SYSTHL"'TI CS A:\D NATURAL HI STORY of the MIDWATER FISH LYCODAPUS NANDIBULARIS GILBERT in Califorciia WATERS

$SYSTHL"'TI CS A:\D NATURAL HI STORY of the MIDWATER FISH LYCODAPUS NANDIBULARIS GILBERT in Califorciia WATERS$

SYSTHL"'TI CS A:\D NATURAL HI STORY OF THE MIDWATER FISH LYCODAPUS NANDIBULARIS GILBERT IN CALIFORCiiA WATERS

A Thesis Presented to the Graduate Faculty of California State University, Hay"ard

In Partial Fulfillment of the RequiremcJtts for the Degree Master of Science in Biological Sciences

Michael Cr1c Anderson

Bet~een March 1974 and Janttary 1976 over 600 specimens of the eelpout codapus mandibztZaris Gilbert were captured using a midwater tra~.;l in the Honterey Submarine Canyon, California. The taxonomic history of this formerly rare fish and its congeners was found to be confused. The type specir::ens of Lycodapus were compar to recent material and the nominal species L. attc~Iiat~ca~

n and L. grosai na (in part) proved to be synonyms of L. ma;zdibularis. Confusion a:nong aodapus species has resulted from the lack of attention to sexual dimorphisr::, variation in gill raker lengths, meristic counts, head pore patterns and coloration. Populations of L. mand~bulcris are kno~n from Pacific coast submari11e canvons and deep water inlets of ~ashington, British Colur::bia and Alaska, where the fish inhabit the upper midwater, generally abo~e

500 meters. In ~lonterey Bay the speci cs underh·ent a dicl vertical migratio11. Individuals were more frequently encountered in an inshore are3 of ~!onterey Canyon than fart]~er offshore. Postlarvae and young juveniles were encounter as deep as 700 meters, but generally were found above 30C meters.

Using otolith annuli, fish 1vcre found to be at lc·e1st five years old, with the largest fish measuring 197 mm totQl length. The growth rate was equal in both sexes a indeterminate after the first year. 01o seasonal cycles fish abundance or sex ration were noted. The greatest frequency of young of the year was found during w.nter months, but trawling effort was also more concentrated 1n those months. The majority of the fish taken throughout the year were two and three year olds, the ages at onset of maturity. Four egg size classes were evident ~nd the}· developed synchronottsly until the largest class reached

·~ r about 0.5 mm in diameter. At this time, one complement ~) j about 100 ova grew maximally to about 1.7-1.9 mm in Jia· meter. Egg size complements measured every month inJic3t2d this maturation occttrs once a )rear, but monthly ovarian weights indicated there is no restricted spawning se~scn.

Juvenile fish consumed fewer prey species thnn adults, hilt both stages relied prim~rily on cuphausiids and copepocs.

Feeding occurred chiefly in ncar surface waters in th0 c~r}~r morning, after which many fish retreated to the deeper areas of the canyon. The initi3tion of feeding was thotJg]:: to be stimulated via two sensory mechanisms. First, dis· turbances 1n the water arc probably detected from stirnn- lation of the larger ccpl13lic latcralis system and the smaller free organ systc1n, then vistial ct:cs arc most likely used when the fish come tn close p1·oxirnity to the prey.

1'Ko parasites were ol)scrvcJ; the nem~ttoJe T nnascQr~s a~:t~c:~m occurred in 29.2~ of the adults and 15.2~ of the ju\·eniles S3mpled, while the copcpod Cardioiectes m2~usce~s occurr~d on 0.9% of the ~Jttlts and 1.4% of the juveniles. Fish in an aquarium were genernlly sluggish, often SK~1m together and lived a maximum of twelve days at the optimal temerature of 7°C. ACKNOWLEDGMDJTS

This study is, 1n part, a result of a midh·:1tc:r :·;.:;:;pl lng program in Monterey Bay, California, but material rren1 outside the Bay, obtained from other sources was also usea.

Any midwater trawling project requires the energies anJ resources of many people not only to organize and handle sampling, but also to help with species identifications, literature problems and previous collections relevant to the study. I am priviledged here to gratefully thank all those connected with this study.

The program was initiated bv Vic E. Breeden, Sr.

3nd .John E. f·kCoskcr, Superintendent of Stc:inhart Aquariun,

San Francisco, and supported bv a grant fro111 the Ch1rlinc l!. Breeden Foundation to Dr. )!cCosker. Otis Burton, c:xplorer and inventor of the bathysphere, helped launch the prograr;·.

~nd inspired In n1e a deep fascination for the crcattircs of the deep-sea. Shi]1tin1e ~tnd schcdtzling ~:1s provided by

Hobert J. Hurley~ DirL~t..:.tor, Moss Landing :.1arinc Laboratori,_:_·s.

A great dc3l of thanks is owed to the intrcpiJ capt~ins 2nd crc~es of H/V ST-908 [dubbed ",\rtemia"), eo;pcciallv John

Snodgrass, Jay ~1ik~clian, Rjch~rJ Keeney, Robert Cayce and all hands aboard.

~lany indivldtJals pro\·idcd information, advice and encouragement throughout the sttidy. I wotrld especiall; like to thank my gr~duatc coJllmittcc, Gregor M. Cuillict, G. Victor

v:ii Morejohn, James W. Nybakken and John E. ~cCos for Lhe-ir interest. Alex E. Peden, Marine Biology Division, British Colambia Provincial Museum, gave much advice on records and systematics, as did Shelly R. Johnson oi rte University of Southern California. John E. Fitch, Calif- ornia Department of Fish and Game provided catch data and helped ~ith otolith age determination in uLar~s. Warren C. Freihofer, California Acad0my o~ Sci- cnces, prep ecimcns for study of the cial nerves. Earl E. Krygier, Oregon State University, and Peter N. Slattery, Moss Landing Marine Laboratories, identified many of the crustaceuns sampled. Milton S. Love, Universit)· of California, Santa Barbara, identified two species of parasites. Robert A. Bchrstock, Humboldt State University, provided specinaens of ~s from \orthcrn California.

Catherine DiNonaco il1ustrated the fish fi res.

Collections were examined, a ice given and T3dio- graphs ~:tkcn with the kind help of: Victor G. ingcr~

Da~iel ~~- Cohen and ~at Gratnblin, U. S. \ational ~use11~ of Nattlrnl l·iistory; Rohert .J. L~lVenllcrg, C:1m1~ C. Swifr nnd

Jerry W. ~~cumann, Los Ang(~les County ~·1uscum of :\atuTal

I·!istory·; Richard II. 1::_oscnb1Jtt, Carl L. 1 r)s and .Joseph

F. Copp, Scrjpps lnstittlticn of Oce3nogr hy; ~illia~ G.

[l•"'':''rL-~" ''Cl~,-,. 1 ..... ~~'- .", ('l'nnon>-1 '-b ···C:t_..+;:. "'~"' iln1"-• -··. _-.._.1: t"'.r, · '1'1.1r ..~. 1" u1·•m ,,;,,. Eschmo\·er~. e,· , l··"· I •

Follett, Warren C. Frciho r, Tomio Iwamoto, Lillian J. 1 '~ of Ichthyology, California Academy of Sciences. Finally, my deepest appreciation goes to a special tea~ who insured the success of this project, esp~ciall~

BrookeS. Antrim, Edwin K. Osucla, Donald M. Balt:, C:a;·,· F..

~lcDonald, Gregor 1-J. Cailliet, Richard G. 1\liever, o'an·; L. Stevenson, David A. Ambrose and Vic E. Brcc n, .Jr. who helped ~c develop the nerve-wracking art of midwatcr tra~l ing in submarine canyons. TABLE OF CO~TE~TS

ACK:\0\iLEDG~tENTS •• , , . • . • ...... • . • . . . • . • • ...... ,. i i

LIST OF TABLES .....•..•...... •.•...•...•......

LIS"' OF FIGURES...... \:i i

E:\TRODUCT I 0:\ .....•.•...... •.•...•......

MATERIALS A"JD ~!ETHODS .•...... •.....•..•....••.••...

RESULTS •.•...•.•...... •...... •..•......

ternatics ...•...... ll

f•li!terial Examined ..•...... •..•...... •...•.

Di.ngnosis ...... •...... •.... lS

~'leristics and i>iorphometrics~ ..... _ ......

Description .•.. , ...... •......

ltet~:a rks •...... ~. ~ ...... ~ ~., ~ ...... ~ ... ~

I\ a tural History ...... ~~ ......

Distribution ...... e an(J G:rot,rth ......

RcproJuction ......

Food IL1bi ts ......

Parasites and Predators ...... , ......

Aquarium Obscrv~!t ns...... '.

DlSCLJSSJl)~ ...... ~ ......

LlTL;L\TURE CITED .•.•...... •.•...•...... LIST OF TABLES

Table

1 .. ~loristics and morphometries of Ly :ts species in California waters, comparing IJc

material to recent (SAMMP) collectio"s. Measurements are percent standard length

except premaxillary teeth lengths and S~

(in mm) ...... •...... •.

2. Numbers of ripe and unripe adttlt

s sampled e:.:tch month .. ~ .....•.... 3. Species of midwator organisms identified

taken in Monterey Bay ...... ~··· ...... J. ..

AppenJiccs

A. Tr~1wl data of the Steir1l1art Aqtlariu:n

\1iCtvater :-ia.intcnancc Progr

B. Partial list of ·anillt~1s caugl1t in the

~lontcrc)· Ray are3 by miJwiltcr ti·awls fished

off the l~otton:, 1972-1076, in water de cr

than 200 Ill.~-···~······ ...... s:,

Xl Ll ST OF FIGURES

1. p of Monterey Bay, California, showing the

t1~o trawling areas. Canyon contours are 1n

fathoms ......

Caudal skeleton of eodapua maniib1tla1ais

sho1.:ing ultimate rays of vcTtical fins .. ~ ... ~ .... J6

pua me_ ·z~bulcu"it: Gilbert 1915. Upper,.

male, 140 rum SL; lower, m~le, 149 mrn SL, fr~m

SAl'·H,!P tow 70, Monterey Bar.~~··~···· ...... 47

4. He::1d of ~?aPia illustrating

cephalic pores and free lateralis orga~s. AbhreviatiotiS arc: AP, articular pore; lOP,

in~erorbit31 pore; ~JP, mandlhular pore~; ~P,

1 nasal pore; roP, postocular pore; P1 ) preoperc~l~T

pores; PrP, preoCtJlar pore .....•......

192 nt!:l SL ...... ::;_._-"

6. Regression of male 3Jld female prc~laxil1nry teeth

lengths in Pre-maxillary

length of 8 mm corresponds to a standard !c t~

(S;..) of about 120 mm, ll m;n to about 160 r~m SL

r both SCXL'~S ...... ~ ••• ~. • • • • • • • • • • 5,!)

7.. Map of the eastern ~orth Pacific showing coll-

ection loc3lities of Ly~~o b14 2'7_..$ •••••

X j i Fig:.Jre :~o. Page:

8. Distribution of vertebral counts in Ly ~-·• from California waters. Sample

s1=e 1n parentheses; single records are circl

Graphics after Hubbs and Hubbs, 1953...... ::~

9. Vertical distribution of Lycodapus ma (left on

[right) 1n Monterey Canyon, day and night. A)

area 1, B) area 2 [see fig. 1). :\umber in parentheses sample size. Greatest bottom depth in area 1 about 700 m; greatest bottom d th in area 2 about 1400 m...... 53

10. Vertical distribution of pelagic zoarci in

Monterey Canyon. A) codapus ma1~dib:rZaris and Sample si:e in each

~epth interval in parentheses; day on left, night following. Catch per hour mirrored ......

11. Vertical distribution of zooplankton found as

prey of

Sergestes spp., D) P~s ~ae~ spp., E) n:ysids and

F) ConcitiJD OHf!;r •• N...... JJ

'""-..:.... 0 r

~on~ere~ B~y as clctcrmined b~ otolith readings.

Right females; left, males. S;Jmple si e>

parentheses ...... •.

xiii Figure :\o.

13. Length frequency histograms of Lycodapua mandibklaris from Monterey Bay. A) October

to Mnrch, cJ=298, B) April to September, ".>261. .. 59

from September to April. Sample size in

parentheses...... tO 15. Monthly gonad indices for adult male mandibularis. Vertical line and end bars represent monthly ranges, horizontal line the

mean and dark vertical bar the 95~ confidence

e rv a 1 ..... * • • • • • • • • • • • • • • • • • ..... ~ ... ~ • ~ •• ~ • • • • • 6 1

16. Monthly gonad indices for adult fe~alc Lya us

Graphics :tftc•r fig. 15 ...... 17. Index of Relative Importance plot of stomach

items 10 juvenile (less than about 110 mm SL)

18. InJcx of Relative l~portancc plot of stomacl1

itc•ms 111 adult

19. Categories of stomJch contents condi:iort (aft r

DC'\'(i tt o l'.:11 . 1 1 lCt,. . 1 'l-~1-') ...... ~~4···~···~···•·· 6S 20, Fullness and digestion state category histogr·ams for

N= 3 8 •••.•••••••••••.•••.•••...... ~ . . . .. ' ...... 66 21. Day-night chronol y matrices (see fig. 19) of

stomach contents in ~odapus il•·,Iaris. :dv Figure No.

Dark bars, night sa~ples; light bars, day samples. Sample size: Day recent, N=86, not recent, N=129; night recent, N=l7, not recent,

:-.'=21 ....•...... •...... •...... 67

22. Lyao us mandibula1~is: lateral view of head showing branches of facial nerve VII that innervate the free lateralis organs and canals.

General cutaneous branches not shown ...... 68

XV INTRODUCTION

The biology of many deep-sea fishes ~n C~lifc-nia waters 1s not well known, although collecting expedit n~ since the late nineteenth century have provided numerous specimens of most forms. Studies of tJ1c :lbundant :oarcic3e [eelpouts) in the eastern North Pacific are few.

ma bulavis 3 a pelagic, nearshore eelpout has received no attention in midwater commtanity stt1dics a11d ina quato treatment in taxonomic accotlnts.

Literattire on the zoarcids of the eastern IJacif~c is generally taxonomic, but recently so~e ecological st~J- ies have been completed. Levings (1967, 1069) exn~ined the life history of co:iops~r-r:. . pce1--;'L::..-o. . ·-. 111. c.onao18n .. 1'/at('f.S.

Klicver (1976] simi.larly studied ~ayn0~ ~a

Monterey Bay. Taxor:omic revieti/S of :o3rcids cons:sr of

Bayliff's (l9S4) anJ Grinols's (1%5) studies of s kno~n ~Lt the time, ;J1thot:glt these :luthors can

tions, Gosli.ne (19C.,8) and Freihoier (1970) used in their d:scussions of "hi.~:)lf'Tlt :oarcid affinities. Ot !"' n1ujo1· easte1·n l'acific :o~rcid st~Jies irl~luJc McAllistc~ and Rees 1 s (1964) revision of MelanostiJ~J ~ YarbcrT)''s

Short notes and descriptions coln]10S2 the rcm:1indcr of the litcrattJrc

ar.d Eor:d, 195~1; Ba)·li.ff, :9~~9; Grinols, 1966; Sc}:ult:. 19fi7; Gotshall, 1971; Peden, 1973). Andriashcv (1954; a Schmidt (1950) provide excellent bibliographies af tjc eelpouts of the Soviet seas, some of whic1: are also fou 1n eastern Pacific waters.

codapus mandibuZaPis populations arc known o~~l~· from Pacific coast submarine canyons, Puget Sound the

~eep water inlets of British Columbia and Alaska. These canyons appear to harbor uni

Barnard, 1966, 1967) and some fish stt•dies (Pereyra et al.,

1969; Osada and Cailliet, 1975; Kliever, 1976; Cailliet and Lea, 1977) constitute the biological literature on Pacific canyons. Geological investigations include

Shepard et al. (1974) and Martin and Emery (1967), which provide excellent bibliographies on related phy5ical studies~ In ~tontere)· Canyon, Galliher (1932) 3nd .J~nscrl (1976) studied the sediments and Caster l19fi9) mcast1red bottom currents.

~lidwatcr trawl sLervcys off tJ1e ~orth AmerJcan

Pacific continPntal slope and CHlifornia Ctlrrent areas have failed to prodtiCC spcci1:1ens of L. n:J~iit~Zaris

(Ahlstrom, 1969; Aron, 1959, 1:162; Bcr·r)· and Perkins~

1966; King Cind l\·erson, 1962; P"ic:pcr, l~ft17).

Previous Monter~y Bny studies also lacked rc~c s

C)f :.,. t;;ond::Z;;i L:L·•:' s (1\ught.ry, 195:1; Harh~m, 1~157; F3::;t,

19t0) althotigh several yot~ the R/V Tage investigations of Hopkins Marine Station.

Heimann [1963) and Best and S:nith (1965) d not r-:port the species from bottom tra1d investigations MonTerey

Bay but net mesh size in these studies did nat favor cnp ture of this diminuitive eelpout.

Surveys in southern California basins provided specimens of us fierasfer only when the tra~ls neared bottom (Lavenberg and Ebeling, 1967; Ebeling e: al.,

1970). Unreport in these papers (pars. serv.) I< ere specimens of £. tinus collected primarily by the R/V Velcro IV cruises of the University of Southern California.

Off Oregon, Pearcy and Laurs (1966) and Pearcy and

~lesecar (1971) did not report L. ma the deep scattering layer but the species was recorded in

Pe~rcy's (1964, 1972) lists from midwater trawling in both offshore and nearshore waters. Those collections were from Astoria Canyon in 200 meters.

Taylor's f1967~1) midv.·atcr stlrvey rc·ported ' 1 unidcr:t-

om Queen C11arlotto Sound nenr known ~ .. m.!n3ibularis collection sit~s but l1e did net report eelpours off the British Columbia continental slope (Taylor, l9G7h,

1968).

The Steinhart Aquarium miJwater collection has provided significant material enabling this and future con tributions to tl1e natural history of submarine canyon-dwell- ing fishes. This study was 11ndertaken to elucld3te the systematics and biology of Lyco marzdibularis. ledge of the food and feeding habits of this species, the detailing of its reproductive biology and growth para meters, as well as morphological characters, provides a framework from which the evolution and relationships of the genus may be discussed .. 1-lATERIALS AolD METHODS

Samples used in the natural history st ~ ~cr:

taken from Monterey Bay during the bimonthly Steinhart

Aquarium }r1irh~·ater .Maintenance Program (SA~L\lP) cruise-s \~ ~ r;~,

a modified, 1.8 m Tucker trawl (Tucker, 1951) outfitt

with a messenger operated, double-release mechanism l!iopkins

et al., 1973) and a flow-throttgh, canvas cod-end bag to

netting was made of 9 rnm stretch mesh knotless nylon wit!"l

a 1 rn wide cod end liner of S mrn square mesh. The net was

towed at abo11t two knots from the R/V ST 908, a 16.5 m

converted harbor tug. Two to four one hour tows per cruise

resulted in 56 discrete depth samples grouped ln 100 m

intervals from the surface to 800 m (Appendix A). Addi

tional specimens were obtained from seven one-half l1our

To\;s in the upper 100 m and open net hauls. Tow dc·pth h.c'

dcterltli11ed by reading the disc charts from a Benthos ·rime

Depth Recorder, model 1170. A total of 604 spocirnons of

ti011S in Monterey Canyon between 4 ~tnrch 1974 and 23 J~~

uary 1976 lfig. 1)~ Most other organisms were identified

to the lowest possible taxa anJ :• pa!ttal list of those

species is prc.::C

food and r tivc studies were )Jrese!·vcd on shipboar~

in 10 formalin and later transferred gradually to 70S

5 6 isopropyl alcohol. Living specimens were returned to the laboratory in darkened containers and maintained in a cold water aquarium (see Robison, 1973; McCosker and Anderson, 1976).

Specimens of L. mandibularis used for systematic analysis were collected only in California waters. Some were radiographed and cleared and stained by the alizarin method (Taylor, 1967) for meristic counts and features of the caudal skeleton. Measurements were taken with dial calipers and an ocular micrometer and recorded to the ncar· est 0.1 mm. Morphometric definitions follow Hubbs and Lagler (1958) with the following exceptions and additions. Measurements including the tip of the snout were measured from the anterior margin of the premaxillary symphysis; orbit diameter and interorbital width are the least bony distance; body depth was measured in a straight line from the anal fin origin to the base of the dorsal fin; pectoral length refers to the length of the longest ray; preanal length was measured from the anal fin origin to the tip of the lower jaw; length of gill slit above pectoral base was measured from the dorsal edge of the first pectoral ray to the dorsal edge of the opercular opening; fourth gill raker ratio refers to the length of the fourth raker on the first arch divided by the distance between the ventral base of the fourth raker and the ventral base of the fifth raker; premaxillary teeth lengths given are an avera of sever::tl of the anteriormost teeth measured in a straight line from the tooth tip to the base of the premaxilla; dorsal and anal fin ray counts (taken from cleared and stained and radiographed specimens) include the last two r::tys not at tached to the hypurals as one, as is customary when they e1canate from a common base (fig. 2); caudal fin rays, then, are only those attached to the hypurals. Since considerable confusion exists in the liter ature on Lycodapus, specimens from all Cali rnia localities were cornpar to the type material deposited :.n the National Museum of Natural flistory, Washington, D.C. and tho California Academy of Sciences, San Fr3nclsco. Institutional abbreviations used in the systematic part are as follows: USNM, Division of Fishes, National Museum of Natural History, Washington, D.C.; BCP~!, ~!arine

Biology Division, British Columbia Provincial ~!useum, Victoria, B.C.; i!SC, School of Fisheries and Wildlife, Humboldt State UniversitY, Arcata, California; CAS, Department of Ichthyology, California Acadmey of Sciences; SU, Stanford University Mttsettm of Natural •listory (now housed at CAS); ~1D1L, Ichthyology collection, ~loss Land ing Marine Laboratories, Moss Landing; LACM, Se~tion of Ichthyology, Natural History Museum of Los Angeles County, Los Angeles; SIO, Marine Vertebrates Collection, Scripps 8 Institution of Oceanography, La Jolla.

The vertical distribution of L. bular was estimated by the use of the closing net. Specimens om tows with gear failures were not included in this a~alvsis.

Lacking a flow meter, the exact fishing a ct of the t:"tn,:l cannot be determined, but all trawls were set, recovered and towed for the duration with minimum variation. Xumbers of L. m2ndibularisJ Mc!a~os~igma pammelas and cephalopods collected per hour of trawling were calculated on a catch per effort basis by plotting the average number taken per hour of trawling in each 100 m interval divided by th~ total number of tows, 63. Catch per effort for crustaceans later determined to be prey of L. mandibularis was similarly plotted except that blotted dry volumes of these organisms were measured, In the laboratory, fresh specimens (mostly those that died in captivity) were examined for age utilizing the annuli on otolith (sagitta) pairs. Becnuse of the poor sample si:es in the summers of 1974 and 1975, fish were selected for aging from the "1vintcr" months only (October to ~!arch). Only otoliths Hith hyaline, or "1·:inter," nuclei were used for age estimati011. l.cngth frequencies constructed from the n\\1 inter" and nsummeru (April to tember) catches were plotted to compare modes corresponding to age groups. Seasonal changes in fish lengths were correlated Cl

\"'ith annulus formation in L. mand·Z:bularis, as discussed by

Dark (1975) for hake and Gregory and Jaw [1976) fer petrale sole, so otoliths were deemed good estimators of age. Preserved specimens were measured, sexed and 0xamin- eel internally. Gonads were dissected, excess water an~ mesentery removed and weighed to the nearest 0.0001 gram on an electronic analytical balance. Gonad indices were calculated according to the formula:

wt. gonad x 10 G. I.~ 10 3 ( S L) ad~1ptcd from an expression 1n general usage (Scl1ncffcr snd

Orange, 1956; Shiokawa, 1962; Moser, 1907; Batts, 10-~i.

These indices were plotted against time to detec~ any spah'ning season. Egg sizes were meastired with an octtlnr micrometer to the nearest 0.1 mm. Egg si:e classes here then plotted ag:tinst standard length for adult femal~s collected during peak spawning months. Parasites were counted and identified to spec1es and their points of attachment were noted.

Stoinach contents were identified to the lo~est taxa possible. The importance of each food category was indicated by plotting the Index of Relative Importance (IRI) for juvenill'S and adults separately (Pinkas et al.,

1971). This formula was constructed so that no single

•:;,:asurc biases the importance cesti.m~te [volumetric, numerical and frcqt•cncy calculations]. The degree of lJ digestion and stomach fullness were subject ly scored

(after DeWitt and Cailliet, 1972) to g an indication of feeding chronol Categories scored were: 0, empty; 1, 1/4 full; 2, 1/2 full; ~. 3/4 iull;

4, full, and for digestion state: 0, empty 1, very digested, nothing recognizable; 2, sonle digestion, parts recognizable to higher taxa; 3, little digestion, much recognizable; 4, intact items, readily i tttifiable. To discover where and when L. ma ibularia fed, a two-by- two contingency table was constructed of the above cat- egor1es and histograms of each matrix category were plotted against time. Lively specimens were selected from the catch buckets and placed in darkened containers cooled tc

~;-10 °C Wlt.. h trozen - sea water on ten occasions (McCosker and Anderson, 1976). They were returned to the labor&- tory and placed in a refrigerated, circulati aquarlun and subjected to temperature and illti1tinllt~cn studies to observe the various species rc~ctions anJ determine, if possible, optimol levels and toler&IICCS. l.ongevity and activity were the subjects of these experiments. RESULTS

SYSTHIATICS.

All of the eastern North Pacific Ly~od~P~d species were described by Charles H. Gilbert of StNnford Lniversiry from material dredged by the U. S. Fish Commission stea~er

Albatross. The generic type, Ly~odapus fie ?· Gilb~r~

1891, was described from a syntypic series of spcci~ens from the Washington coast, southern California and the

Gulf of California. In 1895 Gilbert described two species from the eastern Berirrg Sea, L • .zxt.::nD:J.D Hnd L. :_,:!"£)·>~-'t:'P8:~ and L. dermatinus from the central :nlifornia coast. I11

1915 L~ mandl 7-art~s, L. Zyco n and !.:. att·enu.~:t:.. u:; h:·erc named from Monterey Bay and L. grossii~f.3 from th0 Perin~

Sea and southeast Alaska. Considerable t:1xono:nic C(•ct1:Jsi T• exists in the literature concerning thorough examination of the type material plus matty recc~t ly caught, well-preserved specimens, 2 brief generic over view is included here pending a more comprehensive study.

c us ftcraa r is known from the Bering Sea to the Gulf of Panam3 and is also present in the Gt1lf or

California. Confusion in the litc1·ature with L. ~~ is has been the result of ;1 failure to recognize the dif ferences in hend pore patterns dnd gill ra}:cr leng~hs.

One syntypc from :he Gulf of CaliforJJin (S!J 845) and nine from the COJSt of Nahington (US_\H 43094, thre-e specimc:15-;

11 US(~}-1 53063, SiX Specimens OTiginally identi-fied .~s M·ljr.r::(:. br1 un;--:.

Lycodapua extensus is known only from the holotype

[US''asfer or L. rjltgini Andriashev

1935, although the dorsal fin ray count (stated to be 96} is incorrect. Examination of the type revealed such a thorough state of destruction that until more material lS taken from the type locality the status of no~en dubium should apply to this name.

Lycod~pus parviceps l1ns been kilOWn, ttntil recentl)·, only from the holotype. Recent c:1pturcs arot111d Vancouver

Island, British Coltttnbia, su~gest this speci~s lives in a relatively shallow midwatcr habitat (Alex Peden, pers. comm.). The type specimen CU:->NM 4.3631) is in identifiable condition.

c J''!i·.z>t:/nus is knoHn from northern B~1j a

California to the Washingto11 coast 3ild Hering Sea. It is also present ic the Gulf of California. The type (US~M

53035) is in identifiable condition. There has been ~- l.J

confusion of this species with L. mandibularis 1n the literature.

Lycodapus grossidens has caused a grc3t deal of

confusion in the literature. The holotype (USNM 75824)

is identical to L. fierasfer which was described 24 vcars

earlier. The five paratypes, however, (SU 25634, two spec-

imens; USNM 135596, three specimens) are identical to Mont- I erey Bay specinlens of L. mandibularis. The conftlsion has I arisen from the variability in head pore patterns in L. fierasfer~ the failure to recognize the differences in I! I gill raker lengths, coloration and the variability 1n r tooth sizes with age in the two species. I Two extreme morphs of Lycodapus mandibuZariz were ! given different names from the Albatross Monterey Bay t- material (Gilbert, 1915). These specimens, L. ljeodcr:

and L. a:ttenuatus, exhibit sexual dimorphism typical of

- , the species. A ripe male was designated L. Lyt:odon

("wolf tooth") as it possessed large jdw teeth, thid:er,

darker skin and a [latter head than most fem:•lcs. The

holotype of the morph named L. attenaut;•s is Rlt emaciated,

5flcnt female which gives the upjlearai1Ce of possessing

higher 1neristic counts tl1an other species alt!1ough jt

does not. It differs from male L. ~Jt1·i:b~Zaris of the

same s1:c in h3ving a thinner body and tail, smaller teeth

and lighter color (t3ble 1). l.. -r

Other species of eodapus include L. azts:r·a:~s

Norman 1937 from the Straits of Magellan, L. derJ ''~ Andriashev 1935 from the Bering Sea off Kamchatka and

L. microchir Schmidt 1950 from the Sea of Okhotsk. Two undescribed species from very deep water have been collected from the South Atlantic [Shelly Johnson, pers. comm.) and from the eastern North Pacific (including one from California), also from great depths (Peden and Anderson, MS}. The exact status of the Soviet species is unknown to me though L. derjugini appears to be valid.

L'fCODAPUS MMvDIBULARIS Gilbert 1915 Pallid eelpout

Figs. 2,3,4,5

us mandibular[& Gilbert 1915, p. 305, 369-371, pl. 20, ig.20 (hoJotypc by original designation: IJS~M 75~:3, Monterey 83y, C3lifornia, Albatross sra. 4533; beam t~a~l in 262-533 m.

Ly~o us man;iib:tlaris Gilbert. JorJ~In, Ivcrn1ann and Cla1·k 193 , p. 479; l!ubbs 1941, p. 17; Clc•cu;;ns and \Vilby 1"19, p. 19(1, fig.l33; Bohlke 195:1, p.102; \\ilimovskv 195,1, p. 238; Bayliff 1954, pp. ii lv~ 122-127, 15"1, 173, tab2c 35, pls.Vll,lX; WiUmovsb· 1958, p. 93; B:.ry1iff 1959, pp. -9- 80; Bailey et al. 1960, p. 44; Clc~cns and Wilby 19rl, p. 391, fig.273; Pearcy 1'164, p. 87; Grinols 1965, pp. ,13- lGO; Schultz 1967, p. 4; Fitch and Lavenberg 1968, p. ]l2; Hail et al. 1970, p. 29; l'carcv :1~172, p. 362; Q:1ast and Hall 1972, p. 14; Kuko~Yski l•172,"pp. 11, 27; Han: 1973, p. 241. u:i us r,and.I-but:zP·[;::--:. ;\lver.son 1951, p. 86; Alton 1972, p. 0; DeLacy et ~1. 197:, p. 9; ~iller and Lea 1972, pp. 78-79; Lindberg 19~~. p. ~04, fig.787.

acaan:~.:; a::tenu .. :c:.:.,s Gi1Dert 1915, pp. 372, .':1;,.:,, pl.~l~.;: g. ordan, Evermann and Clark 1930, p. 479; Schult: 19 7, 1:: p. 4; Fitch and Lavenberg 1968, p. 142; Kukowsky 19-2, pp. 11, 27.

Lue tiS ly n Gilbert 1915, pp. 371, 372, p1.21, fi,~. 21; ordan, Evermann and Clark 1930, p. 179; Sc 1t: 1967, p. 4; Fitch and Lavenberg 1968, p. 142; Kukowsk 19 7 2 ' pp . 11 ' 2 7 .

Ly~o us grossidens Gilbert 1915, pp. 371, 373, 374 -a parte, non holotype; paratypes USNN 135596 and SU 25 34, Funter Bay, southeast Alaska, Albatross sta. 4257; heam trawl in 640 rn).

Lye pus gJ•oasiden2 Gilbert [pro parte). Jordan, Ever mann and Clark 1930, p. 479; Andriashev 19~5, p. 422; Amlrioshcv 1939, pp. 51, 82; Bi:ihlkiii· movsk~· 1954, p. 288; liff 1954, pp. ii-iv, 116-121, 126, 154, 177, table 3 , pls.VII,IX; R~yliff 1959, pp. 79, 80; Quast and llall 1972, p. 14.

us fieraefer [pro partel. Schult: and DeLacy 1936, p. 2; Bayliff 1954, pp. 112, 114~116, 153, 177,,tt:bie 33; Bayliff 1959, p. 79.

ccdaD••s ~ran r (rnisiJcntification). Lv~rrnann and Goldsborough 1907, p. 3~5; Hubbs and Schultz 1932, p, 323; Clemens 1933, p. 61; Clemens and Kilby 1946, p. 195, 196; Clen1cns and Wilby 1961, p. 391; llart 1973, p. 239,

Eothracara mollie (misidentification). Chapman 1940, pp. 39-40.

Naterial Examined

Measurements giv~n are sta11dard lengths. Lye. :zpus ma tula1•£s: HOLOTYPE, US\M 75823 [159.3 mm today); Mont-

.~.6°4.0.'" 1 0 crey B·ay, Ca l 1'f orn1a,. ;\'1! Jatross sta. ~j3J,,--- , . L ~ 1"~- Ob.O'W, 28 May 1904; beam tra~l in 262-533 m. PARATYPES:

0 7 MoJ1°~l-n\' USXMA. 14G- .) 14 ('- ' 10'-' .. -' , 1l6' ' .... 1•m)·,ll f ) • L .... .,_ ,{ Bay, Albatross sta 4~61, 36°46.6'N 1Zl 0 53.5'W, 12 May 1904; beam trawl in

519-650 m. SU 25765 [5, 49.0-123.3 mm); Monterey Bav, AI· 16 trawl in 58(?)-562 m. su 22990 (1' 102.1 mm}; c•!onterey

3~ 0 47.~''.' -,~cor Bsy, "'lb a t ross sta. 45-q_.,_, o _ ~ ·' ~~~ 1.~"'\'. ,, .L-,

1904; beam trm-:1 in 846-1108 m. ~OX-TYPE \LATERIAL:

US\M 118118 (1,87.5 mm); Dixon Entrance, British Colusbi3,

International Fish Commission sta. 109A, ~4°19'N 133°13'~. 9 March 1939; midwater trawl, 215-100 m.w.o. HSC 29001-20 (1, 166.3 mm); Eel Canyon off Cape Mendocino, California,

40°53.2·~ Pacific R3iJcr sta. 7 124°33.7'\V, 21 Octcber 19'""'::; otter trawl in 546 m. CAS 3S991 (1, 88.0 mm); Vizcaino

C:lnyon off Cape Mendocino, N.B. Scofield cr. SOIl 88, 39'0

Sl'N 124°06'W, 12 October 1950; beam trawl in 155-lfS m.

BCPM 74-511 (36, 86.0-16~.1 mml; Monterey Canyon, ST·9US cr.~ 36°47~0'N 121°55.srw, :20 Fcbruury 197--1; l.S m Tucker trawl, 0-500 m. LACM 32009-1 (77, 85.1-129.6 wn

0 Canyon) C"a l .l·r· c rn 1· a , ,,'l as k·a s t a . "11 "' _,-- :L~, , ,,~A • t)'. :, • 0'\•.

1~.3'~, 20 April 1971; 15.1 m midwater trawl, 0 ~5 ~~-

LACM 30797-8 (1, 160.1, 176.0 1nm]; l~a Jollu C3n)·~n, l if- ornia, ~.B. Scofield sta. 69 S 5-26, 3

10 August 1969; otter trawl in 564 10. L.\C~! 3 33-1 (.!2~

112.0-1~1.0 mm); Monterey Canvon, Alaska sta. 7 2 A 7-~o.

6 S tembcr 1972; midwate1· tr~twl in :36 m. Uncatalo

mrn); Bering Sea, vic. Pribilof Island~ \lbatrcss sta.

3483, 57°18.0'\ 171°18.0'W, 12 July lN93; beam tr~wl in

101m. US\M 135~96 (3, 91.0-97.0 mml; Lynn (3nal, ~ic. "'- 7 L.

FUJJter Bay, southeast Alaska, Albatross Sta. 4257, 58~

14.1'N 134°57.3'W, 23 July 1903; beam trawl in 63? m.

SU 25634 (2, 9~.8, 94.4 mm); Albatross sta. 4257 [as above). USNM 60109 (1, 90+ mm); Boca de Quadra, south~~s~

1903; beam trawl in 271· 329 Til. USNl-i 60626 (1, 104.,. ;am·,; Eastern Passage, southeast Alaska, Albatross sta. 4250,

7 0 56°'6.:...... 'N! 13'.:. 1-:J. 4· •~u, 1-.:> J uy1 190-·.:)} beam trawl in 111-120 m. USNM 150311 (7, 72.3-126.2 mm); v1c. Un3laska Island,

7 f\_"l·1·ka ._ ~ , Alb a tr o s s s t·a . --~L'L:. '--"' , ::>:;--o-- .:>,) • 8'''~ ;\: ust 1890; beam trawl in 199 m. L'i·: HOLOTYPE, USNM 49638 (83+ mm); Albatross sta. 3324 (as

~.;.~""'-·' abo- "e]v-.• DliCQ~apua' " ar·t 2nU~tuD: ]]'}l.t'.<'':"v[)r::,~- A! L .,_.,,_-,",'.'•'•- :\ -•..• •u

(149.5 mm); Monterey Bay, Albatross sta. 44bl, 36°~6.6'\ 121°53.5'W, 12 May 1904; beam trawl in 519 bSO m. dc:.p us l y co l!OLOTYPE, US\:•! 758c2 (137. 5 m;.,]; ~!o:Jtc:rcv

Bay, Albatross sta. 4509, 36°46.5'~ 121°5S.5'W, 20 May

1904; beam trawl in 277-521 m. Lba;dao~s

SYNTYPES, US\M 43094 (3, 93.0-124.0 mm); Washington cGast,

·to "0~.~2, r"f\7 (\t\11 ;o .J.t1n_n .].~89' ~'"1.u.-l.·']'""tl'C-· ;::,::, ~a. J _ •1707_8'" ...... 0' .....1''"01-\ .:J .:..-..~- 't ~"'' " 1 ''- •· ' beam trawl in 1069 m. Sll 845 (135.0 mn); Guaymas Basin,

~ , f - 11 0 -,..., o ; ... 8 r ,. ~ 1 1 a C ul f or Calr o2·n.ia, Albatross sta ..>1.. , '-' ~-" .• L,

2S.O'W, 20 ~~~rch 16S9, beam trawl in 1839 m. SIO : 4-l t1t)

1 t,'4 6'l-·-- q 1''.;,.J."T !"J")'od ~ t.·"alifoJ·nr'· _._ :; Li L·o-,-• u~ ... , 1"\.~.-l:.''>,<,.,!'-~'--'J·,v·"r·•'·"l"

~~ 0 sa.t 1- 15 , u"1T 1"... ,. "'j:> ...'(" J."~-~ ",,. .. J• ...'1°"5 .)" ••Cl'l•' , 1 Aprl . ·1 1974; otter trawl in 901 m. CAS 35989 (1, 135.1 mm); Nonterev "nay~ ST - 908 cr., -6°46j ~ 7'N~· 171°-o..... ::J~,~:J - ":;'i, _7? ... <::~ep t er:J·c·r... ·b 'o-'~---,_,., ·

1.8 m Tucker trawl on bottom 10-20 min., n-769 ~- ~l\IL

DB-45(7414) (1, 110.0 mm); ~lontercy Bay, USNS De Str ..,ucr

2.1 m McDuck beam trawl in 1197-1409 m. LJco adnus: flOLOTYPE, US:>JM 53035 (109. 0 mm}; Farallon Islar,ds,

California, Albatross sta. 3162, 37°54.~'N 123°30.0'W,

22 March lSB9; beam trawl in 1005 m. SU 5452 (catalogued

(:I, 78.0-107.0 mm); off Pt. Arena, Calif-

tcmber 1890; beam trawl in 828 m.

(housed at CAS) (1, 104.6 mm); off Washington co~L't, A1Lo-

beam trawl in 939 m. CAS 31493 (2, 110.5, 135.1 mm); ~:unt-

ST-908 cr.,

197.:i; 1.8 m Tucker trah'l, U-769 m .. CAS 35990 (1, 1:n.s ~J1);

1.8 m Tucker trawl, 0-580 m, bottom 878-988 m. MUlL \n·:-113

(7417) [1, 91.6 mm); ST-908 cr. as CAS 35990.

::: crpu.s mar; b:tla s can distinguished t~o::1 2.11 other species in the genus by the fcl10\''~:lg comb1 '13.t iD<1 ci characters: gill rakers 5hort, thos0 on the first arch BOt

ous, 15-17+72-80 RS-97; ~ ~ingle interorbital la~eralis pore, 19 m.'lndibular pores 3, articular pore 1, preopcrcul:;r res

4; vomerine teeth 1-5 (usually 2 or 3); posterior m1rgins of dorsal and anal fins become black in older spec ens; jaws pearl-white, lining of mouth and gill cavi dus

tries Counts and measurements were studied to detect specific, allometric and sexual differences in species. Table 1 con1pares the type material of California

eo us to the recent SAMMP specimens. Although growth related proportions are evident, no significant differences warrant the separation of extreme morphs from L. ~c b- uZ.c::"ia. Counts ar.d proportions of the holotype of "' .: sidens are close to L. fierasfeP with the possible cxcc11- tion of the ir.terorbital pores. In L. fierJsfer each pore lies mediad the dorsnl midline or.d above the ont:~ls.

1.;as able to discern this condition in the type of L. ic', ,:-

B7 na. although the skin over the right half of the hcaJ is badly ripped and a second pore, if prescr.t, ca:1not be seen. L. ma1: bul~ris generally h~s higher counts tll3n either of tl1ese nonainal species (except for the paratypcs

a nus differs from both valid species primarily 111 Its loHcr counts, pore patterns ur:.d coloration. Recent specl- mens from California and Oregon support the data taken from tl1e holot e. ' } Description

Body and tail very elongate, compressed, tap :in; posteriorly to a blunt tip. Skin thin and transparent except in some larger adults in which it thickens, becoming opaque 1n preservative. Spines, scales, and pelvic fins absent. Dorsal and anal fins confluent with caudal, the posterior rays coming about half as thick as the anteriormost rays. Pectoral fin shorter than orbit diameter in juveniles less than about 80 mm SL, becoming about tw~~o the orbit diameter in larger adults. Dorsal profile of head generally concave in younger individuals, becoming more evenly rounded with age. Lower jaw slightly projecting beyond upper, ending in a noticeable symphyseal knob. but reduced in the largest adults which have a more evenly rounded head. Mouth sloping ventrally to middle of eve with slig!1t convex tilt ~lnteriorly. VomeTine and pal:t;;ir:·:: teeth in a single series. Palatine teeth weak, rarel)' absent, often not projecting through skin. Vonterine teeth recurved :1nd more prodt:ced in adult males th3ll fer;;a!es.

Premaxillary and dent.3ry teeth in three ill defined T0\-'5 anteriorly, blending into a single r01v of s;naller tcceth in poste1·ior half 0f dcntary 811J a do~~ble row on Jnaxilla.

The formul3 of tl1e cephalic li•tcralis pore pattern is a distinguishing character in this gent1s. Tn L. ma~ii~:,Za?i~ the preopcrculom3ndlbuletr pore series (PMP') consls~s of three pores over the dentary, one over the art lar and cl four over the preoperculum giving the rmula A single interorbital, nasal, prcocular ard postocul2r pore (fig. 4). Smaller free lateralis orga~s are not connected by a canal and run 1n a single series adjace~t to larger pores, below eyes and on nape. circular, protruding into dorsal profile and outline but recedi11g somewhat in preservative. Nostril si11g~e, tubttlar, projecting dorsolaterally. Gill inemhranes u~ited, free fran isthmus along the posterior margin, but attached by a fold of skin anter rly. Gill rakers roughly triangular, short, with about ten teeth on the tips. First raker en upper li[lb reduced to a nub and often absent. Psucdobranch reduced to a si le filament, often absent cntir~1y or present on one side only. Lateral line canal absent; insteEd, a single series of individual 11erves innervating neuromnsts in barely discernible cup-shap pores run dorsolaternlly. This series innervated by branches of facial ~erve VII, as nre free org3ns of hcaJ~ Pyloric caeca reduced to t~o small tubercles. gitta thick 2nJ rounded, with brosd rostrum [fig. 5]. Antirostrum ill defined in all age c;ttcgories; In~rginal sculpturing pront inent in younger fish. ~eJial surface of s3gitta charac teri:cd by dors3l and ventral dcpr~ssions, becorni~g more pronollnced with nge. St1lcus becomes less distincti~0 with age a11d cl1aracteri:ed by a raised central portion separating ostium atld c~uda in older fish. 'fhis feattlre is not as well defined in younger f h since medial surf ace is smoother, Ostium a round bottomed channel, widest anteriorly, sloping steeply at its margin. Cauda :d th round bottom, widest posteriorly. Crista superior some what ra ed anteriorly.

Coloration of fresh ecimens pearl-whi~e anter r- ly, becoming light tan to pinkish on the tail. Large adults with distinct bron3e tinge, disappearing rapidlv after death. Cheeks, gill cover and back silver. Abdomen light silvery blue. Thick-skinned spawning adults lose silvery hue and are uniformly pearl-white. Head, bodv 3nd tail flecked with black melanophores, which arc more ur:i form on tail and denser on nape and head. Margin of dor~al and anal fins black posteriorly in adults. Lining of mGuth and gill cavity dusky. Stomach and peritoneum jet black.

The known length ranges from a postlarva 19.6 mm

TL to a female 197.0 mm TL.

Conside:·able sc>xual dimorphism exists 1n ... r•:.:,:c• ib~~zaris. Although the skin of some females thickens 3t spa~ning time, the n1al~s can generally be distinguishe~ bv their thi- r, more opaq11e skin, larger teeth and flGtter dorsal profile tfig~ 2). !'>1ale premaxillary te:t'th lengths

::1rc gene-rally douhlc the females' at about lC.O m;~, ::;L .•\ standard Model II regression (Sokal and Rohlf, 1969) com- lI

paring the teeth lengths to premaxillary length

a high correlation (r= 0.933 for males and r=0.96~ for females). The ratio of teeth lengths to premaxillarv

lengths were significantly different between sexes Kith the 95' confidence limits of the slopes non-overlapping

above teeth lengths of 0.5 mrn (fig. 6). An F test for

significance in dif renee between the slopes with 59

degrees of freedom gave a value of 4.228 (p

CaJj_forniu specimens cX3lliincd were morphologically similar (except for sexual differences) and there was no signif- .I· icant variation tn the nuJllhcr of verr0brae amor1g these s.pcc.i.rr:cns (fig. fn.

!l i Has found to be in the inshorE' canyon area [area l) \,here

a deeper livi confJnilial, MeZanos g~a pamme~~e, was taken less f1·equcntly Lfig. 9). Farther offsbore, ire : I water about twice as deep, Melanostigma was more frequently encountered, although sampling in area 2 was less intense. Although nighttime samples were scarce in all intervals, a diel vertical migration was detected in L. mandibularis (fig. 10). The deeper living Melanosti]ma, however remained at about the same depths day and night. Some prey species occurred in greater concentration in the upper

200m at night, a pattern similar to that of L. ma~dibula~is

(fig. 11). ~o dicl migration was detected in mysids ~nd amphipods. These crustacean taxa represent several species

[Appendix B) and no species-specific patterns were resoived.

Postlarvae and young juveniles were taken in nearly equal abundance in all depth intervals, although ~hov coc- stituted the greatest proportion of ir1dividuals taken a~ovc

300m in daytime tows. No sexual differences were disc~rn- ible in the species' vertical distribution.

Age and Gr•)Vith

A monthly compar1son of age classes over an entire year revealed the winter marginal annulus formed quickly. Due to tho difficulty in observing tho extent of the marginal annt1lus in olJcr fish bccatJse tl1e otolith grows so1nc·- what SIJherically, ~ny Jctcct~lble winter rn~1rg1n W3S cotlntcJ.

This required the establishment of ~ove•nber as the arbitrarv birth month for the winter spawn.

The growtl1 rate did not differ between sexes, becoming indeterminate after the first year (fig. 12). Because of this and a small sample size at the u1•rer extreme, a Von Bertelanffy growth equation did not fit the observed curve well. A maximum age of five years was obtained from three females.

Otolith ages were compared to length frequency histograms constructed for the warmer and colder periods of the year (fig. 13}. The greatest frequency of :ero age class individuals appeared in the winter months. classes four and five were low in both seasons. The greatest number of individuals taken were two and three year olds, the age at onset of sexual maturity. Of 5~0 fish sexed of all year classes, 54.3% (304) were females and 45.7~ (256) were males.

Reproduction

The testes are elongate, bilobcJ org3ns cf equal si:c lying in the posterodorsal area of the coelom. T'hey arc attached by a medial mesentery dorsally to the peritoneum and ventrally to the inrestinc which is continuous with the perlcardial mesentery. The sper1n duct lies in a Jorsomedial groove along each testis. There is no separate genital pore or papilla, i.e., the urinary bladder and sperm duct exit througl1 a cont1non urogenital sinus.

ltJJJnature testes 3rc white ar1d rod-like, with rc~:nJcJ 26 anterior tips. Ripe males have greatly expanded, leaf like testes that turn a darker gray. In this condition the organs are about one-third the mass of the liver.

A single dorsomedial ovary lies in the posterio~ area of the body cavity in L. mandibuZaris. The OTgJD; attaches to the body cavity with a thin medial mesentery which extends from the pericardium, as in •he males. The oviduct is short and connects with the urinary bladder into a urogenital sinus. Ane at Sexual Maturity. Females under 120 mm SL, or less than two years old (see fig. 12) carried a maximum or 400 immature ova, each less th:1n 0~2 mm in diameter.

As these eggs matured, most of them ceased growth ~t abo;Jt

O.S mm so that eggs in a complement of about 100+25 ~rew as large as 1.7-1.9 mm in diameter (fig. 14). These eg~s were never seen in spent ovaries, i.e., the full complc~ent of mature eggs was evidently spa~ned. The rcmaini11g eva, then, would have grown similarly for the next spawning. size complements measured for the winter samples and compared to some sumn;er ov~1ries indicated complement m:lt uration was once n year. Although few adults were taken in the StlRIIncr, ten were 111 spawning condition ti1c~

(table 2) 1 suggesting no regular sc

The gonad indices for males (f 15) closely paralleled those of females (fig. 16) and indicat pre- tracted spawning. In the winter, most males sampled had recently spawned (testes very shrunken and loose, low index values). Althot•gh few ripe males were taken during summer, spent males continued to predominate then (table

2). Of course, it is impossible to know how long a g en male has been spent but latinous, leaf-like (expanded] testes indicating recent spawning We're encountered in every summer sample except June (of both years). Those males considered ripe had gonad index values above :o.

Food II

Prev Items. Juvenile L. man bul~Pis ate fe~cr

species of prey than odults, hut both groups relied 13ost

heavily on euphausiids and copepods (fios. 17 and 18). Juvenile fish ate large euphausiids and cop0pods in the

same proportion as adult~, b11t other food, such as sq~id

achs. The food items identified from all f h revealed a broad dietary specrrwn (table 3).

into shallower water at night, tl1e exception being mysi

and amphipods (fig. 11). Net mesh size precluded an 28 analysis of the distribution of copepods due to inefficient net filtration and no attempt was made to separate lood items into species categories, with the exception of

Gonatus onyx. That is, the group of amphipods consists of all captures of 17 species whenever encountered, but the group ''Sergestes,'' consists of only two species (Aprcnciix: B). Although daytime samples in very deep water were few, several catches were particularly rich in Lyeodapas and zooplankton. This resulted in the widening effect at the base of some graphs of figure ll due to a just-above botto~ concentration of organisms in the shallower areal. Feeding Chronolog_y. Frequency histograms of the four categories of fullness and recency of feeding (fig. 19) prepared for day and night showed a slight increase 1n stomachs with well digested food primarily at night [cate gory A, fig. 20), while stomachs with more recent itclC1s

(categories B and C) were found most often during the day.

A chi-stluure test for goodness of fit UIJtong c;ltcgorics ro test day-night differences in feeding wns insignifica!!t, however. Correlati11g the two recent and tho n\)t-rec:cnt categories with depth during day and night sl1owed most fish that haJ fcJ t·cccntly were above 400 m during the day (fig.

21). Although snn1pl0 si:c wus small in deeper h3tcr both da)· and night, most fish that haJ not rcccnrly fed here found above 400 m during the night. 29 The broad distribution of this species suggests there is no precise area in the water column wher2 they obtain food, but for a small majority of f:sh feeding occurs during the day. In the day-caught samples, 54~ of 446 fish (241) had food In their stomachs, while at night 42% of 84 fish (35) had stomachs containing food. Sensory Mechanisms of Prey Detection. Anatomical observations of the brain and cranial nerves comparing the optic, narial and facial pore innervations of L. man dibularis with the benthic eelpout Lycodopsis pacifica showed that the facial nerve of Lycodapus is larger than the optic nerve, as opposed to equal sizes in Lycodopsis

(Levings, 1967). The facial nerve innervates the-fish's lateralis system 1n a manner similar to that of the blen nioid Pholidichthys leucotaenia (Springer and Freihofer,

1976). Branches of the ramus buccalis facialis innervate the free lateralis organs beneath the eye and relay vibra tions in the water (fig. 22). The posterior branch of facial nerve VII divides into three separate rami at the level of midorbit. The posteriormost passes along the posterior edge of the preoperculum and sends fibers into the posterior series or free organs there. Tl1e anterior branch, the ramus Inandihularis facialis, passes ventra mediad the gill cover, sending branches into the preoper culomandibular canal and the anterior free lateralis svstcm. The middle branch, the ramus hyoideus, innervates the 30 muscles of the gill isthmus and branchiostegals. Both main branches of facial nerve VII carry visceral sensorv and motor nerves (Romer, 1962). The large canal DOTe'S probably relay stronger vibrations than the sn1aller free organs and a combined effect may aid in more sensitive prey detection. In any case, Lycodapus possesses a 1nore specialized lateralis system than either L:;codousis or

Maynea califoFnica (Warren Freihofer, pers. comm.). Apparently disturbances in the water are very important in the feeding behavior of L. mandibulaFis, with olfaction and gustation probably unimportant and visual contact intermediate.

Parasites and Predators The point of attachment and frequency of parasites were noted on examination of the stomach contents and gonads of 570 L. mandibulaFis sampled. The larvae of the nematode

Thynnascaris ad~lncum were the most abundant parasite encoun tered; 29.2~ of the adults and 15.2~ of the juveniles carried one or more of these worms. In contrast, the copcpoJ

Cal•-iio,:l.:~ete.s mc'd:,is,_Ic-;:u.'_~ was observed in 15 fish (0.9';; of the adults and 1.4~ of the ju~cnilcs). The ncin~ttodc W8S attached to the stomach and intesti11al walls~ fleritonetJm, gon3J, li~er, heart and beneath tl1c skin over the epaxial mt1scles. The copepod was attached to the gill arch or isthmus area.

A small unidentified cestode H:.ls observed lil two fish. !io 31

(1966) reports the bubble snail Janthina gZobosa is the intermediate host of the copepod C. medusaeus. Although there may be several intermediate hosts of Th!tnnasca!•~s,

L. mandibuZaris probably acquires this parasite from its crustacean prey (Milton Love, pers. comm.).

The effect of parasites upon host mesopelagic fishes is poorly understood. Noble and Collard (1970) suggested unencapsulated larv~l nematodes like Thynna3~aris

(or Contracaeoum) were commensals using various midwater fishes as paratenic hosts. Margolis (19701 reviewed the literature and concluded digestion in marine fishes was adversely affected by stomach wall 1n tations due to the inhibition of peristalsis. The thinnest individuals of L. mandibul s from the ~~nterey Bay survey always had the most nematodes. My observations indicated tho~c fishes were morphologically identical to the holotype of

L. att,?nuatzts, hence the erection of this name was due to emaciation caused by a stomach-intestinal nematode infes tat ion.

Predators of L. maniibulai~ia are unknown except that a sablefish (A11 mb1,ia was found to contain a single juvenile [Edwin Osada, pers. comm.). Presumably larvae and juveniles arc fed upon by 1nost larger species that capture other small midwnter fishes. Large adults are probably eaten most often by large squid and benthopelagic fishes. Aquarium Observations

Of the three temperatures of acclimation, ~- ~~:;;l-

ibuLaPis survived the longest at 7°C (twelve days), LlOT'.' than twice as long than at ll°C (five days). At 5.5°C,

corresponding to a capture depth over 800 m in Monterey

Bay, one specimen lived 7~ days, the other eight in this experiment died within three days (McCosker and Anderson,

1976). Specimens in the 7°C experiment were more active and became moribund later than in previous trials (moribund fish lay on their sides at the bottom of the container

motionless for only a day or two, then expired). Sev~n degrees Celsius corresponded to a depth of approximately

350-450 m in the bay. Based on t.hc catch data [fig. 10),

s would experience a temperature range of

about ~<-l'-°C •n" ''!•Jon t crey Bay (.ll roen k·e1< et a l . , lo-s_ 1 ,

~hen first introduced into the tank, hanhazard

sKimming anJ many wall contacts resulted in all the r. ~2n-

dib~Z~ s forming an aggregation which slowly circled the tank. They chiefly employed caudal flexures for propul- sion, but also used alternating pectoral beats to turn the

body sideways. After a few d=lYS !!lOSt fish would rest on t ir \'cntt-al surface' on the bottor.l with their tails coiled

oround :heir l1cads, but ~oon rose in the tank to circle in the current when the container lid was lifted admitting dull light. ilnlike the other captive pelagic zoarcid,

Lana.-:;-:_-';.):·---:::" " ,, always reacted 33 photonegatively even in dull light. Unlike the faagtcoth,

An~plogaster cornuta (Childress and Meek, 1973) L. ma~

bularis always fled when touched with probes especially when stimulated in the head region.

Brine shrimp (A~temia salina) were introduced into the aquarium regularly, as were euphausiids and biolttDin escent copepods (Pareuchaeta japonioa) from the trakl catches, to feed the occupants. Although Melan,st~g~; was observed to feed on the copcpod and brine shr one exception L. mandibuZaris was not. The single feedi~g was observed following a half hour of observation under darkened conditions when a small co ua attacked and

Taking three gulping bites and twisting from side to ~iJ0, the sh ingested the shrimp head first after the shri I S antennae stroked the head and body of the swimming fish. No behavior that could be interpreted as sexual or agonistic was observed in the aquarium, probably due to the stress of confinement. DISCUSSION

The taxonomic study of Lycodapus mandib~lJria revealed an extensive confusion 1n the literature of this species and its congeners. Recent authors (Bayliff, E'5-',

1959; Grinols, 1965) confused the species, chiefly since

~ost specimens were poorly preserved and/or trawl damaged. A careful examination of the type material plus numerous, well-preserved specimens showed that characters used to distinguish the species of Lycodapus (gill r~tker lengths, vertebral counts and head pore patterns) are difficult to detect without radiographs aJJd high magnification, hence the source of earlier confusion.

The association of £. mandibul11ris with suhmariile canyons promises an interesting study of isolation in sub populations as more specimens become available, especially from closing nets towed ncar, but not in, canyons. To date, only the ~!ontcrey Bay population is well sampled and factors governing the distribution aJtd biology of mcsopelagic fishes there may be applied to L. m~ndibtilaris populations.

Seasonal hydrographic changes have a marked in fluence on the temporal distribution of several mesopelagic species I.n ~!onterey Bay (Aughtry, 1953; Barham, 1957; Fc;st,

Fast provided a 1·cvicw of the hydrography of ~ontercy B3y as it affected the distribution of the l3ntern- fisl1 Stcnobrachius leucops:zr':IS. lie reported that during

34 35 the Oceanic period, generally July to October, offshore water 2° to 3°C higher than winter temperatures, lGoved into the bay bringing an "invasion" of offshore plankton including an influx of young Stenobrachius. As the sterns of winter (with winds primarily blowing from the south west) depressed subsurface isotherms, a northward flow was imparted to the coastal current to about 500 m lth~

Davidson Current) and the greatest increase in the pop ulation of Stenobraohius occurred. LAter, from Feb1·uar~ to July [Upwelling period), winds predominated from the northwest, surface water moved offshore, anJ ~ooler ~3t~r from below flowed into the bay through the canyon. A decrease in the population of older Ste~!obrau us ·occ~r- red then. Fast (1960) concluded that as the fish moved into surface waters at night they were swept offshore by this subsurface movement.

During the period of this study (Mar~h 197~ to Jantrary 1976) weak and irregular cycles were recorded in

Monterey Bay (Broenkow et al., 1975, 1976). The 1 isotherm has risen to the surface only twice since 19-1 and north~o:cst Nind intensities associclted with uph·colling have been lower than usual for several )'oars as hell

[William Broenkow, pers. cornn.l. These data suggest ~orne unpsedictability in the hydrographic seasons inside t bay. 36

Although no seasonality of the local L. ma~dib uZaris population was detected throughout this study, during periods of strong summer upwelling in which sur ce water is moved offshore, younger individuals that con centrate in shallow water could be moved out of the bay. During the subsequent Oceanic season, thev then may be returned to inshore areas as Pearcy (1976) showed for micronekton off Oregon. At any rate, the Monterey Bay population of L. manditularis seems to be influenced to a lesser degree by current movements than Fast (1960) found for Stenobraahius. Seasonal hydrographic changes have a marked effect on the ab••ndance of the planktonic food of L. mandibularis (Hebard, 1966; Laurs, 1967; Youngbluth, 1976).

A diel vertical migration in mesopclngic species in order to feed in richer surface waters is well docu mented (Hardy, 1953; Pearcy, 1970; Cailliet, 1972; DeWitt and Cailliet, 1972; l'earre, 1973; Clarke, 1974; Isaacs et al., 1974; Raird et al., 1975; Gorelova, 1974, 1975; Youngbluth, 1975, 1976; Brinton, 197b). The diel cycle noted for most prey species of ~- mcndibk1a•• incl~J~s several species per taxon represented in most instances. With this and small sample size, typical species-specific patterns of migration could be obscurred. For abundant ups like euphausiids, population move:nents are blurrea by individuals remaining at depth on a given night ~tile

-others migrate [Youngbluth, 1975). In a quantitative

analysis, Brinton (1967) considered daytime net avoidance a problem, yet with the larger Tucker trawl used in is study, only adult decapods and some fish and squid should be considered capable of dodging the net.

The ding chronology of L. ma~1ibuZu!~is suggests many fish migrate into the surface waters at night with predominately empty stomachs. Then, as deep scattering layer organisms descend into the canyon 1n the morning,

they are consumed by L. mandibuZaPis which then return to deeper water to ' st and digest'' (Legand and Rivaton,

1969). This pattern of movement is not well defined, however. Actively feeding fish probably rely on cephalic

seismosensory stimuli to orient toward prey with vistt3l cues important when the fish come close ro their prey.

The majority of prey species of L. mandibuZa~is are bio- luminescent, indicating some degree of visual food selec- tion in the fish. The results of the food study are probably not

b3dly biased by net fcedin~. The great proportion of di ted material found in L. mandibuZaris stomachs and th• remarkably active condition of most specimens after capture suggests they swim with the net during tow without 38 much physical stress or early concentration 1n the cod-end where plankton accumulate (Hopkins and Baird, 1975_, l975b).

Very little has been recorded of the spawning habits of eelpouts, with the exception of two investigations of benthic species where parental care in Lycodopsis pacifica (Levings, 1967, 1969) and Maynea califor1:i~a

(Klievcr, 1976) were suggested. ~lead et al. (1964) suggested zoarcids aggregate and produce demersal eggs tha~ develop close to the sea bottom. Slipp and DeLacy (1952) artificially removed most of the eggs from a gravid female

Lycodes palearis whicl1 later laid the rcmajnJer in a sanJ- bottomed aquarium. Tl1e unfertile eggs laid about the bottom singly or in small clumps and no other observations were made. Kliever (1976) suggested the fall and winter decrease in the number of male Maynca cali[or11ica In drif~ kelp habitat traps 1n Monterey Bay may have been due to the I differential movement of individuals engaged in spawning. Also, Kliever thought spawning anJ larval clcvclopmcnt occurred on the bottom near the Monterey Canyon axis ~here the majority of sheltereJ l1abitats (clumps of decomposing seaweed) occurred for this eclpout.

No eviJence exists that L. mandihulaPis utili:es the sea bottom for feeding or spawning, unlike most of its confamilials. Ripe orange-yellow eggs stripped fros the 39 largest ovaries contained dark orange oil droplets and were neutrally or negatively buoyant in dishes ot seawater in the laboratory. Some buoyancy may be added by a ter-

tilization membrane, though. The vertical distributio~ of the smallest individuals ranged widely day and night, so a larval life in the lighted surface waters may not he

the rule in L. mandihuiar•is as it is for most mesopelagic fishes (Marshall, 1954]. With these results, and the assumption of a selective disadvantage to bottom spawning

(where more potential predators li\'e than in rnidwater), egg laying most likely occurs between pairs that chance to meet in midwater. If this occurred chiefly during the day when fish are in deep water, most eggs would drift just off the bottom. The pelagic larvae would then have

to make their way to the surface after hatching unless upwelling water distributed the eggs shallower. AlthcG;t

no spawned eggs of L. man b••laria have actually been identified, postlarvae from Jeep water have been taken, and the above spawning hypothesis precludes r:nrental care .

., . '" ~ . "' What has governed the evolution of • rr: a:-: :2 t- t: -:.: ~,-_ ,: :r· ·: .:1 :

~arshall (1971) suggested that benthopelagic

toh'Jrd a "markedly lo\\'C:r lcveln (d.ecreu:sed metabc.lism 3rL1 activity) on invading bathypelagic dept since there 1s more food on or near the bottom than in midwater. However, 40 if canyons act as concentrating basins for plankton, as Fast [1960) suggested for Monterey Canyon, a reversed abundance of food from that postulated by Marshall (197.) would favor a small benthopelagic species evolving ada~r~ tions to life in midwater. This has apparently occurred only in L. mandibuZaris~.L. paPviceps and L~ a2~straZ~s, since the species of 0 us~ as far as known, live close to the bottom. The selective importance of th~ avoida~ce of predators by assuming a mesopelagic existence in these three species is difficult to assess since the depths tn habited by probable benthopelagic predators ovcrlop the known depths of all codapus species. Besides, the mid water 1s not without its share of formidable predators of small fishes. Predation on the shallower living species may have been more intense in the past than on deeper living species, though, simply because biomass (and hence numbers of predators) decreases with depth. It is interesting to note that the three pelagic species of ll2 today inhabit shallower water than the benthopelagic spcc1es and live closer to shore. The chief cve11t in the evolution of tl1e pelagic Gya liS SJlCcics, it appears, has been the movement from the benthopelagic environment to feed in richer midwater wherever plankton accumulate. '!'able 1.

Koristi~n and roorphomct:ric11 r:tf [.yrmdapwr specie& in C~:~lifornia waters, =mparin; t)'pG lm1l~u;hl t.£l :rocunt (SMIMP) colluction!:l. Mnasurc:munt!l ilro parcont standm::.d length !ll'i'cept pruma:dllacy teeth lenqths and .SL !in r.m),

£, fi.eroll[cv Z:.. daJ":lai;in.uu J.,. grorraidmw [.,, rJI'VIlUl.denu L.""'"'ri1:11l.iibJ 14r"70 L-:--iittemwtuu L. Zycodon L. r.:andibr;lariu Chlln.ctar .synt-.\'1:'

!lead lu.t• u, 15.4 18.1 14.!! 14.)-17.5 14,0 13.9 14.7 l4..:-)_:A!hL_ __ 7 .1.-8.3 ,_, Prc!llllllill len th -- !l.i! 7.0 '·' '·' £.::!-8.4 orb.it <'Ji.,mete:r 3.3 ,~.b-'L 3.5 ____:2.9-4.0 2.1 ,_, ).:.!-4.7

'"' ..J.M~l llod du th -- a., 6.7 5.1 ---- B.! a. 7 " ____ Pectora.l ltmCth 7 .:! 4.J-7.4 ,_, <'tk.Btu) 2-3 2 (soc!!;ats) l 1 l.,:,L.-...... ___ , ___ f!!.,l.,?tln" le(!Lh {L+RJ - ·- 2-15+0-8 ·-~,~--;.l.l:.L~- OH ,., 0-lfl+tl-lD "' ~,.._n :Sranchior.te al!: ' 0 0 ___ Vertchrlle lJ+H-..,eu L4H!!e7r~ lJ.J+69,.U5 1'0-16+73-?S:BS-31' l&+7S"'J4 lfi±J.6"92 Hi+75"'91 15-17+"12-Bo->-S7-':17 ll<:~tl:!lll rar~ "' ·-·-- 7S l!.L.-. 85-ElE " as B7 a:~~_'l-'--~·- l'm.al <'• " " " " 73-75+ 7' " 7S 73-82 ~.Lms a B ' ' '7 ,_,' A ' ,.,' Pectoral razs 7 ' -- ' ' PI'Oi' fnn:mla ------3+1+4 Ul J

""'>-' 42

Table 2.

Numbers of ripe and unripe adult Lycodapus mar~ibularis sampled each month in Monterey Bay.

Ripe Unripe Ripe Ur.ripe !>!a nth males males females females

January 3 9 9 8

Februar 4 14 4 18

!-larch 8 15 21 5

Aril 16 ll 19 ll

May 3 16 2 9

June 0 l l 0

July 3 6 2 ll

August 4 3 5 3

September 9 8 20 5

October 5 4 7 4

November 6 15 14 18

December 2 2 0 4 43

Table 3. Species of rnidwater animals identified from stomachs of Lyccdapus mandibuZaris taken in Monterey aay ~

Phylum: Arthropoda Order: Copepoda Ca Zan us sp ~ unident. calanoids jcmonica Order: Euphausiacea

spinifera unidentified euphausiids

Order: ~phipoda Orchomcne obtu.sa Par>ac~1z Zt:sorna coeeus unident. gammarideans

sp.

unident. hyperiids Order:

Order: Dec._=tpoda

unident. decapods

Phylum: ~1ollusca 44

Table 3. (continued)

Oc topo teu this de Ze-tron unident~ squid Phylum: Chaetognatha Sagitta sp. Phylum: Chordata Salpidae, unident.

Cyclothone si[Jna.ta LeurogZossus stilbius 45

LANDING

0 • I I I I I II KILOMETERS

50'

Fig. l. Map of Honterey Bay, California, showing the two trawling areas.. Canyon contours a.re in fathoms. '"f-'· "' "' n >--' "rt "' ~- "0. "'>--' "'rt (D [/J

ti (D "">--' '< (D "'[/J rt 0 0 "' "

9v - Fig. 3. Lycodapus ma1wibularis Gilbert 1915. Upper, male, 140 mm SL; lower, female, 149 rnrn SL, from SAMMP tow 70, Monterey Bay. 48

NP PrP lOP

MP AP PP

Fig. 4. Head of Lycodapus mandibul=is illustrating cephalic pores and free lateralis organs. Abbreviations are: AP, articular pore; IOP, interorbital pore; MP, man dibular pores; NP, nasal pore; PoP, postocular pore; PP, preopercular pores; PrP, preocular pore. 49

SULCUS r----'------, CA\JDA OSTIUM

...... : ...... ~J'- ·. ·.:. :...... ·::"=;"~<- .:·.·_<·:·:.· ··~~,,~&· ROSTRUM

CRISTA SUPERIOR

Fig. 5. Left otolith (sagitta) of Lycodapus mandibularis 192 mm SL. 50

1.5 0= FEMALES Y=0.0673X-02563 N•2B E r=0.964 E ,,, <>• MALES =;- 1.0 u Y=0.2080X-1.30C z w _j N='32 ::: r•O.S33 ;- w w ;- 0.5

2 4 6 B 10 12 14 PREMAXIcLARY LENGTH,mm

Fig .. 6* of ml.llc: and :emale _pn:•rnaxill21ry tG,:;th len,;rths in iLuL

J Prince William Sound }..._ l fwJ'9v~~ ~6 0"<- ) (~,ynn Canol

~ 7/1:;$ ~ , Glocence S!rOJt ~·' ~ t 't;f. "'~~ ~1Gcorgltl Strait - 50" "-(3 . Puget Sound

} Astoria canyon

~Eel cony on .... 40° ~~z cui no con yon M niarey conyon

Hueneme canyon 11('-' ~\o Jollu canyon i- 30" ~1:\ ' ~ '\", ~\\'" 150° 130° 120° (• ~ I I I I I 'l'J '\

Fig. 7. Map of the eastern North Pacific showing collection localities of Lycodapus mwldibular>is. 52

MENDOCINO CANYONS • •

l MONTEREY CANYONMm

HUENEME CANYON (30)_ I • I LA JOLLA CANYON • BB 89 90 91 9 2 93 94 •5 96 97 96 TOTAL VERTEBRAE

Fig. 8. Distribution of vertebral counts in Lycodapus mandibula:eis from California waters. sample size in parentheses; single records are circled~ Graphics after Hubbs and Hubbs, 1953. 53

0 {7) 100 . I A {8) I E 200 - !! (II) •E 300 . I I ,..£ (II) I I 0. 400 w (10) 0 500 - I (2) 600 - I ' 100 75 50 25 0 2.5 50 75 100 "'/o FREQUENCY

0 (0) 100 {0) B 200 - {I) 300 ~ - ,/! {I) • 400 E - - (3) ",.. 500 - I I w.. (2) 0 600 - I I (3) 700 I I (4) eoo - I I ' 100 75 50 25 0 25 50 75 100 "I<> FREQUENCY

Fig. 9. Vertical distribution of Lycodapw; mandibuZaris (left on graphs) and MelanotrM.gma pammelas (right) in Monterey Canyon, day and night. A) area 1, B) area 2 (see fig. l). Number in paren theses sample size. Greatest bottom depth in area 1 about 700 rn; greatest bottom depth in area 2 about 1400 m. 54

{0,7) 100 Jl l (6,2) 200 I \ ll I ' (9,3) I \ I I 300 I \ J \day \ \ night (9,3) OEPTH,400 I , ( \ meters ~ \, (10,3) 500 "" ~' \ I / \ I (3, I) 600 ~' \ I (// ) J (2, I) 700 ;\ - 2"~ \ L_j (2,2) 1>00 A 0 0<3 0·6 0 0·3 0·6 CATCH, no.;hr.

Fig. 10. Vertical distribution of pelagic zoarcids in l·!onterey canyon. Al Lycodapus mandibu zrn,is and B) Melanostigma pammeZas. Sample size in each depth interval in parentheses; day on left, night follo>ling. Catch per hour mirrored. 55

or------.

I 00 1-

200 1-

300-

DEPTH, 4001- meters 500 f-

6001- day night 700-

800- A I I 0 7.0 14.0 0 7.0 14.0 CATCH, mi./hr.

I 00 200 ~''; /~ 300 I DEPTH,400 <, meters 500

/ 600 / ? day / night 700

800 B

0 .015 .030 0 .015 .030 CATCH, mi./hr.

Fig. 11. Vertical distribution of zooplankton found as prey of Lycodapus mandibularis. Sample size as in fig. 10. A) euphausiids, B) amphipods, C) Sergestes spp., D) Pasiphaea spp. , E) rnysids and F) Gonatus onyx. 56

100 //' 200 / ~ 300 < "; DEPTH, 400 meters \ I \ I 500 night 600 )~ \; ' 700 II ~ b 800 c 0 2·0 4·0 CATCH ,mi./hr.

0 100 j\ 200 / \ / ~ \ 300 / " (./ DEPTH, 400 ( I, meters 500 night 600 II il 700 )C::: / ·- _____ .:::::,_ L.~ D 800 0 2·0 4·0 0 2:0 4·0 CATCH ,mi./hr.

Fig. 11. (continued) 57

0 I I 00 ' 200 }\ I\ 300 j' \" DEPTH, 400 ./~/ /~ meters

700 :I u" 800 E

0 .03 .06 0 .03 .06 CATCH, mi./hr.

0 //~~,~ I 00 ,,I, <" 200 '"' ,/ ''·1 (',// 300 )\ 1 i / '\ I i ' I DEPTH ,400 \ I I I meters J I 500 I I ( day \J 600 / "'· ( " I )"''" 700 I ; ___ j ~j 800 F 0 .05 .10 0 .05 .10 CATCH, no./ hr.

Fig. 11. (continued) 58

t 175 - 150 ++ - ~ ~ ~ 125 - ~ SL,mm 100 - ~ ~ 75 f-

50 f-

25 1- (9) (8) (9) (9) (13) (15) (2) (2) (0) (3)

I I I I I I 2 3 4 5 AGE CLASS, yrs.

Fig. 12. Growth curve for Lycodapus mandibularis in Monterey Bay as determined by otolith readings.

Right 1 females; left, males. Sample size in parentheses~ 59

20 r- A

%F :-- - 10 r- :-- ~ t- I "l .20 40 60 80 100 120 140 160 lBO'

r- 20 - B f- o;., F 10 - -f- - ,...-- t- .-I ~ _I 20- '. 40 60 80 100 120 140 1so 180 SL,mm

Fig. 13. Length frequency histograms of Lyaodapus ~andibularis from Monterey Bay. A) October to !·!arch, N=298, B) April to September, N=26L 60

0 120-129 130-139 140-149 150-165

Millimeters, S L

Fig. 14. Frequency of egg sizes in Lycodapus mandibuZar>is from September to April. Sample size in parentheses. 61

200 f-

175 f-

150 f-

"' f- "' 125 -- ""'"' ""'0 100 f.- <= 0 (!) 75 f- -- 50 - -

25 1- -j -1 " 0 I I ~I:. 0 I 1 i J F M A M J J A s 0 N D (12) (18) (23) (27) (19) (I) (9) (7) (17) (9) (21) (4) Months

Fig. 15. Monthly gonad indices for adult male Lycodapus mandibulal"is. Vertical line and end bars represent monthly ranges, horizontal line the mean and dark vertical bar the 95% confidence interval. 62

BOO I-

700 I-

600 I-

>C I- -.:J 500 "'c: r -r "'CI 400 - c: "'0 C) 300

-- 0 200 -

100 - Ij 1- l- ~!.. f I ;m: 0 I J F M A M J J A S 0 N D (17) (22) (26) (30) (II) (I) (13) (8) (25) (II) (32) (4) Months

Fig. 16. Monthly gonad indices for adult female Lycodapus mandibularis. Graphics after fig. 15. 64

Euphousitds I 60 f- Sag!/ta sp. Cope pods Solp) 40 1- %N I Un1denl FISh/; 20 1- / Cr7~7·$ 0

20 1- o;o V ~J\\\ 40 f- ...

'"""Pasrphaea ~- "' sp, 60 f- \\ My s ids Other Decapads

I I • 20 40 60 0/o F

Fig. lB. Index of Relative Importance plot of stomach. items in adult Lyaoda:pus mandibular_is; N;l44. 65

FULLNESS 2 3 4 A D w 1-

Fig. 19. Categories of stomach contents condition (after DeWitt and Cailliet, 1972). 66

BO - DAY

60 '

•

20 '

0 A 8 c D 80 NIGHT 60 •

%F40 •

20 •

0 A 8 c 0

Fig. 20. Fullness and digestion state category histograms for Lycodapus mandibuUn'iB. Day, N;215; night, N;38. 67

NOT RECENT RECENT 0

200 ..... 1ii.. E -400 :J: 1- Q.. w Cl 600

75 50 25 0 25 50 75 %F

Fig. 21. Day-night chronology matrices (see fig. 19) of stomach contents in Lyaodapus mar~ibuZaris. Dark bars, night samples; light bars, day samples. Sample size: Day recent, N~86, not recent/ N=l29; night recent, N=l7, not recent, N=2l. remus buccelis facie Iilii

/ remus opercularla superficialis faclai111

ramus buccalis ramus hyoldeus accessorius facialis

Fig. 22. Lycodapus mandibularis: lateral view of head showing branches of facial nerve VII that innervate the free lateralis organs and canals. General cutaneous branches not shown. LITERATURE CITED

Alton, M. S. 1972. Characteristics of the demersal fish fauna inhabiting the outer continental shelf ancl slone of the northern.Orogon coast, in, A. T. Prutcr and · D. L. Alverson (eds.), The Colur:;hia Rivc:r estuarv and adjacent ocean waters, Univ. Washington Press, Seatt.e and London, 583-634. Ahlstrom, E. H. 1969. Mcsopclagic and bathypelagic fishes in the California Currcrlt region. Calif. Mar. Res. Comm. Cal C. 0. F. I. rept. (13): 39-44.

Alverson;, D. L. 1951. Ne1v records for mar1ne fishes from SOLothcastern Alaska. Coneia (l): 8(,_ Andriashev, A. P. 1935. On a new fish of family Lycodapod idae from the northeast coast of Kamchatka. Comptes R<:mlues (Doklady), Acad. Sci. USSR ~(69): 422-,124.

1939. Essa)· on tho zoogeography and origin of the fish fauna of the Bering Sea and neighboring waters. Univ. Leningrad, 185 p.

1954. Fishes of the northern seas of the U.S.S.R. Trudy Zool. Inst., Akod. :\auk SSSR, 53. Israel Prog. Sci. Transl., OTS 11160 (1%4), 617 j):"" Aron, l'i. 1959. MiJwater trawling studies in the North Pacific. Limnol. and eanog. 4: 409-418. 1962. The distribution of animals in the eastern ~ortl1 Pacific and its relationship to pl1ysico1 and chemical conJitions. J. Fish. Res. Tid. C3n. ~-.2_[2): 2?1-314.

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Bailey, R. ~1., J. E. fitch, E. S. Herald, E. A. L::~chm;r, C. C. LindseY. C. R. ~ohins and ~- ~. Scott. l9b0. A 1 t of comm~~ and scientific names of fishes from the United States anJ Canada. Amer. Fish. Soc. Snec. Pt1bl. (h): lc\9p. .

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Ho, .J .. -S. 1966 .. Larval stages of Ca i ;;~tes ~~[!. {C2li- goidea: Lernaeoceriformes), a copopod parasitic on fishes. Bull. Mar. Sci. 16(2): 159-199.

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1976. Vertical distribution and dicl migration of euphausiids in the central region of the California Current. Fishery Bull. 7.+[4): 925-936. Appendix A

Trawl data of the Steinhart J\quarium HidwatE!r HaintE!nance Program, 1974-1976, in Monterey Bay.

T0\'1 BOTTON 0 ------5Tl\TION Dl',TE T!..~!Er hr:; DEPTH, m DEPTH, rn NWO START 'I'Ot-v COURSE, T 1A-l 4 !"!ar 74 1040-1335 0-750 841-1005 1500 36°46.3' 121°59.1' 270 2A-2 2 t·1r'"tY 74 1036-lUG 0-3fJO 421-530 750 36°47.5' 121°:)4. 5' 220 3P,-J 3 Nay 74 l 006-1128 0-580 1006-732 llOO 36°45.9' 121°59.4 1 280 4J\,-4 18 Ju1 74 uoc,-1]37 0-330 457-658 750 36°46.8' 121°54 .9' 230 Sl-\-~) 'J" 1 ~- ·- Ju1 74 1155-J22G 0-507 869-1171 1200 36°46.2 122°01. 5' 210 I:. 74 OJJS-0415 50IJ-401J* 98f.l-1l34 1064 36°45.9' 12~ 01.5' 135 llA-1B 7 Oct 74 0~•38-1038 3~~0-190 732-567 500 36°47.1' 121°55.6' 235 llA-ll)** 7 Oct 74 1204-1304 5~H)-52i! 640-548 1 3 ::-,o 36°47 .4' 122°01.2' 240 lll\- 20 7 Oct 74 1447-1'547 520-432 l097-t359 1200 36°4~.6' 122°02.4' 140 12h-21** 4 t-lov 74 0729-CJ[-l29 367-259 6:'!2-SG7 700 36°46.7' 121°59.3' 275 121\-22 4 Nov 74 11•15-1245 301)-360 567-722 ElOO 36°46.9' 121°56.2' 235 12h-23 Lj Nov 74 1354-1454 236-180 786-722 500 36°47.4' 122°01.1' 090 12A-24** 4 nov 74 1555-1655 312-223 768-530 RSO 36°46.5' 121°57.2' OGO 1 ] 37\-:_'!)*1• F ,) Nov 74 \.iRl3-0913 32·1-144 428-73::? 500 36°47.4 l21°5CJ.4' 240 ·1 -~;~_- :·~c p:_ nov 7~ ]<'J]f1-lllll 39(>-345 •184-3fl4 800 36°47.0 1 121°~)fJ. 2' 07() ) 3A-:!7 lf) Nov 74 1216-l :nr:; 410-30S S94-47G 500 36°47.1 1 121°~\6. 1 1 23S l ]l\- 2U 15 nov 74 14 :~h-1 1

141\~ :z~.l 25 t~ov 74 1849-1949 374-364 530-750 600 36°46.6' 121°56.0' 240 l4!'i-30 25 Hov 74 2048-2148 216-202 768-585 400 36°46.2' 121°57 ~ 9 1 090 l•lA-31 25 Nov 74 2241-2341 504-446 594-768 700 36°47.2' 121°55.6' 270 14A-32** 25 Nov 74 0039-0139 250-200 869-658 400 36°46.3' 121°58.6' 070 (' 1 f) A- 3 J+ 16 Jan 75 llU-1122 432-450 <132-450 1200 36°47.9' 121"'54.2 1 270 l5lt-34+ 16 Jan 75 1332-1339 39G-.378 396-378 1200 36°45.2' 121°55.1' 235 167~-35+ 28 .::;an 75 1116-11 2CJ S'J0-615 590-615 1275 36°47.2t 121°56.4' 110 l7A-3b 30 ,Jan 75 1112-1212 2H8-230 494-•102 500 36°47.3' 121°57.1' 070 1811-37 ll Pet: 75 0837-0937 2G2- 210 549-384 50:] 36°47.2' 121°54.2' 250 1 l!lA-3d ll Feb 7:) l~lJS-11:-;5 648-504 650-557 800 36° 46.4. 121°58~1 080 l8A- 3'3 11 Feb 75 1241-1341 173-137 366-311 300 J6°47~7t 121°55.8' 260 0 lBA-40 11 Feb 75 1440~1540 4 25-367 530-585 814 J(-, 47.7• 121°58.4. 065 l9A-[) SBS-4 57 GOO 36°1i7.4* 12 :14~9' 220 1 1 1- n r 1 23A-5l 24 Jun 75 l20H-1:108 252-396 7GB-4S7 650 3Gc <1U.B .12.1 _;(l •.) 0~15 1 24;-\~ 52 28 au] 7 ~l 20t1 :::-::14 2 800-698 l2b'J-1 07 ') HOiJ 'J6°44.] ' l:~:-toc .6' 330 1 241\-5}** 29: Jul 75 0112-02]2 439-33] 65H-5R5 700 36°46.3 1 121°:=-l:J~G,, 060 24A-54** 29 ,)u] 75 0338-0438 3 24-238 FIOS-558 550 36°47.3' t 21- ~lf.L 3' 065 0 25A-S~} 15 f:J.IJ\j 7 ~) 2lll-22Jl 7 so-coa 1171-9<18 '!40 3() 44.2f ] ~::t\·1] . 2. 315 1 2 Sl\- SG .J Auq 75 2314-0024 576-547 lllt.-1225 850 36°44.8 ]_;_;2°01.9' 135 ' ' 0 2~Jl:;-S 7 16 r,ug 75 0139-023 J 43'2-J35 1134-C9S 685 36°43.9' 122 01. ;;:t 31S n 2rd\-~l)! l ;:, A ;v; 75 02 ::)(}-04 ;)g 2r15-22.3 786- 5B:J 400 ](i-th).JI 121°5-7.7 l O~tO 0 26A- c,(j 17 i;ep 75 094H-l0~1H 400-300* 'i27-74l con 3t?4f).9' 121 ~:s-6.0 1 2 ~'. ~) co I-' Ml__;pend:l x A (continued) TOW BOTTO'\ !~~J:~l.']:~~.~l.----~~.:_\TF:__~~-.!;~_!;I:: , hrs DEP'I 1 ~-~ m DEPTH, m Hl'/0 S'l'Jl.,HT rrO\'i COURSE, o,r___

;?(,A- C () 17 7:) 1141-124] 600-~)00* 933-951 BOO 36°46 .. 6t 121°59.9' 275 ~, c ?r~.n,-c 133~ J 17 S>.;p ' ; .. -1435 300-200* 686-466 550 36°46.1' l21°58,C' 050 :17h-62 24 .':~ c~o 75 l34fJ-l.J4il 700-600* SOS-969 900 36°46.2' 121°59~3' 250 2Lt1-G:j 14 Oct 75 0950-10~)0 518-453 786-640 700 36°46.9' l21°5G.4 ' 225 :~~h!\-64 14 Oct 75 1156-L2S6 800-700* 786-942 925 36°46.4 1 122°00.4' 250 :if3l\-(!5 14 Oc;t 75 J4 J.l - 1:5 ll 396~324 C,SfJ-4 7i) 580 36°46.3' 121°58.6 1 050 E5C7_r::,-GG 28 75 2Jfll-2~Lll 0-100 366-110 lF, 36°47.5 1 121°58.4' 070 0 ESC7 ::,-G 7 29 75 o3o2-cqo::: 0-100 3G6-110 125 36 4{'.). 1' 121°57. 9 1 070 (>")nr.: "'"'"lr ESC75-6B 13 Oct 75 "'"· ,,_)-'._,·~j:_) 0-100 4'::l4-/.i21 175 36°47.8' 121°57.7' 245 0 £Sc:75-G9 ll 0::-t 75 OJ2--1-c;J:)o 0-100 293-677 l80 36°45~7' 121 56.9' 270 29!>-70 5 ~-1ov I:; 1922-2(;~2 489-3')6 732-594 G40 36°47.1' 121°56.1' 230 ;::'-))\- 7} ::-1 ~kt'J 7 ~~ 2J.2tl~:C:2?4 44G-34S 695-567 560 :;,c/--)4().7' 121°57.5 1 065 0 ]J;;\-7 :.: J f:i;'r' 7 JiJ0:-:-}102 525-4·-16 8 23-I:__ SlS 700 3G 4h.8 1 121°56. 9' 230 0 3Di\--7~ ::Jec 7 :> 12lG-l:llO 40;8- 396 732-585 750 3() i]6.2' 121°58.3' 055 31!1-7'1 8 Dt::c 75 094;:1-J 4 475-.:103 7J)~()3.1 f,\)0 36()i17~2' 121°55.9' 235 0 1 311~-7 5 B nc~c 75 115f;-l2'>8 J%-2.S8 73:!-47;) 500 3(; <16~G 121°':!7.3 1 060 J2A .. ·76 23 .Jan 76 oqlc,-lfJ16 330-273 402-585 ::so 36°47.2' 121°56~E:' 245 0 1 321\-77 23 ~Jan 76 1133-1733 518- ]~J6 530-64 9 750 3fi 4G.;! 121°5f\. 4 t 080 0 322<.-78 23 J.:.1n 76 1247-1447 504-403 750~-1~)7 7:10 3t?46.8' l:U 56.4 1 230 0SC7S-H'I'1 2~1 :fuJ 75 2;?28-2~5tj 21n-1s5 ~158- 567 350 36°47.3' 121°54.5' 245 1 C!SC7 5-N'f~ 26 ,1 u] 7S 21 :;2-2222 230-173 :::;c;-:1PS 3 :;o 3tt iif .2' 121°5:1.5' 245 ') -J 0 0 OSC7 5-~N'I'S '"' Jul 7'::; 004B-OlJB 94-79 366-.;Js 150 .Jb'' i 17 • 8' 12J 51.6t 240

+ Bcd:tom trav;J~:; with 2.1 m be,~Hn trawl b3

Appendix B

Partial list of animals caught in the Monterey Bay area by :nidi.\ra r.er trawls fished off the bottom, 1972-1976, in water deeper than :200 m.

Phylum: Cnidaria Class: Hydrozoa Order: Leptornedusae Acquorea sp.

Order: Trachymedusae

Order: Siphonophora

Class: Scyphozoa Order: Coronatae

A to Z. la W](l)?.: l Ze i. Par•(xphJ Z ;:?.:na p,~zriDnrn: Periphy lla p.2:.r'1:I/hu llJ. Ctenopbora

Phylum: Annelida

Vanad/s L.a Phylum: Chaetognatha

lurn: Arthropod.:I Order: Copepod3.

(?) {continued}

PaT'~uc:haeta japoniaa

Order: Ostracoda Giga.ntoeypr"Z:s

Order: Hysidacea Boreomys"~3

Order: Amphipoda

forma 'hystr~:;:

sp. E Sti? Appendix B (continued)

Order: Euphausiacea Bentheuphaus-i.a wnb lyor's Euphausia paci['[ca NematosceZis diffici Zis Thysa:rwessa spirdfera Order: Decapoda

Sergestes svnt. l.-u:;

Bentheoger-rrzc:na_ bur~~enroadi

Pa:.:::iplLaea

t.'lLI;n:.a

HymenodolYl: f1' 10rztc Z-is

PandaZ.us Jor>clan-i.

Phylum: ~1ollusca Class: Cephalopoda

Order: Decapoda

' ' _,. Gor:.:-'I. [)t;·Y'G'U L 1.-B B (conti01ued)

Gcnat1.!S orlJf~T Chi t?ote-ut" hi.s Valbyteuthi2

Order: Octopoda Japei::eZta hea·thi Oetopus spp. Orn:n cali;~JPniar..~::z Order: Varr.pyromorpha

Class: Gastrogoda

Cavo sp.

Phylt'"-m; Chorda::a Class: Larvacc:J.

Class: Thaliacea 8-_;

sp ..

Tic' 1. 1 n:. '·''" sp.

Chondrich Family:

I.. ;f>, Appendix B (continued)

Class: Osteichthyes Fw"TLily: f\~lepocephalidae

Family: Searsiidae Pa llisoZus L·is saawm:eh Family: Bathylagidae

CX'L l."LS Family: Sternoptyc:hidae Z.e.:::-Js sp.

r: Ina

Family: Go nos toma t iclae

Far::.i

Z~t.:.

Fo.mily: B (co:>tinwcd)

Fa."TTily: Chauliodontidae

Fami !·1alacosteidae scint·i llans Family: Myctophidae

Family: Paralepididae

Family! NotosudiCae

Family: Hclampaidae

Family~ Sc:ornbc:r e soc idae

Fam:ly: Cen t:ro lo_rhJdue

Trachirteridae

?ami 1y:

:..: ; Apper~clix B (continut::d}

Family:

Family: lopoma tidae

Family:

Family: Brotulidae

?amily: Zo?irc:idae

Farnily: Bothida•:::

Family: P leuronectidae

sp.