. 40.—Four stages in the development of the trilobite Agnostus nudus. A, Youngest stage with no mesosomatic somites; B and C, stages with two mesosomatic somites between the prosomatic and telsonic carapaces; D, adult condition, still with only two free mesosomatic somites. (From Korschelt and Heider.) |
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From Korschelt and Heider, after Barrande. Fig. 41.—Five Stages in the development of the trilobite Sao hirsuta. C, Stage with two free mesosomatic somites between the prosomatic and telsonic carapaces.
D, Stage with seven free intermediate somites. E, Stage with twelve free somites; the telsonic carapace has not increased in size.
a, Lateral eye.
g, So-called facial “suture” (not really a suture). |
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Fig. 42.—So-called “trilobite stage” of Limulus polyphemus. A, Dorsal; B, ventral view. (From Korschelt and Heider, after Leuckart.) |
The most important general change which has affected the structure of the nomomeristic Arachnida in the course of their historic development is the transition from an aquatic to a terrestrial life. This has been accompanied by the conversion of the lamelliform gill-plates into lamelliform lung-plates, and later the development from the lung-chambers, and at independent sites, of tracheae or air-tubes (by adaptation of the vasifactive tissue of the blood-vessels) similar to those independently developed in Peripatus, Diplopoda, Hexapoda and Chilopoda. Probably tracheae have developed independently by the same process in several groups of tracheate Arachnids. The nomomeristic Arachnids comprise two sub-classes—one a very small degenerate offshoot from early ancestors; the other, the great bulk of the class.
Sub-Class I. (of the Nomomeristica). PANTOPODA.—Nomomeristic Arachnids, in which the somites corresponding to mesosoma and metasoma have entirely aborted. The seventh, and sometimes the eighth, leg-bearing somite is present and has its leg-like appendages fully developed. Monomeniscous eyes with a double (really triple) cell-layer formed by invagination, as in the Eu-arachnida, are present The Pantopoda stand in the same relation to Limulus and Scorpio that Cyamus holds to the thoracostracous Crustacea. The reduction of the organism to seven leg-bearing somites, of which the first pair, as in so many Eu-arachnida, are chelate, is a form of degeneration connected with a peculiar quasi-parasitic habit resembling that of the crustacean Laemodipoda. The genital pores are situate at the base of the 7th pair of limbs, and may be repeated on the 4th, 5th, and 6th. In all known Pantopoda the size of the body is quite minute as compared with that of the limbs: the alimentary canal sends a long caecum into each leg (cf. the Araneae) and the genital products are developed in gonocoels also placed in the legs.
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From Parker and Harwell’s Text-book of Zoology, after Hoek. Fig. 43.—One of the Nymphonomorphous Pantopoda, Nymphon hispidum, showing the seven pairs of appendages 1 to 7; ab, the rudimentary opisthosoma; s, the mouth-bearing proboscis. |
The Pantopoda are divided into three orders, the characters of which are dependent on variation in the presence of the full number of legs.
Order 1. (of the Pantopoda). Nymphonomorpha, Pocock (nov.) (fig. 43).—In primitive forms belonging to the family Nymphonidae the full complement of appendages is retained—the 1st (mandibular), the 2nd (palpiform), and the 3rd (ovigerous) pairs being well developed in both sexes. In certain derivative forms constituting the family Pallenidae, however, the appendages of the 2nd pair are either rudimentary or atrophied altogether.
Two families: 1. Nymphonidae (genus Nymphon), and 2. Pallenidae (genus Pallene).
Order 2. Ascorhynchomorpha, Pocock (nov.).—Appendages of the 2nd and 3rd pairs retained and developed, as in the more primitive types of Nymphonomorpha; but those of the 1st pair are either rudimentary, as in the Ascorhynchidae, or atrophied, as in the Colossendeidae. In the latter a further specialization is shown in the fusion of the body segments.
Two families. 1. Ascorhynchidae (genera Ascorhynchus and Ammothea); 2. Colossendeidae (genera Colossendeis and Discoarachne).
Order 3. Pycnogonomorpha, Pocock (nov.).—Derivative forms in which the reduction in number of the anterior appendages is carried farther than in the other orders, reaching its extreme in the Pycnogonidae, where the 1st and 2nd pairs are absent in both sexes, and the 3rd pair also are absent in the female. In the Hannoniidae, however, which resemble the Pycnogonidae in the absence of the 3rd pair in the female and of the 2nd pair in both sexes, the 1st pair are retained in both sexes.
Two families: 1. Hannoniidae (genus Hannonia); 2. Pycnogonidae (genera Pycnogonum and Phoxichilus).
Remarks.—The Pantopoda are not known in the fossil condition. They are entirely marine, and are not uncommon in the coralline zone of the sea-coast. The species are few, not more than fifty (23). Some large species of peculiar genera are taken at great depths. Their movements are extremely sluggish. They are especially remarkable for the small size of the body and the extension of viscera into the legs. Their structure is eminently that of degenerate forms. Many frequent growths of coralline Algae and hydroid polyps, upon the juices of which they feed, and in some cases a species of gall is produced in hydroids by the penetration of the larval Pantopod into the tissues of the polyp.
Sub-Class II. (of the Nomomeristic Arachnida). EU-ARACHNIDA.—These start from highly developed and specialized aquatic branchiferous forms, exhibiting a prosoma with six pediform pairs of appendages, an intermediate prae-genital somite, a mesosoma of six somites bearing lamelliform pairs of appendages, and a metasoma of six somites devoid of appendages, and the last provided with a post-anal spine. Median eyes are present, which are monomeniscous, with distinct retinal and corneagenous cell-layers, and placed centrally on the prosoma. Lateral eyes also may be present, arranged in lateral groups, and having a single or double cell-layer beneath the lens. The first pair of limbs is often chelate or prehensile, rarely antenniform; whilst the second, third and fourth may also be chelate, or may be simple palps or walking legs.
