A New Patagonian Cephalopod (Octopodidae) with Notes on Related Genera

BULLETIN OF MARINE SCIENCE, 28(2): 282-296, 1978

VOSSELEDONE CHARRUA: A NEW PATAGONIAN CEPHALOPOD (OCTOPODIDAE) WITH NOTES ON RELATED GENERA

Francisco J. Palacio

ABSTRACT A new octopodan genus, Vosseledone, and its type species, Vosseledone charrua, is described from the Patagonian Province. The new taxon is characterized by the possession of a degenerate radula and a well developed ink sac. Its position in reference to the closely related genera is reviewed.

In the course of a study of the coastal MORG, Museu Oceanografico de Rio Cephalopoda of Brasil, I encountered seven Grande do SuI, Brazil; BMNH, British specimens which represent a new genus of Museum of Natural History; UMML, Uni- incirrate octopods with uniserial suckers versity of Miami, Marine Laboratory. that occurs in the southwest Atlantic Ocean. I have examined three specimens from a col- Family OCTOPODIDAE Orbigny lection kindly loaned by E. C. Rios of the Subfamily ELEDONINAE Gray Museu Oceanografico de Rio Grande, Rio Vosseledone new genus Grande do SuI, Brazil. Four additional Diagnosis.-Octopod with uniserial suckers. specimens had been sent to G. L. Voss, Ink sac large superficially imbedded, evident University of Miami, by Z. de Castellanos externally. Radula degenerate with tall, re- of the Instituto de Biologia Marina, Mar curved, simple rachidian tooth with narrow del Plata, Argentina, originally believed to base; faint traces of first laterals; second represent Tetracheledone spinicirrus from laterals degenerate with proximal cuspless the coast of Uruguay. A preliminary study prominence; third laterals and marginal suggested the possibility that the specimens plates lacking. Third right arm hectocoty- belonged to the genus Thaumeledone Iized with strong, well differentiated ligula; Robson, 1930. The type specimens of calamus deeply grooved. Arm heteromor- Thaumeledone brevis and T. gunteri were phism absent. Digestive system with nor- made available for study by the courtesy of mal esophageal crop. Funnel organ V V J. F. Peake of the British Museum of Nat- type. Large eggs and spermatophores. ural History. Measurements and indices given in this Type species.-Vosseledone charrua n. sp. paper have been defined by Voss (1963) with the exception of Hectocotylized Arm Vosseledone charrlLa new species Index (HeAL) used here as the mantle Figures 1-5; Tables 1-6 length expressed as percentage of the Charruan octopus. Pulpo charrua. Polvo hectocotylized arm length (MHI, of Pick- charrua. Tetracheledone spillicirrus, Caste- ford, 1964); and Fellow Arm Index (FAI), llanos y Menni, 1968: 18; 1969: 73, 77 (non defined here as the hectocotylized arm Tetracheledone spinicirrus Voss, 1955). length expressed as a percentage of the Material examined.-HOLOTYPE, one male. symmetrical fellow arm length (HAl, of mantle length 49 mm. 35°14'S, 52°28'W, east of Punta del Este, Uruguay, in 200 m, USNM Pickford, 1964). Scanning electron micro- 729454. PARATYPES: 2 males, mantle lengths scope photographs were taken by W. 50 and 53 mm, and one gravid female, mantle length 61 mm, collected with the holotype, Charm, RSMAS. Abbreviations used are: UMML 1681. One male, mantle length 50 mm, USNM, United States National Museum of and one gravid female, mantle length 58 mm, 33° 17'S, 52°40'W, off Albardfto, Rio Grande do Natural History, Smithsonian Institution; SuI, in 100 m; collected by ALINE, October 282 PALACIO: A NEW PATAGONIAN CEPHALOPOD 283

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Figure 1. Vosse/edolle e1wrrua, new genus and species: a, dorsal view of male; b, funnel organ; c, digestive system; d, upper mandible; e, lower mandible, 284 BULLETIN OF MARINE SCIENCE, VOL. 28, NO.2, 1978

Figure 3. Vosseledolle clwrrua, new genus and species. Radula.

erect and strong; they are largest near the Figure 2. Vosseledone charrua, new genus and edge of the web and minute distally. Almost species. Male hectocotylus. half the suckers are found on the distal one fourth of the arm. The web is moderately 1970, USNM 729455. One female, mantle length deep with an average index of 34. Sector C 76.5 mm, from Sao Sebastiao, Sao Paulo, in 10 m, is usually deepest; sector E is usually the collected by J. Ribeiro, 20 December 1945, shallowest. MORG 10701- The third right arm is hectocotylized; it is Description.-Vosseledone charrua is a only slightly shorter than the corresponding small octopod. The mantle is globular and left arm. The hectocotylus is muscular and wide, its length about one third of the total massive with a ligula index of 8.4-11.1. The length. The mantle width is accentuated in ligula is deeply excavated with few gravid females. The head is not as wide as laminae; the lateral margins are strong and the mantle and the neck is not constricted. somewhat rolled inward. The calamus (Fig. The eyes are prominent and occupy a large 2) is large (CLI 43.1-58.8) and strong, portion of the head area. The pallial aper- deeply grooved, and very acute apically. The ture is wide. spermatophoral groove is prominent with a The funnel is stout and muscular, free wide, strongly infolded membrane. for about half of its length. The funnel There are 6-7 gill filaments per outer organ consists of two thick V-shaped pads demibranch. The digestive system of one which are acute apically. The basal funnel male and one female were dissected. The flaps are strong and with the corresponding beaks (Fig. Id-e) offer no special charac- depressions of the mantle constitute a ters. The radula is degenerate (Figs. 3-5), powerful mantle-locking apparatus. possessing a tall recurved and simple The arms are strong, about twice as long rachidian tooth with a narrow base. There as the mantle, with an ALI of 58-67 and are faint traces of first laterals. The second an AWl of about 21. The arms are nearly laterals are degenerate and exhibit a proxi- equal in length and show no particular arm mal prominence without a cusp. The third order. The suckers are uniserial, large, laterals and marginal plates are lacking. PALACIO: A NEW PATAGONIAN CEPHALOPOD 285

Figure 5. Vosseledolle c1wrrua, new genus and species. Lateral view of rachidian tooth. Figure 4. Vosseledolle charrua, new genus and species. Delail of rachidian tooth and atrophied laterals. ulum. Needham's sac contains a few (three in specimen No.2) large spermatophores (Fig. 6a). These are long (SpLI 100-112) The digestive tract is of the normal and slender (SpWI 5-8); the convoluted octopodan type. Anterior and posterior tube of the sperm receptacle occupies 36- salivary glands are present. The esophagus 42% of the tube length; the cylinder and is anteriorly expanded into a crop. The cement body, about 20% and the ejaculatory stomach is smalIer than the spiral caecum. apparatus is almost as long as the receptacle. The surface of the latter has small protrud- The anterior third of the ejaculatory tube is ing nodules which are probably the result arranged as a spiral band, the midportion of undetermined parasitic or bacterial in- resembles flakes disposed along a main axis fections. The caecum is followed by a large and the oral segment is slightly spiraled. intestine which is recurved midway to its The ovary is large with stout proximal junction with the ink sac orifice in the anal oviducts. The ovary (Fig. 6c), of a gravid region. The liver is large and ovoid; the female (specimen No.1), contained about pancreatic region is connected with the 40 differentially developed, closely clustered, spiral caecum by paired hepatopancreatic 10ngitudinalIy striated eggs (Fig. 6d). The ducts. The ink sac is large and fusiform, apparently mature eggs are about 13 mm about half as long as the liver into which it by 4 mm with prominent stalks. is superficially imbedded but externally evi- The sculpture, in preservation, of the dent when the viscera are exposed. dorsal surface of the mantle, head and Male and female reproductive systems arms, consists of numerous well-spaced, were dissected. The penis is thick and small papillose warts; some of these, irregu- tubular with a proximal, recurved divertic- larly distributed, are more minutely papil- 286 BULLETIN OF MARINE SCIENCE, VOL. 28, NO.2, 1978

Figure 6. Vosseledone clwrrua, new genus and species: a, spermatophore; b, penis; c, female geni- talia; d, egg.

lated than the others. The warts are less octopods devoid of an ink sac and in which pronounced on the distal portion of the the esophageal crop is reduced or absent. arms. Two prominent papillose cirri occur This rules out the possibility of Vosseledone over the eye, of which the anterior is being congeneric with Bathypolypus Grimpe, smaller. The eyelids are densely papillated. 1921, Graneledone Joubin, 1918, Thau- The ventral surface of the mantle, head and meledone Robson, 1930, and Bentheledone arms is smooth. Robson, 1932. In addition, the degenerate In preservation in isopropyl alcohol, the radula of Vosseledone is distinctly different animals are of light flesh color. from that of BathypolYPllS, Graneledone, Measurements and indices for the seven and Bentheledone, where the rachidian specimens are given in Tables 1 and 2. tooth is simple and the lateral teeth are well differentiated. The degenerate radula of Type.-USNM 729454, male, mantle length Thaumeledone brevis (known in the litera- 49.0 mm. ture from two small CHALLENGER speci- Type locality.-35 °14'S, 52°28'W, east of mens from off Montevideo), and T. gunteri Punta del Este, Uruguay, in 200 m. (known from one small DISCOVERY female collected in the South Georgia Islands), Discussion.-The new taxon described suggested the possibility that the material above cannot be placed in any of the under study might be conspecific with known genera of the Octopodinae, Bathy- either of these species, more likely T. brevis, polypodinae, and Eledoninae recognized by due to its geographical distribution. The Robson (1932). The Bathypolypodinae possibility arose that because of the small was considered by Robson to include abyssal size of the type specimens of Thaumeledone, PALACIO: A NEW PATAGONIAN CEPHALOPOD 287

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Table 2. Indices of bodily proportions of 4 males and 3 females of Vosseledolle chamw new species ======--======-.c==--=-- -- - Sao Sebastiao Rio Grande do SuI Range Index Brazil Brazil Uruguay x

ML 76.5 N = 1 50.0- 58.0 N = 2 49.0- 61.0 49.0-56.8- 76.5 MWI 91.5 103.4-108.0 86.9-102.0 86.9-98.6-108.0 HWI 64.1 75.8- 88.0 57.4- 82.0 57.4-74.1- 88.0 ALI 58.3 62.4- 63.0 61.1- 67.5 58.3-63.0- 67.5 AWl 17.6 20.0- 20.6 18.8- 24.4 17.6-20.9- 24.4 MAl 52.6 47.2- 48.1 39.3- 51.7 39.3-47.7- 52.6 HcAI 64.9 54.1- 69.4 57.0-61.4- 69.4 FAI 98.7 73.1- 86.0 73.1-85.4- 98.7 LLI 8.4 9.9- 11.1 8.4- 9.9- ] 1.1 eLI 43.1 50.0- 58.8 43.1-50.7- 58.8 PLI 36.0 26.0- 31.1 26.0-30.4- 36.0 SoDI 6.5 10.3- 11.0 9.0- 12.2 6.5- 9.8- 12.2 WDI 35.7 33.3- 39.4 28.0- 36.4 28.0-33.9- 39.4 SpLI 112.0 100.6 SpRI 36.1 42.3 SpWI 5.3 8.1 ------~---~ the ink sac, the lack of which is of generic following genera are included: Eledone significance, was either not well developed Leach, 1817; Velodona Chun, 1915; Tetra- or had been undetected. Examination of cheledone Voss, 1955; Pareledone Robson, the type material from the British Museum 1932; Thaumeledone Robson, 1930; Gran- confirmed the fact that the ink sac was eledone Joubin, 1918; and Bentheledone absent. Larger specimens collected by the Robson, 1932. ELTANIN in Antarctica being studied by Eledone presents heteromorphic arms, the G. L. Voss also lack ink sacs. The examina- tips of the non-hectocotylized male arms tion also revealed that the youngest of the being modified into fleshy papillae or two types of Thaumeledone brevis, listed laminae; the hectocotylus is not differenti- as a female by both Hoyle (1885) and ated into ligula and calamus. Both charac- Robson (1932), was in fact a young male. ters distinguish it from Vosseledone. The hectocotylus, however, is not well de- Velodona is characterized by a vestigial veloped and the species awaits redescription ink sac; the funnel organ has two very by Voss. thick V-shaped pads; the ventral web of Robson (1932) did not recognize the the dorsal arms is greatly expanded; the Eledoninae sensu stricto due to polyphyletic radula is simple with an unicuspid rachidian, divergences of the subfamily's heterogeneous the third lateral remarkably thick and short. generic assemblage, and to his doubts about The well developed ink sac and the degen- the validity of grouping them under the erate radula of Vosseledone differentiates it same subfamily in which only the suckers from this genus. and the large egg size are common charac- In Tetracheledone the funnel organ char- ters. This is in contrast to the evidence in acteristically is disposed in four parallel bars favor of convergent evolution within the or pads; the hectocotylus is small but dis- Bathypolypodinae. Robson believed that tinct and the radula is normal, well de- Thaumeledone belonged with the Bathy- veloped, with a simple rachidian. Th~ polypodinae and placed the remainder of double V funnel organ, the prominent the known genera in the Octopodinae, most hectocotylus, and the degenerate radula of of which have biserial suckers. The Ele- Vosseledone distinguish it from this genus. doninae is a useful subfamily in which the Taki (1961) created the Megalcledoninae PALACIO: A NEW PATAGONIAN CEPHALOPOD 289

Table 3. Indices of bodily proportions of 4 males and 3 females of Vosseledolle charrua new species, grouped by sexes

Range Jndcx Males Females x ML 49.0- 50.5- 53.0 N = 4 58.0-65.2- 76.5 N = 3 49.0-56.8- 76.5 N = 7 MWl 96.2-102.1-108.0 86.9-93.9-103.4 86.9-98.6-108.6 HWI 69.8- 80.4- 88.0 57.4-65.8- 75.8 57.4-74./- 88.0 ALI 62.3- 64.8- 67.5 58.3-60.6- 62.4 58.3-63.0- 67.5 AWl 20.0- 22.3- 24.4 17.6-19.0- 20.6 17.6-20.9- 24.4 MAl 39.3- 60.8- 50.5 47.2-50.5- 52.6 39.3-47.7- 52.6 SnDI 9.0- 10.7- 12.2 6.5- 8.6- 10.3 6.5- 9.8- 12.2 WDI 28.9- 35.0- 39.4 28.0-32.3- 35.7 28.8-33.9- 39.4 for Megaleledone senoi, from Antarctica; it The rachidian, in combination with one of was characterized by having a small "abor- the heteromorphic series of lateral teeth, is tive" ink sac, but lacking an esophageal of taxonomic importance. The marked crop and by a small unicuspid radula. Voss heterogeneity between the multicuspid (pers. comm.) considers it synonymous heterodont radula of Pareledone and the with Pareledone, based on his study of Ant- degenerate radula of Vosseledone is con- arctic octopods. sidered to indicate a significant phylogenetic Vosseledone is closely related to Parele- gap in maintaining a generic distinction. done Robson, 1932. The generic diagnosis Nixon (1969) showed that in Octopus for the latter is brief: uniserial suckers, the radula serves primarily for the physical large eggs, normal hectocotylus and well transport of food and it seems likely that developed ligula and calamus; restricted to this is the case in most octopods. The same the southern hemisphere. Having erected role was suggested for the radula of Nautilus Thaumeledone for a form with a degenerate by Griffin (1900). However, I find no radula and lacking an ink sac, and Benth- evidence to support the view that the eledone for a form lacking a crop and an cephalopod radula represents a primary ink sac, also possessing a peculiar radula, it adaptive character or that its characteristics is surprising that Robson did not mention can be associated with any particular ecolog- any of these characters in his diagnosis of ical adaptation. In my opinion, the radula Pareledone. Based on the type species, the should be regarded as a secondary adaptive nominal species and material in the UMML character possibly resulting from a non- collections, the diagnosis of Pareledone is specific ecophysiological adaptation to biotic expanded: or abiotic environmental factors as suggested Octopods with uniserial suckers, ink sac by Golikov (1973). Nevertheless, its com- well developed, deeply imbedded but ex- plex heteromorphism in cephalopods externally visible; radula with unicuspid presses underlying genetic differences and (tricuspid) or pentacuspid rachidian; three similarities of high taxonomic value which well defined lateral teeth, marginal plates are probably consistent with phylogenetic usually present. Third right arm hectocoty- lineages. lized with well defined ligula and deeply In his discussion of the primitive or grooved calamus. Normal crop present. specialized condition of cephalopod radulae, Funnel organ of the V V or W type. Large Rob:on (1932) did not benefit from the eggs. Restricted to the southern hemisphere. more recent discovery of Neopilina gala- Eurybathic. theae, the anatomy of which was described All the nominal species of Pareledone by Lemche and Wingstrand (1959). This have well differentiated radulae (Table 4). Cambro-Devonian mollusc possesses a 290 BULLETIN OF MARINE SCIENCE, VOL. 28, NO.2, 1978

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Table 6. Synopsis of eledonine species with general geographic and vertical distribution -----==c======-==------'-''-'--=-- Approximate Vertical Bottom Species Geographical Distribution Distribution Type

Eledol/e carpati Adam, 1950 West Africa 60-170 m Dark ooze Eledol/e cirrhosa (Lamarck, 1798) N.B. Atlantic, 10-800 m Muddy Mediterranean Eledol/e massyae Voss, 1964 Patagonian 60-170 m Muddy to calcareous E!edol/e moschata (Lamarck, 1798) Mediterranean, N.B. Usually not Atlantic below 300 m Eledone thy.mnophora Voss, 1962 South Africa Tide pools Bel/the/edone albida (Berry, 1917) Antarctic 3,110 m Thick ooze and rock Bewhe/edolle rotunda (Hoyle, 1885) Kerguelen Is.-W. 3,568-4,071 m Diatom ooze Australia Gralle/edolle alltarctica Voss, 1976 Ross Sea 2,341 m Grandedone clwllel/geri (Berry, 1916) Kermadec Islands 1,153 m Gralldedone macrotyla Voss, 1976 Falkland Islands 1,647-2,044 m Grandedolle verrucosa media (Joubin, 1918) North Western Atlantic 1,458 m Gral/dedone l'errucosa \'errucosa (Verrill, 1881) North West Atlantic 2,300 m Parc!cdol/c adeliealla (Berry, 1917) Antarctica (Ade1ie coast) 300-549 m Ooze Pardedol/e carlgreni Thore, 1945 South Africa Pardedolle clwrcoti (Joubin, 1905) Antarctica. One record 0-1,500 m Sandy bottom from Rio de Janeiro Pare!cdol/c Iwrrisolli (Berry, 1917) Antarctica (Shackelton 24-595 m Ooze Ice Shelf) Pareledolle lIigra (Hoyle, 1910) South West Africa Shallow water rocks Pm'dedol/e pol)'morpha (Robson, 1930) South Georgia Islands 273 m Mud Pardedol/e turqueti (Joubin, 1905) Antarctica. One record 549 m usually from Rio de Janeiro in depths not exceeding 4,000 m Pardedol/e umitakae Taki, 1961 Antarctica (Riiser- 630-680 m Larsen Pen.) Tetrachdedollc spillicirrus Voss, 1955 Gulf of Mexico, Florida, 183-544 m Cuba Tlwumeledollc brcvis (Hoyle, 1885) Uruguay 1,098 m Green sand Tlwllme/edone gUllteri Robson, 1930 South Georgia Islands 400-410 m Velodolla togata capell sis Robson, 1929 South Africa (Natal) 402-457 m Vdodolla /Ogata togata Chun, 1915 Tanzania 748 m Vasse/edol/e charrua Palacio, 1978 Brazil-Uruguay 10-200 m 294 BULLETIN OF MARINE SCIENCE, VOL. 28, NO.2, 1978 radula with 11 longitudinal rows of teeth; probably offer us more sound bases for the fourth lateral or comb-tooth is modified comparative studies. aborally by possessing about 40 denticles. A synopsis of eledonine genera is pre- A brief review of the radulae of some sented in Table 5. cephalopods reveals the following: the primitive Nautilus has 13 longitudinal rows Etymology.-The new taxon is named in of teeth with two strongly modified laterals honor of G. L. Voss and N. A. Voss for (Vayssiere, 1896: pI. 18; Naef, 1923: pI. their contributions to the advancement of 25, Figs. 1-3; Robson, 1932: Fig. 3A; Teuthology. The specific name is given Solem and Richardson, 1975: Figs. 14-19); after the Charrua Indians who inhabited Vampyroteuthis infernalis has seven rows Uruguay, Argentina, and Brazil; the name of well differentiated unicuspid homodont is indicative of the occurrence of the species. teeth (Berry, 1912: 273, Fig. 1; Pickford, 1949: Fig. 3; Solem and Roper, 1975: ARTIFICIAL KEY TO THE GENERA WITH UNISERIAL Figs. 15-18); Argonauta and Ocythoe pos- SUCKERS OF OCTOPODIDAE sess seven rows of unicuspid teeth for the la. Tips of the non-hectocotylized male arms former and multicuspid teeth for the latter modified into small fleshy papillae or laminae; hectocotylus not differentiated Eledollc (Robson, 1932: Figs. 3D, 3F). Naef lb. Tips of non-hectocotylized male arms with (1921) and later Aldrich et a1. (1971), normal suckers; hectocotylus well differen- have shown that the Sepiolidae have uni- tiated into ligula and calamus . . 2 2a. Radula well differentiated into seven teeth 3 cuspid teeth and lack marginal plates; and 2b. Radula degenerate, lateral teeth atrophied 4 that the Loliginidae and Ommastrephidae 3a. Esophageal diverticle or crop well devel- have tricuspid rachidians, bicuspid laterals oped _ __ S and unicuspid inner and outer marginals. 3b. Esophageal diverticle or crop reduced, un- conspicuous or absent Grancledonc In the Octopodidae there is a trend towards 4a. Ink sac well developed Vosselcdonc n. gen. reduction of the first lateral, and in deeper 4b. Ink sac lacking . Tllaumeledonc water forms there is a trend towards a uni- Sa. Funnel organ disposed in four parallel pads ______.... T ctraeIIcIcdonc cuspid rachidian versus a multicuspid ra- Sb. Funnel organ V V or W . .. 6 chidian in shallow water forms. Evidently, 6a. Ink sac well developed, deeply imbedded primitive molluscs and cephalopods possess ...... ______.Parc Iedo nc well differentiated and specialized radular 6b. Ink sac degenerate or absent______7 7a. Esophageal divertide or crop well devel- teeth. In the modern octopods, however, oped; very thick V V funnel pads _Yelodona there is an overall trend to a simplification 7b. Esophagus lacks diverticle or crop; funnel of the radula. It is now possible to conclude consists of single, thick V. ..Bcntheledollc that a complex radula constitutes a more primitive condition and that a more evolved ACKNOWLEDGMENTS and specialized condition is represented by I am grateful to Prof. E. C. Rios, Dra. Z. de a simple, strong and functional structure for Castellanos and Dr. J. F. Peake for making the the transport of food. material available for study. Special gratitude is Most radulae illustrated in the literature due to C. Gordillo for her encouragement in the course of the work. Finally, I am indebted to are inadequately depicted (Aldrich et aI., Professor G. L. Voss, who is preparing a mono- 1971 ), and radular orientation has lacked graph on Antarctic benthic octopods, for the uniformity. This circumstance, in addition to availability of representatives of most of the ele- my view that cephalopod radulae do not donine species referred to here and for distribu- tional and diagnostic information. I am also constitute a primary adaptive character, grateful for the use of his library, for confirming suggest that this structure has limited value the validity of the new taxon and for his correc- in the study of octopodan speciation. Other tions and suggestions while reviewing the manu- script. This is a scientific contribution from the structures, such as spermatophores, and the Rosenstjel School of Marine and Atmospheric digestive and reproductive systems, will Science, University of Miami. PALACIO: A NEW PATAGONIAN CEPHALOPOD 295

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1962. South African Cephalopods. oda), from the Southern Ocean. Proc. BioI. Trans. Roy. Soc. S. Afr. 36: 245-272. Soc. Wash. 88: 447-458. 1963. Cephalopods of the Philippine DATE ACCEPTED: January 18, 1977. Islands. U.S. Nat. Museum, Bull. 234. 180 pp. ADDRESS: Division of Biology and Living Re- 1964. A note on some cephalopods SOl/rces, Rosl'nstiel School of Marine and Atmo- from Brazil with a description of a new spe- spheric Science, University of Miami, 4600 Rick- cies of octopod, Eledone massyae. Bull. Mar. en backer Causeway, Miami, Florida 33149. Sci. Gulf Carib. 14: 511-516. PRESENT ADDRESS: Marine Policy and Ocean 1976. Two new species of octopod of Management, Woods Hole Oceanograpllic Institu- the genus Graneledone (Mollusca: Cephalop- tion, Woods Hole, Massachusetts 02543.