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<Br>On the Presence of Light Organs In BIOLOGICAL RESULTS OF THE UNIVERSITY OF MIAMI DEEP-SEA EXPEDITIONS. 54. ON THE PRESENCE OF LIGHT ORGANS IN SEMIROSSIA STEENSTRUP, 1887 (MOLLUSCA: CEPHALOPODA)l S. V. BOLETZKY2 University OF Miami, Rosenstiel School OF Marine AND Atmospheric Sciences ABSTRACT The presence of light organs in the species Rossia TENERA (Verrill), R. EQUALIS Voss, and R. PATAGONICA Smith (R. TENERA subsp. PATAGONICA [Smith)), which together form the subgenus Semirossia Steenstrup, is stated in the present study. The organs are briefly described; their function as light orgall.s is mainly deduced from the anatomical similarity to those of other sepiolids, as observations on living animals are still lacking. INTRODUCTION Although the presence of light organs in the mantle cavity of some species of Rossia has been mentioned and discussed in the past, the lack of such or- gans is generally regarded as a character of the monotypic subfamily Ros- siinae, in contrast to the other two subfamilies of the Sepiolidae (Sepiolinae and Heteroteuthinae) which comprise numerous species equipped with light organs. During a cruise to the Caribbean aboard the R/V JOHN ELLIOTT PILLS- BURY of the University of Miami, in March 1968, I found light organs connected with the ink sac in specimens of Rossia (Semirossia) TENERA (Verrill) trawled off Yucatan. Further investigations in the collections of the University of Miami Ros- enstiel School of Marine and Atmospheric Sciences and of the U. S. National Museum showed that light organs are always present in R. TENERA (Verrill) and R. EQUALIS Voss, as well as in R. PATAGONICA Smith, which was referred to by Thore (1959) as Semirossia TENERA PATAGONICA (in the following, how- ever, this species is referred to under its old specific name, because of a dif- ference in the configuration of the suckers of the hectocotylus, ignored by Thore, that first has to be cleared up). The presence of light organs in R. PATAGONICA has already been observed by R. E. Young and E. S. McSweeny, Jr., of the Rosenstiel School of Marine and Atmospheric Sciences. They kindly gave the abundant material to me for further examination. 1 Contribution No. 1190 from the University of Miami, Rosenstiel School of Marine and Atmo- spheric Sciences. This study was supported by National Science Foundation grant G B 5729X. This paper is one of a series resulting from the National Geographic Society-University of Miami Deep- Sea Biology Program. 2 Present address: Laboratoire Arago, 66 Banyuls-sur-Mer, France. 1970] v. Boletzky: Light Organs IN Semirossia 375 ACKNOWLEDGMENTS I wish to give my sincere thanks to Dr. Gilbert L. Voss, Chairman of the Division of Biological Sciences of the Rosenstiel School of Marine and At- mospheric Sciences, as well as to Dr. Richard E. Young and Mr. Edward S. McSweeny, Jr., and to Dr. Clyde F. E. Roper of the U. S. National Museum for valuable advice and help in carrying out this study. I particularly ac- knowledge the assistance of my wife who performed all histological prepara- tions. During my work at the Rosenstiel School, I was the recipient of a post- doctoral fellowship of the Swiss National Fund for the Advancement of Sci- entific Research. All laboratory expenses were covered by National Science Foundation grant GB 5729X. HISTORICAL RESUME Meyer (1906), in a note on the light organ of the Sepiolini, mentioned an inconspicuous swelling on the right side of the anus in Rossia MACROSOMA which, he supposed, was a reduced light organ. Chun (1915) and Sasaki (1920) described papilliform organs on either side of the rectum in R. MASTIGOPHORA and R. bipapillata, respectively. Naef (1923: 575-76), in his monograph, mentioned both records and suggested that these organs might be rudimentary light organs, and that the second pair of papillae in the males of R. mastigophora, as described by Chun, might be rudimentary accessory nidamental glands. However, when characterizing the subfamily Rossiinae (Naef, 1912), he stated (1923: 573) that no light organs are present in the mantle cavity, but then (p. 574) he again described the differentiation of the front part of the accessory gland as a luminescent gland. These contradictory statements were evidently due to the uncertainty of whether or not the described organs really were light organs. Nevertheless, Naef appeared to have had a very precise idea about a conceivable pathway of formation or reduction of light organs in Rossia; he evidently supposed the presence of distinct light organs in Semirossia. In a free translation, his hypothesis (pp. 575-76) reads: "1 assume that Rossiinae with well-devel- oped light organs lived in greater depths. '" R. MASTIGOPHORA then might be an intermediate form. Unfortunately, the entire subfamily is quite insuf- ficiently known. The situation in Semirossia, for instance, would be interest- ing as it is in some regards strikingly close to the Sepiolinae." As to the functioning of the light organs in Sepiolids, Meyer (1906) be- lieved that the organ was a gland producing a luminous secretion. Skowron (1926) was able to show that the luminescence was due to luminous bacteria dwelling in the organ. The function, anatomy, histology, and bac- teriology of these light organs are treated in Harvey's "Bioluminescence" 376 Bulletin OF Marine Science [20(2) (1952), based on numerous original investigations, the references to which are given there. MATERIAL AND METHODS The material examined included 36 specimens of R. TENERA (Verrill) (14 <t, 22 '?) from depths mainly between l60 and 350 m; 8 specimens of R. EQUALIS Voss (7 <t, 1 '?) from depths of about 450 m; 3 specimens of Semirossia (1 <t, 2 '?) which could not definitely be identified as TENERA or equalis; and 37 specimens of R. PATAGONICA Smith (8 0, 29 '?) from South Atlantic stations southwest of the Falkland Islands at depths of 120-230 m. Most of the specimens of R. TENERA and R. EQUATIS were collected by the research vessels ATLANTIS, GERDA, OREGON, and JOHN ELLIOTT PILLSBURY (Figs. lO, 11). All of them had distinct light organs more or less involved in the anterior part of the ink sac. No light organs associated with the ink sac have been found in specimens of Rossia pacifica, R. molleri, R. megaptera, R. mollicella, R. sublevis, R. glaucopis, R. caroti, R. macrosoma, R. tortugaensis, R. bullisi, and R. anti/- lensis. The male specimens of R. BULLISI had, in addition to the kidney papillae, two small bodies on the front of the kidney sac. One male specimen of R. CAROLI had no ink sac at all. Nine light organs and six accessory nidamental glands of the three species of Semirossia have been investigated histologically. The material was em- bedded in Paraplast, sectioned at 6-fL thickness, and stained with hematoxy- lin-eosine or Masson's Trichrome. ABBREVIATIONS USED IN ANATOMICAL DRAWINGS aa, anal artery la, artery to light organ ang, accessory nidamental gland ld, duct of light organ ct, connective tissue 10, light organ h, systemic heart lp, papilla of light organ la, artery of ink gland lv, vein from light organ id, ink duct rna, mantle artery ig, ink gland r, "reflector" kp, kidney papilla vc, vena cava I, "lens" GENERAL MORPHOLOGYOF THE LIGHT ORGAN IN Semirossia The light organ of Semirossia is a bilobed body more or less surrounded by the anterior part of the ink sac; its position is transverse to the longitudi- nal axis of the animal, ventral to the ink duct (Fig. 1). In the four fresh specimens of Rossia (Semirossia) lenera, in which the organ was first observed by the author, its free part could be seen as a very light, whitish, transverse band on the ink sac (Fig. 1) when the mantle 1970] v. Boletzky: Light Organs IN Semirossia 377 FIGURE 1. Fresh dissection of mantle cavity of female of Rossia tenera, ML 27.5 mm. Free part of light organ is visible on top of the ink sac. was slightly pulled back. After preservation, this striking aspect of the organ disappears as the transparent integument and "lens" tissue turn opaque; this might be the reason the organ has gone unnoticed in the past. In several small male specimens of R. tenera, the organ strongly protrudes into the mantle cavity on either side of the hindgut, and shows two bulbs 378 Bulletin OJ Marine Science [20(2) 1970J v. Boletzky: Light Organs IN Semirossia 379 •... 'u; -< ,5 Q) ~ '0.. c...o:! Q) -5..... o:!t: ..co:!•..• 0., Q) Q) •..•Q) 0 ••• Z<+-< o:! E~ E.e V)U •....• ~ '0 ...:IQ) ::E-5 Q)"!XI (;1:: Ei';;; , o:! <::l•.. •..Q) <:u Q) s::.e ~ ~ .d' ~o:! bJ) " •.. iXlO..•... E~ E:.:: v:] 0\ •..• •.••.• <+-< 0 ...:I", ~:9;::l .,)'.0 -Q) Eo:!.e•... <+-< <::l"0 •.. 0., <:u 0 s::•..• ~ I:: <::l 0 ';;; 0'" R:: -<" ..; ~ ~ ;::l ~" 380 Bulletin OF Marine Science [20(2) • A Q '1 C o FIGURE4. Different dispositions of the light organ on the ink sac, and variations in the type of duct, following the definitions given in the text: A, class 1 disposi- tion of the light organ, and ducts of type a (absence of papillae); B, class 2 dis- position, and ducts of type b (with papillae); C, class 1 disposition and type b; D, class 3 disposition and type b. (A and B are for females; C and D are for males. The symbols beside the drawings are also used in Figures 9 and 11). which can hardly be overlooked (Fig. 3, A). Even when not so evident, the organ often can be easily recognized by its appearance as an opaque struc- ture that hides the anterior part of the ink sac, uncovered by the renal ap- pendages of the kidney (Fig. 2). In large female specimens however, it often cannot be seen when the accessory nidamental glands and the nidamental glands extend far anteriorly and overlap the entire ink sac.
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