Opisthoteuthis Californiana Berry!

Home , Cirroteuthis, Opisthoteuthis, Umbrella octopus

New Records and Observations on the Flapjack Devilfish, Opisthoteuthis californiana Berry!

WALTER T. PEREYRA 2

IN JUNE 1961 the U. S. Bureau of Commercial respect to the development of secondary sexual Fisheries joined with the Atomic Energy Com characteristics ( Berry, 1954 and 1955 ) . The mission to undertake an investigation of the four specimens reported by Berry (two mature deep-water ocean fauna in the area contiguous females, one mature male, and a juvenile fe to the Columbia River mouth at depths from male ) represent the only previous records of 50 to 1050 fathoms (91-1920 m). The main this species known to the author. objectives of this project are to describe the Throughout the world, six other species of benthic fish and invertebrate communities in this genus have been described . Two species habiting the study area, and to define their are known from the Atlantic (the genus type bathymetric distribution on a temporal basis. O. agassizii Verrill, 1883 and an apparent pe Associated with the faunal investigation is a lagic species, O. m edusoides Thiele, 1915), one monitoring of the various demersal forms to from the Indian ocean off the west coast of evaluate the biological transport of radionucle Sumatra ( 0 . extensa Thiele, 1915 ), two from ids which may have their origin in Columbia Australian waters ( 0. persephone Berry, 1918 River waters. and O. pluto Berry, 1918), and one which has Thirty-one specimens of the flapjack devil been taken repeatedly in Japanese waters ( 0 . fish, Opisthoteuthis californiana, have been depressa Ijima and Ikeda, 1895). captured since initiation of the field program Despite wide interest in the taxonomy and in 1961. Records of these captures, together comparative morphology of this divergent ceph with records of this species from other areas alopod group (Robson, 1925 and 1929), very in the northeastern Pacific, are used here to de little is known of their feeding habits, mode of scribe the bathymetric and geographic distribu life, or behavior. As late as 1952 Berry (p. 187 ) tion of O. californiana and to augment knowl wrote of his hope ". .. that we may not have edge of its biology. long to wait before further examples of so The flapjack devilfish was described and strange and interesting an animal will be cap named by Berry in 1949 from tWO mature fe tured and that something may then be learned males captured in 188 fathoms ( 344 m ) by a of its appearance in the living state, its behavior commercial trawler off Humboldt Bay, Califor and habits." nia. A second paper by Berry (1952) illus trates and gives additional descriptions of these ACKNOWLEDGMENTS specimens. A male and a juvenile female flap I am grateful to Dr. Paul Illg, Zoology De jack devilfish were taken tWO years later in 280 partment, University of Washington; Mr. Wil fathoms ( 512 m) off Humboldt County, Cali liam High, International Pacific Halibut Com fornia, also by a commercial trawler . The male mission; Mr. Clifford Fiscus, Marine Mammals specimen was described in detail, especially with Branch, Bureau of Commercial Fisheries; and

1 The work was supported by the cooperative Bu Miss Patsy Mclaughlin, Oceanography Depart reau of Commercial Fisheries-U. S. Atomic Energy ment, University of Washington, for loaning Commission deep-water investigation of the benthic specimens and/or data for several Gulf of marine fauna in the area contiguous to the Columbia Alaska and Washington coast records. The loan River mouth. Manuscript received December 11, 1963 . of specimens of Opisthoteuthis depressa and 2 Fishery Biologist, Exploratory Fishing and Gear Research Base, Bureau of Commercial Fisheries, U. S. Stauroteuthis albatrossi from the U. S. National Fish and W ildlife Service, Seattle, Washington. Museum by Dr. Fenner Chase and Dr. Harold

427 ~ N TABLE 1 00

COLLECTION D ATAFOR THIRIT-NINE Opistboteuth is calijorniana T AKEN IN THE NORTH EASTERN P ACIFI C

PO SITION DEPTH ~~~: SIZE DATE TEMP . ( OC) I I N O. I SEX REMARKS (fathoms) (mm) Lat. NLong. W ~ I Bot. %0 ------'------I SW of Columbia River off Oregon

1 July 1961 46° 0.5' 124° 49.0' 300-303 16.1 _...... 1 undetermined 3 July 1961 45° 51.8' 124° 48.5' 425 16.9 _. . . _... 1 mat. F 225 X 250 3 July 1961 45° 50.3' 124° 50.0' 450- 500 16.3 .... -- -- I immat. F 180 X 200 16 Sept. 1961 45° 56.3' 124° 47.8' 372-374 16.1 ...... -. 1 mat. M 460 X 440 1 mat. F 225 X 224 0 0 10 Dec. 1961 45 58.5' 124 49.1' 350 10.0 .- ...... 1 undetermined 0 0 10 Dec. 1961 45 52.2' 124 53.7' 455-475 10.0 ..-. ._.- 1 mat. F 195 X 175 W t. 21b 0 0 11 Dec. 1961 45 58.4' 124 49.6' 335 9.9 .... . _.- 1 mat. M 280 X 260 0 0 11 Dec. 1961 45 55.9' 124 48.1' 372-380 10.0 ._. - - ..- 1 mat. F 340 X 330 1 mat. M 280 X 260 1 mat. M 420 X 380 1 mat. M 205 X 200 6 Mar. 1962 450 52.4' 124 0 49.7' 446-455 9.2 4.6 . 34.192 1 irnma t. F 170 X 180 Wt. 1.5 lb ~ 7 Mar. 1962 45 0 58.5' 124 0 52.1' 415-427 8.3 4.6 34.137 1 mat. F 210 X 150 W t. 1.5 lb 1 irnrnat. F 190 X 180 Wt. 1.5 Ib c 0 0 :!l 11 Mar. 1962 45 55.5' 124 46.1' 318-333 7.8 5.8 34,021 1 immat. M 145 X 140 Wt. l Ib o 0 0 26 May 1962 45 58.8' 124 50.6' 350 13.3 ---- .... 1 mat. M 440 X 420 en 0 0 27 May 1962 45 58.1' 124 51.5' 375 12.2 ... . _ _ e . 1 mat. M 290 X 290 o 0 0 1. 7 May 1962 45 59.3' 124 52.2' 400 12.8 .... - -- - I mat. F 205 X 205 0 0 ~ !.7 May 1962 45 54.1' 124 54.5' 450 13.3 _._ . .... 1 mat. F 210 X 205 0 0 30 Aug. 1962 45 58.0' 124 44.7' 250 16.7 .... . _.- 1 mat. M 250 X 240 .Q ~ 1 Aug. 1962 450 55.6' 124 0 46.2' 350 16.7 .... __e. 1 mat. M 285 X 260 1.' mat. M 200 X 220 ~ 0 0 1 Sept. 1962 45 54.0' 124 47.2' 375 16.7 . _.- .... 1 undetermined small 0 ~ 23 Jan. 1963 45 50.9' 124 0 52.8' 440-450 8.3 4.7 34.13 1 irnmat. M 140 X 125 1 mat. M 360 X 350 .X 23 Jan. 1963 450 54.2' 124 0 53.8' 405 8.3 5.1 34.09 1 imrna t, M 145 X 128 24 Jan. 1963 450 55.0' 124 0 47.4' 348-354 8.9 4.6 34.18 1 mat. M 260 X 230 >? 0 0 3 9 May 1963 46 00.1' 124 50.8' 300 11.7 .... ._-- 1 mat. M 13 May 1963 450 55.8' 124 0 49.8' 375 12.2 _._. __e. 1 mat. M 310 X 300 ~ 13 May 1963 45 0 52.3' 124 0 52.0' 450 12.2 .-_. .... 1 mat. M 360 X 340 ...... \0 0\ VI Opisth oteuth is californiana-PEREYRA 429

Rehder is greatly appreciated. Special thanks ...... ~ 80 ~ ~ I ~ 0 are due Dr. Grace Pickford, Bingham Ocean 8..0 o tt:: ographic Laboratory, Yale Uni versity, and Mr. ~ 0 Clifford Fiscus for critical review of the manu ..0..<::o . tt ~ script . 0 ...... c:: ,-n . ~ ~ METHODS AND MAT ERIA LS ~o.OJ " ~ ~ 1 All the specimens agreed with Berry's de 1- - scription of Opisthoteuthis californiana, but no comparison was made with species types. Speci mens were compared with repres entatives of X X eJl X X ~ 0 0 IA IA two closely related species, O. depressa and IA '

00 '0 lected in the north east Pacific are reported in 0 IA ,-n 0 ONO oo oo this paper (Table 1) . Of this total, 31 speci I IA ,-n I 0 ...... '0 '<1' ...... N Or('\.....-I 0 IA mens were taken on the Columbia River track ,-n N line by the MjV "Commando" and "John N . Cobb," two off the coast of W ashington by the ~V: 0",, 000 ...... IA Oo\v-\tr\ r---: ...... ,-n OO "<:f'~...... -I "Cobb," and six in the Gulf of Alaska by the

o 0 0 0 0 o 0 "Cobb" and chartered vessels of the Interna IA IA V"'\O O\lI"'\ V"\ N N 'OlAIA '

N ,-n ...... NNNC"I fore, although all of these captures, with one IA '0 '0 '0 '0 '0 '0 0\ 0\ 0\ 0\ 0\ 0\...... 0\...... excep tion, are treated as occurring on the bot ...; >- >->- . ~ ...... tom, the possibility exists that they could have 0. ", cod t'1 O-t A o, ~ ::E ::E ~ -< ,,"$ ,,OJ been taken as the gear was being set or hauled IA '

Observations on living animals were made depths (5 87 and 1058 fathoms [107 3 and in a 20-gallon aquarium with a sealed top 1934 m], respecoively) than O. californiana. aboard the M/V "John N. Cobb" during a Th e bathymetric ranges of the other four spe cruise in January 1962. Motion pictures were cies, O. extensa, persephone, pluto, and mede taken at this time which allowed a more de soides, fall within that of O. californiana. tailed behavioral analysis to be m ade on shore. The temporal distribution of males and fe males by depth along the Columbia River OBSERVATIONS trackline is presented in Figure 1. Although Geographic Distribution the number of specimens is small, males appear to OLCUpy a greater bathym etric range than Prior to this study the known geographic females. The latter were never captured at distribution of O. californiana was recorded depths shallower than 375 fathoms (6 86 m ). from four specimens reported by Berry (1 949: There are, however, three known records of 24; 1952 :18 3; 1954:29; and 1955:219,223). females from levels shallower than 375 fath The exact locations for these captures are not oms (686 m) : one off California in 188 fath given, but from Berry's descriptions they ap oms (344 m )( Berry, 1949 :24 ) , one off Wash parently were confined to a small area off Hum ington in 150 fathoms (274 m ), and one from boldt Bay, California (approximately 45° lO'N, the Gulf of Alaska in 68 fathoms (1 24 m ) . 125°1O'W). On the trackline the data suggest that the The data in Table 1 extend the geographic males may move seasonally, being found at range northward 2200 miles along the conti shallower depths in the summer. Perhaps mat nental shelf to off Unimak Island, Alaska ing takes place in late winter when the sexes that is, from northern California to the Aleu occupy similar depth zones, but this is only a tian Islands, Alaska. surmise. Bathym etric and Seasonal Distribution In cephalopods a separation of the sexes by depth has been noted by several authors. Is The known bath ymetric range of this spe grove ( 1909:472) notes a disparity in the sex cies off northern California as recorded by ratio of Eledone. She suggests that the dis Berry (ibid.) is from 188 to 280 fathoms (344 parity is caused by the females breeding in 512 m ). Specimens collected from the Colum littoral waters, in which they are more fre bia River trackline were taken at depths from quently taken. Robson ( 1925: 1325 ) presents 250 to 450 fathoms (457-823 m ); those off figures for Octopodidae picked at random from W ashington from 150 and 308 fathoms ( 274 selected reports, where inside of 200 fathoms 563 m ) ; and those from the Gulf of Alaska (366 m ) the percentage of females is greater from 68 to 300 fathoms (124-548 m ) . The and outside the percentage of males is greater. Both of the above findings, where the female above records extend the known bathymetric Ocropodidae predominate in shallow water, range shoreward to 68 fathoms (12 4 m) and are reversals of the situation which has been seaward to a depth of 450 fath oms (823 m ). found in this study for Opistboteuibis, where As has been observed for the western Pacific the males predominate in shallow water. congener ( 0. depressa ) , O. californiana has Of the specimens collected off the north been collected from shallower water at the Oregon coast for which the sex is known, 18 northern end of its range, with the bathymetric males and 10 females are recorded . Since greater range being wider to the north despite infre trawling effort was expended at the depths quent trawling in water deeper than 300 fath where the males predominate ( in the shallow oms in northern waters. part of the range ) , the above ratio probably The known bathymetric ranges of the other does not reflect the actual sex ratio . species of Opisthoteuthis are not all similar (Robson, 1929:167-173 ). Both O. depressa Availability from Japanese waters and O. agassizii from Catch-per-hour trawling for O. californiana the N orth Atlantic have been reported at greater at various depths is given in Figure 2. The data opisthoteuthis californiana-PEREYRA 431

LEGEND 01 Male .2 .. D 1 Female 250 .2 " • Male Dist'n

C/) 300 ~ 0 ::I: ~ LL: 350 ~ ::I: t- o, 400 W 0 ~ ~ 450

5001...--....I..._...... J._-'"--_....I..._...... J._-'"--_....I..._~--.!.--....I...----:-'--:-----:=-_-l J F M M J MONTH

FIG. 1. Temporal and bathymetric distribution of male and female Opistboteutbis calijorniana based on capture of 28 individuals on a trackline off the mouth of the Columbia River, July 1961-May 1963. suggest that Berry (1949 :26 and 1952:183) The bimodal nature of the catch-per-unit-of was correct in surmising that these anima ls are effort curve may result from several factors. As pro bably not unduly rare where suitable bot shown earlier, the sexes are distributed un tom conditions exist. The average catch rate evenly, with the males found in shallower water over the depth range where they were taken than the females. This could create the appar on the Columbia River rrackline was approxi ent non-normal distribution. Another factor mately 0.4 individuals per hour of trawling which may be responsible is the direct or in (Table 2 ) . Within the range of depth where direct effect of bottom type on distri bution. the species were found, they were most fre Green mud was found to predominate over quently encountered from 375 to 450 fathoms the range where the species was encountered; (686-823 m). Due to the low effort from 475 but inside of 325 fathoms (594 m ) and at 400 to 575 fathoms (868-1051 m ), where only fathoms (731 m), where the catch per effort two drags have been made, its relative abun was low, the sand/mud ratio was higher dance in deeper water is not well known. ( > .08) than was found at adjacent stations 432 PACIFIC SCIENCE, Vol. XIX, October 1965

rounded eye prominences, which are noticeable 1.00 on the upper surface of preserved material , '"

FIG. 3. Behavior of live Opistb otesahis californiana in an aquarium. a- d, swimming off bottom ; e-], set c:ing back to bottom. the vessel rolled, flapjack devilfish moved back cephalopod group and allows questions as to and forth in the aquarium . A small Octopus the function of these enlarged suckers. Assum sp., placed in the aquarium at the same tim e, ing that these animals behaved normally despite had no trouble "hanging on" or clinging to the the fact that they had been brought to the sur aquarium. The inability of the flapjack devilfish face, it would not seem that this cephalopod can to attach to the aquarium glass, even though apply itself to a rock as stated by Borradaile the males possess many greatly enlarged suck et al. (1 935:602), or attach to a proj ecting ers, sheds more light on the degeneracy of this substratum as stated by Ijima and Ikeda ( 1895: 434 PACIFIC SCIENCE, Vol. XIX, October 1965

328), or is a creeping or clinging form as Ver cion, There was no concentric streaking as Berry rill (1896:74) thought. observed for preserved material, and no aboral The highly specialized dorsal arm-pair of the spotting as is reported for several congeners males, regarded as the hectocotylized or nuptial (Berry, 1918 :286, 29 1; Sasaki, 1929: 11; and arms, with their 6-7 greatly enlarged globular Verrill, 1883: 114 ) . The oral surface was like suckers distally, is characteristic of this species wise dark reddish brown but without the breaks (Berry, 1955 :221-222). This condition, of in the background coloration. This dark reddish course, reaches its highest degree of develop brown coloration diffuses to a pale reddish tan ment in the mature males; but even with the color in the region of the cirri , suckers, and smaller immature males, three of whi ch are the central portion of the arms. After preserva present in this series, the distal suckers of the tion the coloration was simil ar to that reported dorsal arm-pair are slightly enlarged. Th e sexes by Berry (1 952: 186 ). are, therefore, always distinguishable by this The light blue breaks in the reddish brown coloration were due to splitting of the delicate male nuptial arm character. outer skin of the aboral surface by trawl abra The internal structuring of this group is more sion and from the spines of fish in the catch. striking than the external. Many organs are This point was verified by purposely splitting completely lost with no new structures replac the outer skin of freshly caught specimens . The ing them. There are no salivary glands, no ink light blue colorati on became evident immedi sac, no well defined crop, no radula, and no ately. The degree of skin splitting varied with right oviduct (Berry, 1952 :185 ). The stomach the individual, some being so badly abraded is quite small, and is divided into two parts; that the brown coloration was almost com the liver is bilobed. The reproductive apparatus pletely wanting. of both sexes is massive, and in mature indi viduals it constitutes a considerable porti on of Fecundity the visceral mass ( Figs. 4 and 5 ). Egg counts and measurements were mad e Color on three mature females captured at different times of the year. These data are presented in In life the coloration differed slightly from Table 3. that reported by Berry (1949:24 and 1952: The total egg count varied from 225 to 475 186 ) for preserved material C.. . light dull eggs. In none of the individuals examined was drab, very heavily concentrically streaked with the egg size uniform; it varied from less than dull dark reddish brown . .."), being a more one mm to 11 mm in greatest diameter (length). evenly distributed dark reddish brown with The largest egg examined measured 11 by 6 breaks of light blue in the background colora- mm, which is slightly larger than the largest

FIG. 4. Reproductive organs of a mature female FIG. 5. Reproductive organs of a mature male Opis Opisthoteuthis californiana, X .45. th oteuthis californiana, X .65. opisthoteuthis californiana-PEREYRA 435

TABLE 3

F ECUNDITY OBSERVATIONS ON THREE MATURE FEMALE Opisthoteuthis californiana TAKEN ON THE COLUMBIA RIvER T RACKLIN E

LARGEST TOTAL NO. EGGS NO . EGGS DEPTH SIZE % EGG (mm) DATE VOL UME > 3mm < 3 mm LARGE (fathoms) (mm) LENG TH X EGGS (ml) (length ) (length ) EGGS WIDTH . 3-7- 62 425 210 X 150 5.0 98 150-200 33-39 10 X 5 7-3- 61 425 255 X 250 3.3 125 150-200 46-56 9 X5 12-11-61 I 375 I 340 X 330 \ 23.0 324 I 125-150 I 68-72 I 11 X 6 egg reported by Berry (1952: 185) (9 by 5 mm). identification was not possible. A few arnphi The percentage of large eggs (greater than pods, isopods, and a masticated shrimp, pro b or less than 3 mm in length was arbitrarily ably a hippolytid or crangonid species, were chosen to designate large and small eggs, re also recognized. N o quantities of mud were spectively) differed for each individual, with a mixed in with the stomach contents, but a few range from 33 to 72 %. Likewise, the volume sand grains were found in the stomach of the of eggs in the ovary ranged from 3.3 ml to 23.0 Unimak Island specimen. In some stomachs a ml. In the specime n with 72 % large eggs (egg brown organic material was found mixed with volume of 23.0 rnl}, the majority of the large the food, but the exact composition of this eggs were located in the upper part of the material is not known. Fecal remai ns left in ovary with ripe eggs spirally arranged in a the aquarium by the live individual appeared compact gro up just under the oviducal gland to contain this same material. ( Fig. 4). In no case was there any indication The above summary of gut-contents par· of stalk development. tially substantiates the speculation by Berry The extent of egg development was used as ( 1952:187) that this species feeds mainly on circumstantial evidence indicating the season of micro-pla nkton or detr itus swept down the disc ovulation. The highest degree of egg develop by the radial series of cirri toward the mouth. ment was observed in a female containing ripe The detrital element of its diet has not been eggs, captured on the 11th of December. The substantiated, but the presence of the small least development was found for the March crustaceans in the gut verified rhe planktonic capture, while the female captured in July showed an intermediate degree of development. feeding habit. The size of the plankton indi With only three samples represented no un cates that the animal is more of a macro- than equivocal conclusions are permitted, but indica a micro-plankton feeder. tions are that ovulation occurs during the A few mysids found in th e esophagus of one winter. individ ual were entire, as were most of those found in the first stomach of the others. Only Food one organism was noticeably bitten, the small The stomachs of eight specimens were ex hippolytid or crangonid shrimp. Thus it ap amined to ascertain the food organisms (Table pears that except for a few of the larger or 4). Only one stomach was found completely ganisms the majority were taken entire ' and emp ty and the remaining stomachs contai ned not first reduced by the strong beak. As Rob very little, never more than 5 ml. The material son (1925 :1348-1349) has suggested, the food in greatest abundance was crustacean remains, is probably ground down in the muscular first mostly small mysids (approximately 7 mm in stomach and digested in the second, instead of length) and large copepods. Only one non· being partially comminuted by the radula with crustacean organism was found, a polychaete; preliminary digestion in the crop, as occurs in and this was partially digested and positive species that possess the latter two structures. 436 PACIFIC SCIENCE, Vol. XIX, October 1965

Locomoti on powerful downward stroke, which reduced the opening into the concavity of the now closed Observations made on live individuals in an umbrella to about one-half the diameter of the aquarium revealed a most interesting locomo animal (Fig. 3c ). This actio n forced the ani tory pattern. Although most octopods, with the mal off the bottom by (1) the actual contact exception of the pelagic species .such as Cirro of the downward thrusting arms with the bot teuthis, move about pr incipally by two meth tom and (2) the interaction of the closing oral ods, "crawling" and rapid ejection of water surface with the water. This pattern of opening from the siphon, the flapjack devilfish, owing and closing the umb rella was repea ted so that to its unusual anatomical features, is not able the animal remained off the bott om. The clos to move effectively by these means. Th e very ing action was about twice as rap id as the open weak suckers and the loss of free arm move ing. With each stroke the devilfish would move ment -due to webbing prevent crawling; and from 4 to 12 inches depe nding on the thrus t of the reduction in the mantle aperture consequent the closing stroke, and the tempo and direc to the posterior placement of the siphon greatly tion of movement. T he swim ming motion was decreases the effectiveness of the siphon as a o bserved to progress as rapidly as one stroke locomotory organ. .every two seconds, although the usual tempo The principal means of progression, as ob was once every four seconds. With movement served in the aquarium, was by opening and at the higher rate the animal was carried to the closing of the webbed "umbrella" together with surface of the aquarium in three to four strokes. fin movement. Th e general patt ern of this method of locomotion, whi ch was analogous to Subsequ ent to termination of swimm ing, the that of a jellyfish, is seen in Figure 3 a-f. anim al opened its umb rella into the convex From a resting state the animal initiated shape, thereby descending slowly ( Fig. 3f) . moveme nt by extreme opening of the um When swimming the animal occasionally made brella ( Fig. 3b) . This caused the oral surface partial strokes instead of complete ones. to take on a convex shape with the arms With contraction of the umbrella a jet of curved slightly upwards and anter iorly. Fol water was ejected from the siphon ( Fig. 3c ). lowing this, the umbrella was closed by a This jet could be directed by the siphon left

TABLE 4

STOMACH CONTENTS OF E IGHT Opistboteiabis calii orniana COLL ECTED INTHE NORTHEAST PACIFIC

DEPTH ANIMAL STOMACH DATE LOCATION CONTENTS (fatho ms) J SIZE (rnm ) VOL. (ml ) 5-14-61 425 Gu lf of Alaska 350 X 340 <1 crustacean remains including 1 amphipod ; some sand gr ains 7- 3- 61 425 off north 255 X 250 <1 noth ing whic h could be made out Oregon coast 12- 11- 61 335 off north 420 X 380 empty Oregon coast 12- 11- 61 375 off north 280 X 260 2 crustacean remains; 3 mysids approx. Oregon coast 7 mm long 12- 11-61 375 off north 340 X 330 4 8 mysids, largest 10 mm ; 4 amphi Oregon coast pods, 9 copepods, 2 isopods, 1 crango nid or hippolyrid shr imp approx. 22 mm 3-7- 62 425 off north 190 X 180 crustacean remains, mostly copepods Oregon coast 3-7- 62 425 off nort h 210 X 150 3 same as above plus partly digested O regon coast polychae te? 8-30- 62 250 off north 250 X 240 crus tacean remains, mostly copepods Oregon coast opisthoteuthis californiana~PEREYRA 437

or right, to the posterior, or down. But be- . Thus , by moving the fins alternately or together, cause the ejection was feeble, it added only and by adjusting the axis of movement and the slightly to the net movement. Verrill (1896: strength of the stroke, the animal had consid 74 ) and Berry ( 1952: 184 ) were correct in erable control over the direction of movement stating that the siphon appeared to be toO small' created by the powerful undulations of the to be used for efficient .locomotion. umbrella. The animal might swim up, side The tWO dorsal fins, which heretofore had ways, or down depending on its attitude at the been considered as "inefficient looking" (Berry, time of the umbrella closing. 1952: 183), moved in a highly coordinated The tempo of 'fin movement was always manner, augmenting the effect of the um greater than that of umbrella movement. Usu brella movement. (This is consistent with their ally it was in the ratio of three fin strokes to morphology, inasmuch as they are supported every umbrella stroke, bur this varied. Also, by a cartilaginous rod and possess ample rnus the fin movement was coordinated so that a culature. ) Balancing and orient ation of the ani down-stroke coincided with the closing of the mal were the main functions of the fins, bur umbrella. during certain phases of swimming they ap When the animal was at rest on the bottom, peared to assist locomotion. fin movement usually continued, but at a When at rest the fins were positioned at slower rate and in various directions. Also,.the the side of the head just behind the eyes in a pattern of fin movement was generally alternate. down position, with the free-edge which con As . the swimming undulations of the um tained the cartilaginous rod posterior (Fig. 3a brella ceased and the animal settled to the bot left). The fins could point slightly posteriorly tom, the tempo of fin movement increased con but never anteriorly. siderably in an alternate pattern. This increased Fin movement began with an up-stroke ( Fig. fin activity probably helped to slow the ani 3a right) . The supported posterior edge led on mal's descent and provided for better attitude this stroke, resulting in the fin moving through control. '. the water with the least resistance (Fig. 3d) . The alternate fin movement pattern was sur This stroke was terminated after the fin had prising. Considering that both fins are sup moved upward almost 180 0 ( Fig. 3e) . On the ported by a common cartilaginous rod, this rod down-stroke the supported posterior edge again must be able to undergo considerable torsion moved the fin, but this time the transverse axis for the fins to be moved alternately. was perpendicular to the direction of fin move The use of phorofloods to take pictures re ment , offering the greatest surface area to come vealed a negative light reaction. Prior to turn in contact with the water (Fig. 3c) . Because ing on the photofloods, the animals actively of this, fin movement on the up-stroke was moved in the aquarium under subdued light. faster than on the down-stroke. This pattern When the phorofloods were turned on, activity of least resistance on the up-stroke and great continued for 15 to 20 seconds and then all est on the down-stroke created an upward re movement ceased. As long as the photofloods sultant force. Both fins were usually moved were on, the animal remained perfectly still. together in the same direction, but occasion Several authors have hinted at the existence ally they were moved alternately, or one might of the pulsating locomotory pattern from be moved while the other was not. (Fig. 3b, [), knowledge of the gross morphology of the Although the basic fin movement was always species. Ijima and Ikeda (1895 :328) stated the same, the axis of movement could be shifted ".. . that alternate closure and expansion of the 90 0 forward from the up-down to an anterior arms is of much greater moment to Opistbo posterior direction or to any degree between teuthis than to most other Cephalopods, since these extremes. The greatest surface area of the the ejection of water from the comparatively fin was exposed on the posterior or down small branchial chamber and siphon must be of stroke. This permitted net movement in any subordinate significance." Berry ( 1952:184 ) direction from upward to forward or anterior. offered similar speculations, stating that the 4 3H PACIFIC SCIENCE, Vol. XIX, October 1965

undulations or opening and closing of the arms Another possible functi on might be food con might be quite significant. Robson (1929:22, centration. At tim es the animals were observed 28) also stated that the web is probably used just off the bottom of the aquarium making in in locomotion. He commented that the web complete swimming motions. Could this pos becomes deep independently in the Cirromorpha sibly represent a method whereby food is con and the abyssal Ocropodidae and that this con centrated by means of the produced currents? vergent resemblance is suggestive of some iden Once the food was concentrated, the animal tity of adaptation. could settle to the bottom with the umbrella Verrill ( 1896:74-75 ), by deducing function open. This might trap the food organisms from structure, pUt forth several possibilities as under the umbrella and permit feeding. La means of movement for members of this An escape function is indicated from the .group that conflict with the above opinions. He reactions of provoked indi viduals in an aqua felt that Opisthot euthis was a crawling or creep rium: they moved rapidly upwards when ing form, having concluded that the union of prodded. the eight arms with the web formed a disc-like Habitat ventral foot, and therefore, was analogous to that of a chiton or limpet. The small lateral fins It is generally felt that members of this i ndicated to him that Opisthot euthis could group (w ith the exception of O. medusoides ) .swim, more or less, and that undulatory move are bottom dwellers. This conclusion has been ments of the lateral edge of the pedal disc deduced by several authors through arguments might be possible. But he felt that a pulsating from structure to habitat ( Berry, 1952:184; swimming motion by the umbrella, similar to Dalla, 1912; and Meyer, 1906) .The pigmenta that of Cirroteutbis, would not be possible be tion, depressed form, dorsal eyes, and general cause adhesion of the arms and web to the body body shape all are indicative of a benthic exist would prevent their free use. It can be seen that, ence. Observations and data from this paper by approaching function in this manner, his are further supporting evidence for a benthic deductive reasoning was, for the most part, existence. erroneous. .Even though numerous drags along the Co Certain Octopodidae have also been ob lumbia River trackline have been made shal :served to swim by opening and closing the web. lower and deeper than the known bathymetric Verrill (1882:373) states that Bathypolypus range of this species, the captures of flapjack .arcticus swims in this manner, as does Eledone devilfish are restricted to the interval from 250 moschata, according to Orbigny and Ferussac to 450 fathoms (4 57:-823 rn ). This indi cates (18 40 ) . Wells (1962: 3) comments that Cirro thauma employs a jellyfish-like locomotion. An that the species is benthic; for if it were pelagic, -orher species of Eledone, E. cirrosa, uses its web and if the trawl captured these animals while only when sinking downwards (Isgrove, 1909: being set or hauled, we would expect to have ' 4 72-473). captures over a wider depth range. The one off The main function of the swimming pattern bottom capture, which might be argued as could be to facilitate travel from one area to being pelagic, should still be classed as a bot the next. This is indicated by the fact that one tom capture, because of its proximity to the specimen was apparently captured three fath bottom. oms above the bottom in an off-bottom trawl. Extensive midwater trawling off central and 1£ bottom currents were strong enough, the ani northern Oregon by Oregon State University mal could travel considerable distances by personnel has not produced a single flapjack .swimrning off the bottom and allowing the cur devilfish (Dr. William Pearcy, written commu rents to carry it passively. Shorter movements ' nication ). This is further supporting evidence ·could be made by orientation forward together of a non-p elagic existence. with umbrella pulsations. A combination of Observational data further supp ort a benthic these two methods would allow the greatest for existence. The negative buoyancy, together with ward progress. the manner in which the animal swims off the Opisthoteuthis californiana- PEREYRA 439 bottom and settles down again, are supporting Evolutionary Considerations evidence. From what is known of the morphology and Although the animal is primarily a bottom habits of Opisthoteuthis, most authors have felt dweller, it is doubtful that it burrows in the that members of this group are highly evolved, mud. The locomotory behavior of specimens exhibiting numerous degenerate conditions in an aquarium, the capture of one specimen (Berry, 1952; Dollo, 1912; Meyer, 1906; Rob JUSt off the bottom, and the lack of mud and son, 1925 and 1929; and Verrill, 18%). The infauna in the gut imply epibenrhic existence. evolutionary significance of some of these con Although most of the food organisms recovered ditions is not thoroughly understood and is from the gut were not epi- or infauna, indicat questioned by Robson (1929). ing a truly benthic feeding habit, none ap The following observations are further addi peared morphologically to be pelagic. tions to the concept of specialization and de Hydrographic data have been taken coinci generation for Opisthoteuthis: (1) the pres dent with some of the captures of O. californi ence of large suckers which are ineffective for ana. Bottom water temperature ranged from holding on, and (2) a pulsating swimming 0 4.6 to 5.8 C, and salinity was approximately motion through use of the web, together with 34.1 %0 , and bottom type consisted primarily highly coordinated fin movement. of greenish silt, occasionally mixed with small The adaptive significance of the deepening amounts of sand and clay. Since this bottom of the web convergently in many of the OCto type is associated with a smooth bottom which podidae is suggested by the locomotory pattern is, therefore, more conducive to trawling, flap exhibited by Opisthoteuthis and others (Bathy jack devilfish may inhabit rocky areas which polypus arcticus, Eledone moschata, E. cirrosa, have not as yet been sampled. Cirroteuthis, and Cirrothauma) . . All have an Authors in the past have classified Opistbo extended web and use it in some phase of teuthis as an abyssal animal ( Dollo, 1912 and swimming. When the behavior patterns of Meyer, 1906) . According to Hedgpeth (1957: more deep-webbed species are known, we prob 21) the limits of the abyssal region are from ably will find that the extension of the web is 2000 or 3000 m (1092-1622 fathoms) down primarily a locomotory adaprion, ward to about 6000 m (3241 fathoms) . The SUMMARY upper limit may better be defined as the region 1. Thirty-nine records of the flapjack devil in which the temperature never exceeds 4 0 C. fish, Opisthoteuthis californiana, from off the Bruun (19~7: 643) remarks that in the Atlan coasts of Oregon, Washington" and Alaska are tic Ocean rhisremperature is not reached until reported. about 2000 m (1082 fathoms ), whereas in the 2. The known geographic range of this spe Indian and Pacific oceans it is as high as 1500 cies is extended from northern California to or 1000 m (811 or 541 fathoms). If we note the Aleutian Islands, Alaska, and the known the upper limit of the abyssal region in the bathymetric range is now defined from 68 to areas of capture, then we must conclude that 450 fathoms (124-823 m) . 3. Males occur in shallower water than fe Opisthoteuthis has not been taken in the abys males, and there is an indication that the males sal zone. With few exceptions, captures of move seasonally, being found in shallower Opisthoteuthis have been reported from the water during the summer. The overall sex ratio continental slope. These exceptions were a few appears to be close to 1:1. specimens of O. depressa taken on the conti 4. At the depths of greatest availability nental shelf off japan at the higher latitudinal (375 and 450 fathoms [686 and 823 m}) off limits of this species' range (Sasaki, 1920: northern Oregon a catch rate of one specimen 170 ) where bottom temperatures are lower per one-hour drag was recorded. (Dr. Felix Favorite, personal communication) ; 5. Observations on living animals revealed and one specimen of O. californiana captured that the general appearance was more nearly a off Alaska. bell-shape than a plano-convex disc, with the 440 PACIFIC SCIENCE, Vol. XIX, October 1965 eye prominences not noticeable; that at rest deavor" in the Great Australian Bight and there was a horizontal curvature of the distal other Southern Australian localities. BioI. portion of the arms anteriorly and that the Results Fishing Exper iments carried on by oral surface was not completely in contact with F.I.S. "Endeavor," 1909-1914, Fisheries 4 the bottom; that the suckers were ineffective ( 5) :201-298. or not used by the animal to hold to the bot --- 1949. A new Opisthoteuthis from the tom; that contraction and expansion of the um eastern Pacific. Leaflets in Malacology 1 (6 ) : brella together with coordinated fin movement 23- 26. were the means of movement, with ejection of --- 1952. The flapjack devilfish, Opistho water from the siphon being of very minor teuthis, in California. Cal. Fish and Game importance; and that the color in life was dark 38 (2) :183-188. reddish brown with the bluish blotches being --- 1954. The male flapjack devilfish. ( Ab the result of rough treatment in the trawl. stract) Amer. Malacol. Union, Ann. Rept. There was no concentr ic streaking. 1953:29. 6. Hecrocotylization of the dorsal arm-pair --- 1955. The male flapjack devilfish. Cal. can be detected in all males, regardless of Fish and Game 41 (3) :219- 224. maturity. BORRADAILE, 1. A., et al. 1958. The Inverte 7. Th e total egg count of mature females brata. 3rd ed. Univ. Press, Cambridge. 795 pp. examined ranged from 225 to 475 eggs, with BRUUN, ANTON F. 1957. Deep sea and abys the largest egg found measuring 11 by 6 mm sal depths. In: Treatise on marine ecology at the greatest diameters. Ovulation was tenta and paleoecology. Geol. Soc. Amer., Memoir tively concluded to occur in the winter. 67, Vol. 1, Ecology, Chap. 22:641-672. 8. Gut-content analysis showed that these DOLLO, 1. 1912. Les cephalopodes adopres 11. la animals feed on small crustacea, mostly large vie nectique secondaire er 11. la vie benthique copepods and small mysids. A brown organic terti are. Zool Jahrb. Suppl. 15 ( 1) :105-140. substance was found in several stomachs. H EDGPETH, JOEL W . 1957. Classification of 9. The benthic existence of this species was marine environments. In : Treatise on marine further substantiated from catch records and ecology and paleoecology, Geo!. Soc. Amer., observational data. Memoir 67, vol. 1 Ecology, Chap. 2:17-28. the presence of the large ineffective suckers IJIMA, I., and S. IKEDA. 1895. Description of 10. Th e specialized locomotory behavior and Opisthoteuthis depress» n. sp. ColI. Sci., further substantiate the degenerate and special J. Imp. Univ ., Tokyo, 8 :323-335. ized nature of Opisthoteuthis. ISGROVE, A. 1909. Elydone. 1.M.B.C Memoirs, ADDENDUM London (not seen ). MEYER, W ERNER T. 1906. Die anarornie von After the present paper was submitted, Dr. Ooistboteutbis -depressa (ljima und Ikeda) . D. B. Quayle, Fisheries Research Board of Can Zeits. wiss. Zoo!. 85 ( 2) :183-269. ada, Nanaimo, B. C , provided the author with D'ORBlGNY, A.,and A. DE FERUSSAC. 1840. His the following tWOadditional records from the toire naturelle generale et particuliere des northeastern Pacific: ( 1) Off Unirnak Island, cephalop odes acetabuliferes, vivant er fos Alaska (53 °39'N, 164°44'W) at 238-252 siles. Paris, 1834-48 ( not seen ). fathoms on August 13, 1964; and (2) off north ROBSON, G. C 1925. The deep-sea 'Octopoda. ern Washing ton (47°58'N , 125°47'W) at Proc. Zool. Soc. London, 1925, Part 2:1323 505-510 fathoms on September 6, 1964. The 1356. latter record extends the bathymetric range to --- 1929. A monograph of the recent 510 fath oms. Cephalopoda, based on collections in the British Museum (N atural His tory) Part I, REFERENCES Octopodinae, 236 pp. Part II, Th e Octopoda BERRY, S. STILLMAN. 1918. Report on the (excluding the Octopodinae) (1 932 ), 359 Cephalopoda obtained by the F.I.S. "En- pp. London. opisthoteuthis californiana-PEREYRA 441

SASAKI, MADOKA. 1920. Report on cephalo Rept. U. S. Comm. Fish. for 1879. Part 7: pods collected during 1906 by the United 211-445. States Bureau of Fisheries Steamer "Alba - -- 1883. Supplementary report on the tross" in the northwestern Pacific. Proc. U. S. "Blake" cephalopods. Bull. Mus. Comp. Zool., Nat. Mus. 57(2310) :168-170. Harvard College, 11(5) :105-115. --- 1929. A monograph of the dibranchi --- 1896. The Opisthoteuthidae, A remark ate cephalopods of the Japanese and adjacent able new family of deep sea Cephalopoda, waters. Hokkaido Imp. Univ., Sapporo, J. with remarks on some points in molluscan ColI. Agric., vol. 20, 357 pp. (suppl. no.) . morphology. Amer. J. Sci. II, art. 11:74-80. THIELE, J. 1915. See Chun, C.,'1915. WELLS, M. J. 1962. Brain and behavior in VERRILL, A. E. 1882. Report on the cephalo cephalopods. Stanford Univ, Press, Stanford, pods of the north-eastern coast of America . California. 171 pp.