THE RELATIONSHIP OF THE CHELICERATE ARTERIAL SYSTEM ...

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50 THE JOURNAL OF ARACHNOLOGY4. In all chelicerates which lack book lungs there exists a perivisceral arterial sinus.This sinus persists in its most primitive state in pycnogonids. In Limulus and theapulmonate arachnids, it persists principally in the form of a periganglionic arterialsinus.5. The thoracic sinus arterial System of pulmonate arachnids in an arrested stage ofarterial development and is interpreted here äs neoteny. This stage occurs in thelarval Limulus, and presumably it occurs also during the embryogeny of otherchelicerates.6. In the apulmonate arachnids, the development of the mesodermal endosternite isinversely proportional to the development of the apodemal endoskeleton. Hence,the more highly developed the apodemal endoskeleton, the more vestigial is theendosternite.7. The pycnogonid horizontal vascular septum is probably homologous to the ancestralprogenitor of the endosternite in merostomes and arachnids.8. The primitive function of the endosternite may have been vascular. Movements ofthe endosternite may have augmented cardiac contractions in effecting blood circulationthrough the perivisceral arterial sinus and into the appendicular arteries.9. Probably, neoteny has been involved in the origin of arachnids from merostomes,and in the origin of apulmonate arachnids from pulmonates.10. The Pulmonata is a natural monophyletic category. It includes the Scorpionida,Thelyphonida, Schizomida, Amblypygida and Araneida.11. The Apulmonata is a natural monophyletic category. It includes the Palpigradida,Acarida, Opilionida, Ricinuleida, Pseudoscorpionida and Solpugida. The ancestralapulmonate stock diverged to give rise both to modern palpigrades and to thenonpalpigrade apulmonates.ACKNOWLEDGEMENTSThis research is dedicated to the memory of Professor Gordon F. Ferris, underwhose influence it was inspired. Professor Ferris believed, äs I do, that comparativeinternal morphology is the most fruitful approach to deducing the complicatedphylogeny of the Arthropoda. I wish to acknowledge also my sincere gratitude to Dr. D.P. Abbott, whose very helpful counsel filled in the gap left by the passing of ProfessorFerris. Moreover, I am grateful to Dr. J. W. Hedgpeth, who supplied me not only withlarge specimens of pycnogonids, but also with the counsel and constructive criticism thatI needed in order to develop what I feel is the most important contribution of thisresearch: namely, the homologies of circulatory morphology common to the Pycnogonidaand the rest of the Chelicerata. I am thankful to the following colleagues forProfessional advice, technical assistance, and specimens: Dr. J. F. Oliphant, Dr. C. J.Goodnight, Dr. I. M. Newell, Dr. E. I. Schlinger, Dr. E. T. Röche, Dr. B. J. Kaston, Dr. W.J. Gertsch, Dr. R. W. Mitchell, Mr. J. M. Rowland, Mr. V. D. Roth, and Miss M. J. Moody.Finally, I am thankful to the librarians, Florence Chu (Stanford), and Mrs. S. M. Walton(Riverside), without whose cooperation I could not have collected the literature necessaryfor this research.LITERATURE CITEDAndre, M. 1949. Order des Acariens, p. 794-892. In P. P. Grasse [ed.], Traite de Zoologie, tome6. Masson et Cie, Paris.
FIRSTMAN-CHELICERATE ARTERIAL SYSTEM AND ENDOSTERNITE 51Appell, V. 1900. Studie o nervove soustave Phalangiinu. Sitzungsberichte Königlich Böhmischen Ges.der Wiss. 38 p.Baker, E. W. and G. W. Wharton. 1952. An introduction to acarology. Macmillan, New York. 465 p.Barnes, R. D. 1968. Invertebrate zoology. 2nd ed. Saunders, Philadelphia. 743 p.Beck, E. J. 1885. Description of the muscular and endoskeletal Systems of Scorpio. Trans. Zool. Soc.London 11:339-360.Beklemischev, V. N. 1958. Grundlagen der vergleichenden Anatomie der Wkbellosen, bd.1-2. Deutscher Verlag Wiss. Berlin.Benham, W. B. S. 1885. Description of the muscular and endoskeletal Systems of Limulus. Trans.Zool. Soc. London 11:314-338.Berland, L. 1949. Ordre des Opiliones, p. 761-793. In P. P. Grasse [ed.], Traite de Zoologie, tome6. Massen et Cie, Paris.Bernard, H. M. 1892a. The Apodidae. Nature Series. Macmillan, London.Bernard, H. M. 1892b. The apodemes of Apus and the endophragmal System ofAstacus. Ann. Mag.Nat. Hist. 10:67-74.Bernard, H. M. 1894a. Some observations on the relations of the Acaridae to the Arachnida. J. Linn.Soc. London (Zool.) 24:279-291.Bernard, H. M. 1894 . Notes on the Chernetidae, with special reference to the vestigial Stigmata andto a new form of trachea. J. Linn. Soc. London (Zool.) 24:410-430.Bernard, H. M. 1894c. The endosternite of Scorpio compared with the homologous structures inother Arachnida. Ann. Mag. Nat. Hist. 13:18-26.Bernard, H. M. 1896. The comparative morphology of the Galeodidae. Trans. Linn. Soc. (Zool.), Ser.2,6:305-417.Blanchard, E. 1852-60. L'Organization du regne animal. Victor Masson, Paris.Bonnet, A. 1907. Recherches sur l'anatomie comparee et le developpement des Ixodides. Ann. Univ.Lyon, New Ser., 1:1-185.Börner, C. 1901. Zur ausseren Morphologie von Koenenia mirabilis Grassi. Zool. Anz. 24:537-556.Börner, C. 1904. Beiträge zur Morphologie der Arthropoden. I. Ein Beitrag zur Kenntnis derPedipalpen. Zoologica 42:1-174.Borradaile, L. A., F. A. Potts, L. E. S. Eastham, and J. T. Saunders. 1961. (4th ed., revised by G. A.Kerkut). The invertebrata. Cambridge Univ. Press, London. 820 p.Buisson, M. du 1925. Recherches sur la circulation sanguine et la Ventilation pulmonaire chez lesscorpions. Bull. Acad. Roy. Belgique, Brussells 11(5):666-680.Causard, M. 1892. Sur la circulation du sang chez les jeunes araignees. Compt. Rend., Acad. Sei.,Paris 114:1035-1038.Causard, M. 1893. Sur l'appareil circulatore de la Mygale caementaria Walk. Compt. Rend., Acad.Sei., Paris 116:828-830.Causard, M. 1896. Recherches sur l'appareil circulatore des Araneides. Bull. Sei. France et Belgique29:1-109.Cole, L. J. 1910. Peculiar habit of a pycnogonid new to North America with observations on the heartand circulation. Biol. Bull. 18:193-203.Comstock, J. H. 1948(2nd. ed., revised by W. J. Gertsch). The spider book. Comstock, Ithaca, NewYork, 729p.Croneberg, A. 1888. Beitrag zur Kenntnis des Baues der Pseudoscorpione. Bull. Soc. Imper. Nat.Moscou, New Ser., 2:416-461.Daday, E. von. 1880. Ueber den Circulationsapparat der Pseudoscorpione. Termeszetrajzi Füzetek4:331-339.Dohrn, A. 1870. Ueber Entwicklung und Bau der Pycnogoniden. Jenaische Z. Med. Naturwiss.5:138-157.Dohrn, A. 1871. Utersuchungen über Bau und Entwickelung der Arthropoden. 12 Zur Embryologieund Morphologie des Limuluspolyphemus. Jenaishe Z. Med. Nat. 6:580-640.Dohrn, A. 1879. Neue Utersuchungen über Pycnogoniden. Mittheil. Zool. Sta. Neapel 1:28-39.Dohrn, A. 1881. Die Pantopoden des Golfes von Neapel und der angrenzenden Meeresabschnitte.Fauna Flora Golfes Neapel 3:1-252.Dufour, L. 1862. Anatomie, physiologie et histoire naturelle des Galeodes. Mem., Acad. Sei. Paris17:338-446.Evans, G. O., J. G. Sheals, and D. Macfarlane. 1961. The terrestrial Acari of the British Isles, vol 1.British Mus., London. 219 p.
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