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VIMS Articles

1986

The Abundance And Distribution Of The Family Macrouridae (Pisces, Gadiformes) In The Norfolk Canyon Area

Robert W. Middleton Virginia Institute of Marine Science

John Musick Virginia Institute of Marine Science

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Recommended Citation Middleton, Robert W. and Musick, John, "The Abundance And Distribution Of The Family Macrouridae (Pisces, Gadiformes) In The Norfolk Canyon Area" (1986). VIMS Articles. 623. https://scholarworks.wm.edu/vimsarticles/623

This Article is brought to you for free and open access by W&M ScholarWorks. It has been accepted for inclusion in VIMS Articles by an authorized administrator of W&M ScholarWorks. For more information, please contact [email protected]. THE ABUNDANCE AND DISTRIBUTION OF THE FAMILY MACROURIDAE (PISCES: GADIFORMES) IN THE NORFOLK CANYON AREAl

RoBERT W. MlDDLETON2 AND JOHN A. MUSICKs

ABSTRACT

The Norfolk Canyon off Virginia and the adjacent slope areas were sampled with 13.7 m otter trawls in June 1973, November 1974, September 1975, and January 1976. Trawl depths ranged from 75 to 3,083 m, and 22 species of macrourids were captured during the study. Co-I'Yph.a.enoides 1'Upestris demonstrated seasonal movement to shallower water (ea. 750 m) during winter. Nezumia bairdii, N. aeqtwli$, and Cory­ phaenoides carapi1!lUl exhibited a significant positive correlation between head length and depth (,-2 = 0.47, 0.37, and 0.35, respectively). Neru,7IIia bairdi-i apparently spawns in July or August, and reaches an age ofabout 11 years. New size records were established for Ne1!lt1nia aequalis (64 mm head length (HL» and N. bairdii (66 mm HL). New depth records were established for Coelorin{"hlUl c. carlllinatus and N. aeqtwli$ (884 and 1,109 m. respectively). The known geographic ranges for CoelorinrhlUl carib­ beus, C. occa, Ne:ul1lia cyrano, Corypha{Bathygadus macrO'ps, Macrourull IJergla;e, and GadonlUB diBpa.r were extended to the Norfolk Canyon area.

The Macrouridae (Pisces: Gadiformes) includes some important in the western North Atlantic. Experi­ of the most abundant archibenthic deep-sea fish mental commercial trawling was initiated by the species (Marshall 1965, 1971; Marshall and Iwamoto Soviet Union in 1962, and many studies directly 1973; Iwamoto and Stein 1974) and attains greatest related to the commercial fishing of macrourids have abundance and diversity on the continental slopes been subsequently published by Soviet workers of the world oceans (Marshall and Iwamoto 1973). (Podrazhanskaya 1967, 1971; Savvatimskii 1971, Present knowledge of the life history and ecology 1972; Grigor'ev 1972) and to a lesser extent by Polish of macrourids has been accrued piecemeal from researchers (Stanek 1971; Nodzinski and Zukowski faunal lists and taxonomic works (Gunnerus 1765; 1971). Gunther 1887; Gilbert and Hubbs 1920; Farron 1924; The present study examines the seasonal distribu­ Iwamoto 1970; Okamura 1970; Marshall and tion and abundance of the macrourid species cap­ Iwamoto 1973; Iwamoto and Stein 1974), or from tured in the Norfolk Canyon area. In addition aspects studies on physiology, anatomy, and life history of age, growth, and reproduction of selected domi­ (Kulikova 1957; Marshall 1965; Phleger 1971; Ran­ nant species are also described. nou 1975; Rannou and Thiriot-Quiereaux 1975; Haedrich and Polloni 1976; McLellan 1977; Merrett MATERIALS AND METHODS 1978; Smith et al. 1979). The meager literature on reproduction and growth of macrourids and other Gear deep-sea anacanthine fishes has recently been reviewed by Gordon (1979). With the advent of in­ The data presented in this paper were obtained creasing expertise in deepwater trawling, some on four cruises to Norfolk Canyon and the adjacent macrourid species, such as Coryphaenoides 'rUpestris open slope to the south (Fig. 1) conducted by the RV and Macrourus berglax, have become commercially Columbus [sel·in (June 1973) and RV James M. GiUis (November 1974, September 1975, January 1976). 'Contribution No. 1226 from the Virginia Institute of Marine On all cruises a 13.7 m semiballoon otter trawl with Science. 1.3 cm (stretched) mesh in the cod end liner and 5.1 2Virginia Institute of Marine Science, College of William and Mary, Gloucester Point, VA 23062; present address: Minerals cm (stretched) mesh in the wings and body was Management Service, U.S. Department of the Interior, 1951 Kidwell employed. Steel "china V" doors at the end of 22.9 Drive, Vienna, VA 22180. ·Virginia Institute of Marine Science, College of William and m bridles were used to permit spreading of the net Mary, Gloucester Point, VA 23062 from a single warp (Musick et al. 1975).

Manuscript accepted March 1985. 35 FISHERY BULLETIN: VOL. 84. No. 1. 1986. FISHERY BULLETIN: VOL. 84, NO. 1

1I7·3(1 !

!! .: roo'.-- ...... i • 0:Eo // 0' ."Ni ....••. l······ ._~.... 1I7"00' 1J.'-~..c.-ooi- -.- - • E; . • ~~ • ,..-;-" --js \ <:• ...../ c.> .,,,/\\", 1i6. E: C:;w ...... 0,' , ... ~ • /0.. ,t • :IIe" ! I ~ 2, /' ./ ,: I I." ... I, f~~· ~1. .: (._- ) ....." ... i ; ,So I .~. • ,. .j. • .(: "~;,, •" i.M \0'· 1I6.~.:::tt-:":','-"'t-:/If---(,...;;~~;:-I~·l--~-.,Jr+-----!~..;.,; -+---(,-....-...7'.'-----I--.-'-ISI----4--H

) II: \ :

7S"Or:! n·lIr:! 7l1"Or:!

FIGURE I.-Map of the Norfolk Canyon study area with station locations indicated.

Sampling Design 1.0-1.5 g under all typical shipboard conditions. The sex and gonadal conditions of freshly captured Norfolk Canyon and an adjacent open slope were specimens were noted. Gonadal samples for histo­ divided into five sampling strata: 75-150 m, 151-400 logical processing were stored in Davidson's preser- . m, 401-1,000 m, 1,001-2,000 m. and 2,001-3,000 m. vative and later mounted using standard paraffin Six stations were then randomly assigned in each techniques. Sections (5 mm) were stained with depth stratum. The duration of all tows in depths Mayer's hematoxylin and eosin counterstain. of <2,000 m was 0.5 h (bottom time). Where the Saccular otoliths and a scale sample were removed depth exceeded 2,000 m, the tow times were ex­ from all Nezu1nia bairdii and stored dry. Represen­ tended to 1 h. Station depth was determined from tative otolith samples were chosen randomly from a sonic precision depth recorder when the net was individuals over the entire size range of fish set and then every 3 min for the duration of the 0.5 captured. h tows (every 6 min for the 1-h tows). Mean station The length-weight relationships for Nezumia bair­ depth was then determined by averaging the 11 d:ii, CQr/jphaenoides armatus, and C. rupestris were resultant values. analyzed using log transformed weights regressed against head length (Fig. 2). Data Collection and Analysis Regression analysis of head length on depth of cap­ ture was performed for each species to determine Head lengths instead of total lengths were any significant change in head length with change measured because macrourids have slender whiplike in depth. Thsting of the hypothesis that (J = 0 for tails that are easily damaged during trawling. The the regression line ascertained whether there was head length (HL) was measured to the closest a significant change of size with changing depth. The millimeter, from the tip of the snout to the posterior coefficient of determination (r) was also calculated edge of the opercle using Helios4 dial calipers. The to determine what proportion ofthe variance of head fish were weighed with an Ohaus dial-a-gram scale. length could be attributed to change in depth. Calibration showed the scale to be accurate within The a posteriori Student-Newman-Keuls analysis of means was used as a second method for inter­ 'Reference to trade names does not imply endorsement by the preting the size/depth relationship. This method National Marine Fisheries Service, NOAA. calculated the mean depth of capture of each head- 36 MIDDLE'lUN and MUSICK: ABUNDANCE AND DISTRIBUTION OF MACROURIDAE length interval, combined the head lengths in subsets oogenesis was evident. The oocytes were fully whose mean depths did not differ significantly from developed in the females and the male testes con­ each other, and defined the constituents of each tained milky-white seminal fluid. subset. Stage 7-Spent. The testes and ovaries were spent. Due to the large size and thickness of the macrou­ The reproductive organs were flaccid and had rid otoliths, standard age determination techniques recently released sperm or eggs. proved unsuccessful (Christensen 1964; McEachran and Davis 1970). Therefore, a thin section was re­ RESULTS AND DISCUSSION moved from each otolith and, using a dissecting microscope, the number ofbands presumed to be an­ Species Accounts nual were counted and recorded. Gonads of the specimens were classified into repro­ Coe/oN"chlts c. carm;nalus (Goode 1880) ductive stages for analysis. The criteria for these stages were as follows: Coelorinckus c. carmina.tus is a relatively shallow water macrourid reported from depths of 89-849 m Stage I-Undeveloped. The gonads were immature (Marshall and Iwamoto 1973). In the study area this and no development was evident. The reproduc­ species was captured in depths of 210-884 m (Fig. tive organs were difficult to distinguish within the 3). Marshall and Iwamoto (1973) reported C. c. car­ body cavity. minatus from northern Brazil to the Grand Banks, Stage 2-Early Immature. The reproductive organs but absent in the Bahama Island chain. The largest had enlarged slightly. The sex could be determined, specimen captured in our study had a head length but no vascularization of the ovaries was apparent. of 70 mm, while Marshall and Iwamoto (1973) The organs of both sexes had a highly translucent reported specimens with 73 mm HL. appearance. During our study, a maximum of 188 individuals Stage 3-Immature. The ovaries were enlarged and and 4 kg of C. c. carminatus were captured in a 0.5-h vascularization had begun. The testes had become trawl. This species also contributed as much as 34.2% discernibly "sausage shaped". The organs of both of the number and 27.8% of the biomass of benthic sexes were opaque. fishes captured in individual samples. Stage 4-Late Immature. The reproductive organs Figure 4 shows the depth distribution of C. c. car­ of both sexes were full size. The ovaries were about m·inatus incremented by 2 mm size groups. A slight 90% vascularized. The testes had become milky increase in head length with increase of depth was white in color. apparent. The slopes of the regression lines were Stage 5-Mature. The reproductive organs were shown to be significantly different from zero. The developed completely. Ovaries were fully vascular­ coefficient of determination ('Thble 1) also showed a ized and had a granular appearance. correlation between head length and depth. There Stage 6-Ripe. Advanced spermatogenesis or was variability among cruises, but this may be at-

d'n = 305

_ 2 •... 2

9~

~ N.rumio bairdii Coryp/HMnoidtJs ormofus Coryp/IfMnoidtJs rrJPBsfris O+--r---,.-r-..,...-~--l o 10 20 30 40 50 60 0 10 20 30 40 50 60 70 eo 90 100 0 30 50 100 150 2 HEADLENGTH I mml

FIGURE 2.-The log (wt) versus head length regressions for Ne.zulIlia bairdii. ClYrIJPkaenoiM.! arlllat·us, and Coryphaenoide.s rupe.stris. 37 FISHERY BULLETIN: VOL. 84. NO.1

A Coe/ormcl/us C carmino/us B Ntlrumio boirdii ,~ "",0 ,<1' ,0' .$) r!' 0" 0' '\~. ,\" ,\t) '\'0 ,,4{ ,,-.' ".; '\rO

Dopth M,n 252 260 210 226 Depth Mi. 270 315 277 310 Iml Mo. 776 750 I 828 I 884 1m.! Mo. 1350 .525 164411470 I

Tomp Mon 47 49 4.3 45 Tomp. Min 4.1 4 I 3.7 3.7 (oC J Mo. II 3 106 II I II 0 1"C I MOl IQO 9.6 9.2 7. I MOdo 9.0 7.4 8.7 78 Modo 5.1 5.1 4.4 5.3

c Nerumio aequo/is D Corypl/otlnD/des ruptJs/r/s

,0 0" 0" 0' ;P ,r!' ,It ,\0{ ~. J1' <:> ,\..' -(' ,\" "t) ,,'0 Depth M ,n 367 578 330 452 Depth Min 636 578 616 828 Im.l Mo. 986 912 1109 884 CmJ MOl 1591 152511108 Jl 6981

Min 45 45 43 4.3 Min 41 45 3.7 4.1 Temp I----+--+-+_~ Tlmp. 1"C1 Mo. 5.7 7.1 7.8 80 C"CI MOl 49 57 4.3 5.0 Modi 4.6 5.3 4.0 4.8

E Coryphotlnoidtl5 caropinu5 F CoryphotJnoidtl5 arma/us

,0 0" p" 0' ,,0 .0" ,0" .IS' ,,4J' ,\...' ",," "\rO '\'" ,," "t) ,,'0

Oepth MI. ~1~03 118911108 Dlpth Mi. 2100 2257 22.50 I m I Mo. [187612642 2767 2679 (m.l Mo. I 12642 3083129201

3.5 2.9 2.5 2.9 2 Tlmp. Man Tlmp. Mon 1--+_2.5---1r=.34 .::2;.;..4-j FIGURE 3.-Minimum and ma."imum depth C·CI Mo. 4.1 4.2 40 39 C"CI Ma. 3.3 2.8 32 I--+=-+:-+--I of capture, with minimum. maximum, and M.do 3.8 39 37 3.8 Modo 1--L_2_.9...J.,.;2:....4-L2=.,;..,J modal temperatures of capture for each species and each cruise.

TABLE 1.-The coefficient of determination «(1) for the change in significantly different subsets; one 10-50 mm HL and head length With change in depth regression lines. the other 51-70 mm HL. Cruises Other macrourids (N. bairdii and N. aequabis) had Jan. June Sept. Nov. Combined high biomass but low numerical abundance at the Species 76-01 73·10 75·08 74-04 cruises deep end oftheir ranges, indicating the presence of Coe/orinchus c. a few large specimens there. This was not the case carminatus 0.006 0.23 0.13 0.44 0.23 for C. c, carrnina.tus (Fig. 5). The occurrence of fish Nezumia aequa/is 0.45 0.15 0.62 0.14 0.37 distributing by size can be obscured if the larger Nezumia baird;; 0.12 0.50 0.44 0.49 0.47 members of the population traverse the entire range. Coryphaenoides The biomass of the species would be elevated at the rupestris 0.04 0.19 0.08 0.11 0.02 C. carapinus 0.59 0.005 0.30 0.37 0.35 shallower depths so that a consistent biomass level C. armatus 0.000 0.05 0.000 0.14 is present throughout the depth range. Comparison of Figure 4 with Figure 5 shows that although the mean depth of capture for this species increased with tributed to sampling artifacts and the relatively nar­ head length. the larger fish occurred over the en­ row depth range (674 m) of this species. tire depth range. This pattern is important because The analysis orvariance showed a significant dif­ it shows that for some fishes the "bigger-deeper" ference in mean depths of the head length groups phenomenon described by Polloni et al. (1979) may (F = 35.9, F(table; a = 0.01) = 1.79). The Student­ really be a "smaller-shallower" phenomenon. A plot Newman-Keuls test divided the group into two of mean fish weight against depth as used by Polloni 38 1000 3300 CotJlorinchus C. corminofus s: 2100 8 2100 c t'" 76-01 JAN. 2600 73-10 JUNE I.zJ 2600 ~ 2400 2400 Z n 548 .. n 435 2200 = c.. 2200 = "s: 2000 t 21 0::: 2000 t 12 = rn = (=i 1100 1100 DEPTH DEPTH ~ 1600 1600 >to c:: 1400 1400 Z ~ .200 1200 (')z 1000 I.zJ .000 > Z lOa 800 c c lOa 600 !!j 400 400 eto B II + ,1,11,11 11!.l 111.111 i, hil'! c:: III -'l 200 200 III!!II 0 Z 10 20 30 40 !l0 60 0 .0 20 30 40 50 60 0 0 "'l 5000 3000 is: 2800 2100 [!; 1:5 2100 75-08 SEPT. 2800 74-04 NOV. c:: :

0 .0 20 30 40 50 ~O 0 '0 20 ;'0 40 50 60 HEADLENGTH I mm) HEADLENGTH (mm) cc ~ FIGURE 4.-Graph of head length versus depth. by cruise, for CoeWrin~kus c. car- the lines each enclose the range n = the number ofspecimens and t = the number minatus. The dot is the mean, the rectangle is the 95% confidence interval, and of trawls. FISHERY BULLETIN: VOL. 84. NO.1 Coe/orinchus c. corminotus

3.0 u • 73-10 UJ 0 74-04 0._ CJ)- • x 6 75-08 •>c 2.0 • ·66 • X 76-01 ~­ 0 • 00' X 6 o o~.ti 0 -l 0 0 10· X • 6 • • • 1,0 • X X • • X X ~ 0 • 0 60 6 0 0 6 6 00 X X

. I

4.0

X X

o 06~~ 3.0 0 0 0 X 0 ~ 0 0 I- 0 0 X CO liD • 0 0 ::I: + 0 6 0 (!) 0 0 >C 6 X 0 X W-2.0 X 0 0' • 6 0 6 0 II ~ II 0 0 0 6 -l 6 6 0

1.0

100 200 300 400 500 600 700 800 900 1000

DEPTH (m)

FIGURE 5.-The distribution of log transformed (log (x + 1)) abundance and weight of CQelorinrh"lls r. rarminat"lls at each station, plotted against depth. et al. (1979) may have a highly positive slope, but N. baird-ii and is found primarily south of the study these data are impossible to interpret without infor­ area (Marshall and Iwamoto 1973). Nezumia ae­ mation about length-frequency patterns with depth. {fItalis attains a head length of at least 53 mm and The temperatures at which C. c. cat"minatus were has a depth distribution of 200-1,000 m. Its captured varied from 4.3° to 1l.3°C (Fig. 6). The average temperature of collection was 7.6°C. FIGURE G.-The temperature range for each species, by cruise. Th dot designates the modal temperature, Ccc. . CoelorinchlUl I Nezumill tUJfJ_lis (Gunther 1878) carminatU8, N.b. . Nez"Ilmia bairdii, N.a. - Nezllmia aequalis, C.' . Coryphaenoicks rllpestris, C.c. • COr1JPhaenoides carapinU8. CJ Nez-u.mia aequali.s is a closely related congener of . C01·yphanwide.s a1'1natlUl. 40 MIDDLETON and MUSICK: ABUNDANCE AND DISTRIBUTION OF MACROURIDAE

JANUARY 76-01 JUNE 73-10 12 12

II II

10 10

9 9 I• 8 • 8

1 1

6 6

5 I 5 1 T 4 1 4 -r 1 .1 I I 2 2

0 0 e.ee. N.b. N.G e.r. e.e. e.a. c.e.c. Nob. N.G. c.,. C.e. C.a. SPECIES SPECIES

SEPTEMBER 75-08 NOVEMBER 74-04 12 12

II II

10 10

9 1• 9 8 8 • 1 7 T 6 6 T 5 • 5 1 11 T 4 4 1 T ·C 1 J. ·C 3 3 1 l' I I l. 2 2

oJ-__,.----,.--r--,.----,.-.., o.L..-.....---.....--_..,....._...-_...---. C.c.c. N.II Na c.,. C.c Ca. C.c.c. N.~ N... c.r. c.c. c... SPECIES SPECIES 41 FISHERY BULLETIN: VOL. 84. NO. 1 geographic range is listed as from the Faroe bank a half hour tow was 76 fish and the greatest biomass to northern Angola in the eastern Atlantic, the per half hour tow was 5.7 kg. Nezumia ba.irdii com­ Mediterranean, and from Davis Straits to northern prised up to 30% of the demersal fish catch in Brazil in the western Atlantic (Marshall and Iwamoto number and up to 15% of the biomass. 1973). In the January plot (Fig. 9), the head length in­ In the Norfolk Canyon area the depth of capture creased slightly with depth. The regression line of ofN. aequaUs was from 330 to 1,109 m. The greatest the mean depth of each head length class showed number in a trawl was 40 in November of 1974, and a positive slope significantly different than zero. By the highest biomass per trawl was 300 g in Septem­ June (Fig. 9) the regression line showed a highly ber 1975. Nezurnia. a.equalis comprised up to 8.9% significant positive slope and three distinct size ofa trawl catch by number and 3.1% by weight. The groups separated by depth were evident. The first analysis of variance of the mean depths of the head group included those fish <30 mm HL, the second length groups gave a F value of 3.32 (p(table; a = group was from 30 to 42 mm HL, and the third group 0.01) = 2.11). The Student-Newman-Keuls analysis was >43 mm HL. The head lengths at the start of showed only one subset, probably because of the low matul;tyfor females (27 mm) and males (32 mm) cor­ sample size. Examination of Figure 7 suggests head respond well with the dividing line between size length increased with depth, and the slope of the line groups one and two, as defined by depth distribu­ was significantly different from zero. tion. Also, N. bairdii females and males can be fully Although its bathymetric range was extensively mature at 44 and 45 mm HL, respectively (Fig. 10). sampled, densities were low and few mature speci­ These values are close to the division between the mens were captured (Fig. 8). These findings are in second and third size groups noted above. The three contrast to the distribution and abundance of its size groups appear to reflect maturity stages as well cogener, N. bairdii, suggesting competitive exclu­ as size differences, and this may contribute to the sion. Alternately, Norfolk Canyon populations ofN. bathymetric differences. The first group consisted aequa./.is may represent expatriation from denser of all immature fish that were not found in deep populations in the Gulf of Mexico or on the Blake water in June. The second group could be termed Plateau. the transitional group because it included fish that The temperature range for N. a.equalis captured were just starting to mature and those more highly in the Norfolk Canyon area was from 4.3° to 8.0°C developed. Since this group included such a diverse (Fig. 6). The average temperature of collection was spectrum of maturity, it encompassed portions of the 5.3°C. depth ranges of both immature and mature fish. The third group consisted ofall mature fish and was not Nezum;a hairdii (Goode and Bean 1877) found in water shallower than approximately 600 m in June. In September, the larger fish had reached NezU'm'ia ba:it'd-ii is a relatively small macrourid their deepest limit, and immature N. bai-t'dii were with a reported head length of up to 60 mm (Mar­ virtually absent deeper than 1,000 m. By November shall and Iwamoto 1973). During our study the head (Fig. 9), the largest fish were returning to shallower lengths varied from 12 to 66 mm with the weight water to complete what appears to be a seasonal of the largest specimen being 295 g. The geographic migration cycle. range of N ba·i·rd·ii extends from the Straits of Examination of histological sections of gonads Florida north to the Grand Banks (Marshall and showed that the only spentN. bait'dii were captured Iwamoto 1973). Nezumia. bairdii is captured com­ on the September cruise. Although no ripe fish were monly between 90 and 183 m in the northern part caught on any cruise, these spent fish suggest that of its range and appears to undergo tropical sub­ N. bairdi.i spawns in July or August, coincident with mergence because it is found primarily between 548 the time when the mature fish are inhabiting their and 731 m in the southern parts of its range. The deepest level. inclusive depth range is 90-2,285 m (Goode and Bean Marshall's (1965) hypothesis concerning reproduc­ 1885; Marshall and Iwamoto 1973). One anomalous tion of certain macrourids states that fertilization catch ata depth of 16.5 m was recorded in Vineyard takes place at the bottom. Subsequently the eggs, Sound (Bigelow and Schroeder 1953), but this was which are buoyant, develop and hatch on their way most likely a discard from a commercial fishing upward to the seasonal thermocline. The larvae then vessel. maintain themselves just below the thermocline, in Within the study area the depth of capture ranged order to take advantage of the plankton that tends from 270 to 1,644 m (Fig. 3). The largest catch in to accumulate there in the density gradient. In con- 42 3000 3000 Nezumio oequolis is 2100 0 2100 0 73-10 JUNE t"' 2600 76-01 JAN. 2600 l.ZJ 2400 2400 ~ n 80 III 2200 n = 19 2200 = "" t II is:: 2000 t = 7 2000 = c: 1100 1100 5 DEPTH DEPTH P:l 1600 1100 > c:tll 1400 1400 Z 1200 1200 ~ C"l 1000 1000 l.ZJ ~ 100 100 0 ~ llll 0 600 600 !tll ! ~ 400 400 ~ !!! iill tll c: 200 200 g Z 50 0 10 20 30 40 50 60 0 0 10 20 30 40 60 "'l '000 3000 Iii: 2100 2100 ~ 2600 75-08 SEPT. 2600 74-04 NOV. g !!! 2400 2400 ~ l.ZJ 2200 n = 46 2200 n = 95 2000 t = 8 2000 t = 10 1100 1100 DEPTH DEPTH 1600 1600 1400 1400 1200 1200 1000 ,Qr~ 1000 100 100 ~ftt~¥O~~U 600 BI 600 400 ii 400 200 200

0 '0 20 10 40 50 60 0 10 20 30 40 50 60 HEADlENGTH ( mm) HEADlENGTH (mm)

"'"Co' FIGURE 7.-Graph of head length versus depth, by cruise, for Nezumia (U!. the lines enclose the range. n = the number of specimens and t = the number q'ltaii8. The dot is the mean, the rectangle is the 95% confidence interval, and of trawls. FISHERY BULLETIN: VOL. 84, NO. I Nezumio oequo/is

U> Z 2.0 UJ ~ • 73-10 U o o 74-04 UJ o 75-08 0._ o U>- x x 76-01 + o 0 o )( o u.. ­ 1.0 or)Jo 0000 00' o o x .J o Ox 0 x a:: x UJ c. 0 CD o o ~ x :::> z

3.0

o 0 o o o • o o I- =2.0 o x :I:~ C) )( o -- o x 0 x x

x 1.0 00 o

o

100 200 300 400 500 600 700 800 900 1000 1100 1200 1300 1400 1500 DEPTH (m)

FIGURE S.-The distribution of log transformed (log (x + 1)) abundance and weight ofNezumia aeqll.alis at each station. plotted against depth. junction with Marshall's hypothesis, the advantages travel through the thermocline and be removed from of the type of seasonal migration suggested by our the area by the more a<;tive surface currents data are twofold. First, the migration concentrates (although egg density could be such that neutral the reproductively mature fish in a limited area buoyancy occurs at the thermocline). If these sug­ thereby increasing the probability of a sexual en­ gestions hold true, it would be expected that the lar­ counter. Second, itallows additional time for develop­ vae would benefit from the high productivity and ment of eggs on their rise to the upper layers. and warmer temperatures of the surface waters and have concurrently lessens the chance that the egg will enhanced growth. AI; productivity declines in the late 44 !OOO Nezumio boirdli" 2800 ::::~ 73-10 JUNE 2600 76-01 JAN. 2600 2400 2400 n = 506 2200 n = 516 2200 2000 t = 14 2000 t = 20 1800 1800 DEPTH DEPTH 1600 1600 1400 1400 1200 1200 1000 1000 800 800 600 600 400 400 200 200

o 10 20 40 50 60 o 10 20 :so 40 50 60 :S000 3000 2800 2800 2600 75-08 SEPT. 2600 74-04 NOV. 2400 2400 n 453 2200 n = 450 2200 = 2000 t = 14 2000 t = 15 1800 .800 DEPTH DEPTH 1600 1600 1400 1400 1200 1200 1000 1000 800 800 600 600 400 400 200 200

o 10 20 :so 40 50 80 o 10 20 :so 40 50 80 HEADLENGTH (mm) HEADLENGTH (mm)

FIGURE 9.-Graph of head length versus depth, by cruise, for Nezumia bairdii. The dot is the mean, the rectangle is the range. n = the number of specimens and t = the number of trawls. FISHERY BULLETIN: VOL. 84. NO. I fall and the larvae become larger, they would drop recruitment of young occurred between the months out of the water column to the bottom. Length fre­ of November and January. No small N. ba:irdii were quencies of N. ba.i·rdi-i (Fig. 11) suggested that captured benthically between the proposed deep­ water spawning time and the shallower January Nerumia bairdii recruitment spike. 6 d' The larger N. bairdii occurred deeper than the small ones (Figs. 9, 12) demonstrating the "larger­ deeper" phenomenon.

4 The age and growth analysis of N. ba:irdi-i presented many problems. Due to the thickness of the sacculus otolith a thin cross section had to be removed from each. After examination of the thin sections, two problems became apparent. First, all ... of the smaller specimens had two hyaline zones. ~ Oi----.-...... -r--r--r----r-... Because the specimens were obtained on the winter en (January; 76-01) cruise, all had hyaline zones around ... the perimeter as expected. There was, in addition, ..i :> a well-defined hyaline zone in the interior of all the ~ 7 ~ ~ otoliths obtained from the smallest fishes available ::E ("27 mm HL). Subsequently two hypotheses were 6 IEE-i proposed: 1) a period of hyaline zone formation (slow 5 I E$:JI growth) occurred between June-July (spawning) and January, and 2) young N. bairdi'i were not available 4 I E$31 to our trawl until the second winter hyaline zone was 5 1c:LJ I forming (age about 1.5 yr). I $1 The first hypothesis was discarded because a period of slow growth within the first 6 mo would I E$3 I have no apparent selective advantage. It should be

O+----.-...... -r-...... -~--,..-~_ noted, however, that since the larvae of N. bairdii o 10 20 50 40 50 10 were probably pelagic, a change from planktonic HEADLENGTH feeding to benthic feeding would have occurred dur­ FIGURE lO.-The gonadal maturity stages plotted against head ing that time. Such an ontogenetic change occurs in length for NezU'fIl.ia bai·rdii. related gadid fishes. Musick (1969) described the

10

60

50 506 453 450

40

~ E 50 ~ c 20

10

o 10 10 10 40 to 10 10 10 10 40 10 10 10 10 10 40 10 10 to zo 10 40 10 10 H.L.lmml

FIGURE n.-Head length frequency distribution for Nezumia bairdii by cruise. The number above each cruise indicates the number of specimens. 46 MIDDLETON and MUSICK: ABUNDANCE AND DISTRIBUTION OF MACROURIDAE ontogenetic transition for Urophtycis chuss and sug­ The second problem was that in the older fish (>4 gested that the transition from pelagic to demersal yr) the outer bands were very difficult to define with adaptations in morphology and behavior occurred any degree of confidence. The percentage of within a period of 12-24 h. This short time span unreadable otoliths increased from about 5% in fish would be unlikely to be reflected in macroscopic <4 yr to about 50% in fish >4 yr. The mean head hyaline band formation. Therefore, the second hy­ length of N. bairdii with four bands was 42.7 mm, pothesis appeared more likely, and led to the con­ the size at the onset of sexual maturity. Growth may clusion that the juvenile N. bairdii remained pelagic have slowed down to compensate for the energy until the second winter and then descended from the needed for reproduction, and produced spatially water column to the bottom where they were close and obscure hyaline zones. Therefore spawn­ captured. ing checks may have had considerable influence on Nezumia bairdi; en Z 3.0 w ~ - 73-10 u [J : 74-04 W 0._ o 75-08 en - 2.0 .. : 76-01 + 0 )( \ alto Cljl-J- It ,§ 0 LL - 10 It_ tJ-o It 0 Oeo 0 o o - ~ _ DC.[J 0 0 ..J 0 - a: 1.0 ~~ [J w [J- - [J o 0 It lD - - It ~ It 11 It :::;) Z -- -

4.0 0 [J 0 [J - 0 It [JlQult- 3.0 '\ It ~ lt~-- 0 Co - [J - [J [J 0 [J --[J 0 - ~- It 0 [J .. :I: t!)+ _ )( 2.0 ~o lIJ- ~g ..J 04 - - 0_ \.0 - .. -It -- - 200 400 600 800 1000 1200 1400 1600 1800 2000

DEPTH (m)

FIGURE I2.-The distribution of log transformed (log (x + 1)) abundance and weight ofNezumia bairdii at each station plotted against depth. 47 FISHERY BULLETIN: VOL. 84, NO.1 the interpretation of the hyaline zones. and Iwamoto 1973), and is found on both sides of Using the length at age data, a Walford growth the North Atlantic. In the eastern North Atlantic transformation graph was plotted (Beverton and it ranges from the Trondhjem area to the Bay of Holt 1957). Instead of calculating the L .., we used Biscay. In the western North Atlantic it is reported our largest specimen (66 mm HL ). The estimate of to occur from Davis Straitto ca. lat. 37°N (Marshall Brody's coefficient (K) obtained from this graph was and Iwamoto 1973), although two specimens (81 and 0.276. Using the Walford graph, the head lengths 100 mm HL) were captured by C. Richard Robins5 for those presumed ages >4 yr could be iteratively at lat. 23°29.8-32.0'N, long. 77°05.5'W. The depth generated. This method gave a maximum age ofap­ distribution of C. rupestris varies from about 180 proximately 11 yr. The von Bertalanffy growth equa­ to 2,200 m (Leim and Scott 1966) with highest abun­ tion for length was dance occurring between 400 and 1,200 m (Marshall and Iwamoto 1973). = O•276 (T+O.16J). L t 66 (L _ e- Coryphaenoides rupestris is rarely used as a food fish in the United States, but the German Rannou (1976) studied the age and growth of a Democratic Republic, the Soviet Union, and Poland congener (N. 8clerorhyncus) that occupies a similar fish commercially for it in the western North Atlan­ depth range in the western Mediterranean. He tic. In 1968, the Soviets recorded a harvest of 30,000 calculated a K coefficient of 0.16 and an L.. of 42.31 tons of C. rupestris off Labrador, Baffin Island, and mm HL. Thus, although this species is smaller than Greenland (Nodzinksi and Zukowski 1971). The N. bairdii, it has a much slower growth rate, prob­ catches ofthis macrourid were reported to increase ably attributable to lower productivity in the western during the second half of the year as the catches of Mediterranean compared with the slope off the mid­ redfish and cod decreased (Savvatimskii 1971). Atlantic coast of the United States (Koblentz-Mishke Coryph.aerwides rupestris was captured in the Nor­ et al. 1970). folk Canyon area at depths of 578-1,698 m (Fig. 3). The length-weight regression for N. bairdii (Fig. Savvatimskii (1971) reported that C. rupestris is 2) was analyzed. The solution of the line for N. bair­ known to form dense aggregations off the coast of dii males was log (weight) = 0.038 (head length) Labrador. In November 1974 an anomalous catch of + 0.083, r2 = 0.810, and for females it was log over 6,000 C. rupestris with a total weight >1,000 (weight) = 0.035 (head length) + 0.216, r2 = 0.760. kg was obtained in a half hour tow in the Norfolk These length-weight relationships are not unlike Canyon area. A random subsample of 1,000 speci­ those summarized by Gordon (1979) for other small mens was examined and no sexually mature fish macrourids (Coelorinch:u8 coelorinchus, C. occa, and were found. Although the head length ranged from Nezumia aequalis). 59 to 110 mm, the length-frequency curve was In summary, larger N. bairdii were captured strongly unimodal at 76 mm. The greatest number deeper and the minimum and maximum depths of and biomass of C. rupestris caught in "normal" half capture off the mid-Atlantic coast were 270 m and hour tows was 128 fish comprising 39% of the in­ 1,644 m. The fish seasonally migrated to deeper dividuals and 68 kg, and representing 65% of the water with the mature fish occurring deeper than total catch by weight. The largest specimen captured immature fish. The males matured at about 45 mm had a head length of 155 mm. HL and the females became mature at 44 mm HL. The head length distribution by depth and by Nezumia bairdii probably spawned pelagic eggs in cruise (Fig. 13) suggested a mass movement of C. July and August and the young apparently remained rupestris toward deeper water during the summer pelagic until the second winter (January), when they months, and a reciprocating movement to shallower first appeared in bottom trawls. The maximum age water in the winter. In January, the majority of C. of N. bairdii was presumed to be 11 yr. The rupestris was captured between 700 and 800 m, temperature range for N. bairdii was from 3.7° to while in June and September there appeared to be 10.0°C, with the average temperature of capture a movement toward deeper water. By November the being 5.3°C (Fig. 6). depths of capture decreased and were similar to those of January, and the slope of the head length­ Coryphaenoides rupestris (Gunnerus 1765) depth regression for C. rupestris was significantly

CorypJuumoidR,s rupestris is a large macrourid that "e. Richard Robins, Rosenstiel School of Marine and Atmospheric reaches a total length of about 100 cm (Savvatim­ Science, Division of Biology and Living Resources, 4600 Ricken­ skii 1971; Nodzinski and Zukowski 1971; Marshall backer Cau~ Miami, FL 33149. 48 3:100 Coryphaenoides rupesfris 1000 2800 ~ 2800 0 t'" 2600 16-01 JAN. 2600 13-10 JUNE l.zJ ~ 2400 2400 Z n 83 l; 2200 n =274 2200 = ... a: t 7 2000 t 8 c::= 2000 = = en (i 1800 1800 DEPTH DEPTH ?'l 1600 1600 i;; c:::: 1400 Z 1400 ~ 0 1200 1200 ~ C'l 1000 1000 l.zJ ~ 800 1 •• 1•••••••••11 • illllAt!11I 800 ~ 0 1 11 ••• • iiUllw,! 51 600 600 ~ 400 400 ~ 200 200 g Z 411 55 65 711 811 95 105 1111 45 55 611 75 811 95 lOll 0 "'l 3000 3000 li: > 2800 2800 C'l 15-08 SEPT. 14-04 NOV. ~ 2600 2600 c::= 2400 2400 n 70 n 306 >a 2200 = 2200 = l.zJ 2000 t = 6 2000 t = 13 1800 1800 DEPTH DEPTH 1&00 1600 1400 1400 8H 1200 1200 ~~~ vBli!! !! Ii 1000 1000 800 800 600 600 Bfti&ile~I!.~.~~~'8 at 400 400 200 200

45 55 611 1lI 85 15 1011. 45 511 65 75 811 .5 105 HEADLENGTH (mm) HEADLENGTH (mm) II>- co FIGURE 13.-Graph of head length versus depth, by cruise, for CCYr1JPhaen.oides and the lines enclose the range. n - the number of specimens and t = the number rupestris. The dot is the mean, the rectangle is the 95% cconfidence interval, of trawls. FISHERY BULLETIN: VOL. 84. NO.1 different from zero. There was no apparent seasonal of 60-62 and 78-80 cm, respectively. Podrazhanskaya's size segregation evident as in Nezumia bai1'dii, but (1971) modal-length data for each area in conjunc­ the graph of numerical abundance against depth also tion with Savvatimskii's (1971) age and growth data indicated a general seasonal movement down slope reveal that the modal-length fish off the Newfound­ in September (Fig. 14). Similar seasonal movements land banks are about 6 yr old, off Baffin Island they have been shown by Savvatimskii (1971) off are 9-10 yr. around Greenland they are 15-16 yr. and Newfoundland. at Iceland they are over 20 yr. Ifa spawning migra­ Females may be mature from about 104 mm HL tion occurs, it does not preclude spawning by some and males from 71 mm HL (Fig. 15). members of the population not undergoing migra­ Podrazhanskaya (1971) supported Zarkharov and tion, thereby accounting for the small percentage of Mokanu's (1970) theory that C. rupestris spawns in ripening fish to be found outside of their primary Icelandic waters. She stated that C. 1"/tpestris spawn spawning area. near Iceland and the Irminger Current could IfPodrazhanskaya's migration theory is valid, some transport the eggs and larvae to Greenland. From interesting observations can be made. First, the C. Greenland the western branch of the West Green­ 1"/tpestris found on the east coast of the United land Current would transport larvae to Baffin Island States may be derived from the larvae that failed where the Labrador Current would move the fish to metamorphose by the time they reached the New­ down to the Newfoundland banks. When the fish in foundland banks and continued to drift southwest. the Newfoundland area attain a size of 40-50 cm total The predominant currents move south and west from length (TL), they start to migrate back to Iceland. Newfoundland to Cape Hatteras (Worthington 1964; Podrazhanskaya gave the modal lengths for C. Webster 1969; Gatien 1976), thereby affording a rupest1'is in each area. The smallest fish (modal TL means of transport for unmetamorphosed larvae of 45-47 cm) were found on the Northern Newfound­ (Wenner and Musick 1979). Additionally, the modal land bank and the largest (modal TL of 98-100 em) length for the 7,011 C. rupestris caught in the Nor- were found around Iceland. Fish from between Baf­ fin Island and West Greenland had modal lengths

6 d' Ir----E~~" 200 II 400 LO. GI I I " • I E?t=3 I 600 3 I I BOO I e:::t:::3 1000

1200 JAN. ... ~ 0 +---,.--r--...--.--..---..-...----.-...... -...--, 1400 :::,.. III .." ~ 1600 Q JUNE NOV. IBOO 6 -9 SEPT. 2000 I • I I I I 3 II C",ypllunDid.. ,u"..'''6 c:::::j::::J I abundan.. - 109(-'1) I I I I o+---..--r----.--r----,...--.--..---..-....---,.---, 3000...... ------~ ~o ID 70 10 lID 100 II0 IZO 130 140 150 lID HEADLENGTH FIGURE 14.-Diagram of depth plotted against the log transform­ ed (log (s + 1» numerical abundance, by cruise. for C()T'yphae'lloides FIGURE I5.-The gonadal maturity stages plotted against head TUpestris. length for CoryphaffioiM8 rl~pe8tri8. 50 MIDDLETON and MUSICK: ABUNDANCE AND DISTRIBUTION OF MACROURIDAE folk Canyon area was 46 em, exactly that which was tured at 1,108-2,767 m (Fig. 3). The largest number found for C. rupestris in the Newfoundland bank caught in one trawl was 37 (total weight 550 g). area. However, no small C. rupestris were captured These were captured in September 1975 at a depth in the Norfolk Canyon area. We found only 2 fish of 1,803 m. CO't"IJPhaenoides carapinus comprised up with a head length <40 mm (24 cm TL) and only 10 to 23.4% of a catch in number, but only 4.3% in fish with head length <50 mm (30 cm TL). biomass. The maximum size captured was 90 mm The regression line for head length against log HL. (weight) (Fig. 2) was analyzed. The solution for C. CorIJPha.enoides carapinus tended to be larger at rupestris males was log (weight) = 0.023 (head the lower end of its depth range (Fig. 17). The slope length) + 0.82. r2 = 0.898, and for females it was of the regression line for head length with depth was log (weight) = 0.018 (head length) + 1.16, r2 = significantly different than zero. The coefficient of 0.885. determination was 0.346. Unfortunately these length-weight data cannot be Figure 18 displays low numbers and high vari­ compared directly with those summarized by Gor­ ability in the capture of C. ca.ra.pinus in relation to don (1979) because we measured head lengths in our depth. The phenomenon of fewer, larger fish at the study and he gave standard lengths. We do not have deeper part of the bathymetric range was evident the data at present to compute the regression for but obscured because of the relatively small size of head length on standard length for this species. C. ca.rapinus, low numbers, and contagious Temperatures atwhich C. 1"upest't'is were captured distribution. near Norfolk Canyon ranged from 3.7° to 5.7°C (Fig. 6). The average temperature was 4.9°C. CoryphaenO'ides carapinus was taken at temper­ CorIJphaenoides rupest't'is does not follow the atures of 2.5'l.4.2°C with the average temperature "larger-fewer-deeper" pattern shown for N. bai't,­ being 3.7°C (Fig. 6). Some overlap in distribution dii in Norfolk Canyon because it migrates seasonally with depth and temperature occurred among C. (Fig. 16) and the larger specimens traverse the en­ carapinm, C. Q.rrnafm, and C. rupestris. Because tire bathymetric range (Fig. 13). C. ca.rapin-us is a small species and mostly a ben­ In summary, C. ru:pestris migrated seasonally to thic feeder (Haedrich and Polloni 1976) and C. ar­ shallower water in the fall and early winter. Catch 'matus and C. 't"Upestris are large species that forage into the water column (Podrazhanskaya 1971; per unit effort increased in the fall and winter, and a dense aggregation was found in the fall. Podra­ Haedrich and Henderson 1974; Smith et al. 1979), zhanskaya's (1971) spawning and migration theory competitive interaction is probably low. appears feasible but further intensive study is need­ ed. No ripe, running, or spent fish were captured Coryphaenoides armatus (Hector 1875) in the Norfolk Canyon area out of 7,011 individuals examined. There was a trend for the larger C. Coryphaenoides armatus is cosmopolitan in distri­ rupestris to range deeper but not to the degree that bution, being found in all oceans except the Arctic. was found inN. ba.i't'dii. Itappears that the distribu­ It commonly is found from 2,200 to 4,700 m, with tion of C. 'rupestl'is was more closely related to a few specimens being captured as shallow as 282 temperature than to depth, the species being found m (Marshall and Iwamoto 1973). Larger individuals mostly within the 4°_5°C range. have been shown to forage off the bottom for pelagic prey (Haedrich and Henderson 1974; Pearcy 1975; Smith etal. 1979). Coryphaenoides a:rrnatus attains Coryphaenoides campinus (Goode and Bean 1883) a size of 165 mm HL and over 870 mm TL (Iwamoto Coryphaenoides cara.pinus is another small and Stein 1974). The largest specimen captured in macrourid which grows to about 390 mm TL, and Norfolk Canyon was 146 mm HL. Although C. a:r­ is found on the lower slope and abyss from 1,000 to rna.tus is one of the deepest living macrourids, it is 3,000 m (Haedrich and Polloni 1976). In the western rather well-known biologically because of its broad North Atlantic it has been found between Nova distribution and availability to deepwater trawls Scotia and Cape Hatteras (lat 37°N) and in the (Haedrich and Henderson 1974; Pearcy and Ambler eastern Atlantic from lat 50oN to the Equator. Cory­ 1974; McLellan 1977; Smith 1978). phaeno-ides carapinus has also been reported Coryphaenoides armatus was taken in every suc­ from the mid-Atlantic ridge (Marshall and Iwamoto cessful trawl from 2,100 m to our deepest trawl of 1973). 3,083 m in the Norfolk Canyon area and virtually In the Norfolk Canyon area C. cara.pinu..s was cap- was confined to below the 3°C isotherm (Fig. 3). In 51 FISHERY BULLETIN: VOL. 84, NO. I en Coryphaenoides rupestris z LU ~ 3.0- (.) • : 73-10 LU 0 ll._ 74-04 0 en:; 2.0- r:!' .. 75-08 K II 76-01 00 0 u..- 0 0 00 0 00 0 0 0 0 0 ...J 0 1.0 x 0 a:: 0 o • x x 0 LU x 0 0 0 at 0 0 .. 0 0 ~ I ::::J I I I I Z

6.0

5.0 .. x 000 0 4.0 0 0 0 0 II 1-- 0 0 • ::1:- II 0 +- 0 0 ~ 0 M 3.0 0 0 0 0 lLI- II 0 ~g 0 II 0 ...J ... 2.0 0 0 0 1.0

200 400 600 800 '000 1200 1400 '600 1800 2000 DEPTH (m.)

FiGURE 16.-The distribution oflog transfonned (log (x + 1» abundance and weight of Coryp/uumnides TUpestris at each station, plotted against depth. one trawl C. armatus comprised 92.7% of thebentho­ The distribution of numerical abundance. and pelagic fish numbers and 93.4% of the biomass. In weight with depth are shown in Figure 20. Cor­ a 1-h trawl the maximum number captured was 76 yphaenoides armatus increased in abundance from and the maximum biomass was 21.2 kg. 2,100 to 2,600 m, beyond which its abundance re­ No increase in fish size with increased depth was mained constant. evident in the data (Fig. 19)(Thble 1), and the slope The regression lines for head length against log ofthe regression line for head length with depth was (weight) were analyzed (Fig. 2). The solution for not significantly different from zero. However, known males was log (weight) = 0,017 (head length) + depth range of C. armatus was incompletely sam­ 0.956, r2 = 0.967, and for females it was log pled in this study, and further samples from greater (weight) -= 0.016 (head length) + 1.029, r = 0.972. depth may lead to other conclusions. The maturity stages of C. armatus against head 52 3:100 1000 Coryphaenoides corapinus I!:: 2800 2800 73-10 JUNE ~ 2600 76-01 JAN. 2600 l"l 2400 2400 ~ n 40 2200 n 65 .. 2200 = = "l>. t 7 ill: 2000 t = II 2000 = ~ 1800 1800 c:; DEPTH DEPTH ;>::: 1600 ~ ~~i~ 1600 1!llii~ili~i ~ 1400 jlil 1400 ~ 1200 1200 ~ (') 1000 1000 l"l 800 800 § 600 600 Sl 400 400 '"~ D3 200 200 c::: ~ 10 20 30 0 10 20 30 40 50 60 0 40 50 60 ~ 3000 3000 ill: 2800 2800 ~ 75-08 SEPT. 74-04 NOV. ~ 2600 2600 c::: :>:l 2400 2400 n ~ 2200 n = 76 2200 = 69 l"l 2000 t =10 2000 t = 13 1800 1800 DEPTH DEPTH i .600 HI! 1600 1400 !!!Ii 1400 i I!i Ii! 1200 1200 1000 1000 800 800 600 600 400 400 200 200

0 10 10 SO 40 50 60 0 10 10 30 40 50 60 HEADLENGTH (mill) HEADLENGTH (mm)

C1l c.:l FIGURE 17.-Graph of head length versus depth, by cruise. for Corypkaenoida! and the lines enclose the range. n = the number ofspecimens and t = the number ca:rapinWl. The dot is the mean, the rectangle is the 95% confidence interval, of trawls. FISHERY BULLETIN: VOL. 84, NO. I

Coryphoenoides coropinus

3.0 (I) z w ~ • z '73-10 u o z 14-04 ~ 2.0 4 ,. 15-08 (1)- IL ,. 16-01 --+ >C 0 IL ­ Oeo o • o 0 0 o 0 • 0 0 -oJ II a:: 1.0 [J)IL II 0 0 W x o 0 0 m 0 00 0 ~ 0 II x :::) o 0 0 0 Z x 0 0 0 X X II x 0 • • . I

3.0

0 0 0 0 0 o 0 0 0 0 f 0 0 0 0 II 0 ~ -::2.0 110 x 0 X II 0 x :I: + 00 00 (!) l&J '" II --eo 0 II 0 ~ 0 0 -oJ x 0 1.0 x 0 0 0

1200 1400 1600 1800 2000 2200 2400 2600 2800 3000

DEPTH (m)

FIGURE lB.-The distribution of log transformed (log (x + 1) abundance and weight of Co-ryphaenoides cara-pilluS at each station plotted against depth. lengths are shown in Figure 21. No mature males Distribution of Macrourids With were found, but the females matured at about 78 mm Temperature HL. Coryphaen.oides a'Nnatus was captured in temperatures ranging from 2.3° to 3.3°C (Fig. 6). Depth distribution has been used commonly The majority of individuals, however, were caught throughout the literature to delineate the habitat of between 2.4° and 2.9°C during the study and the various fishes, including macrourids (Macpherson average temperature was 2.6°C. 1981). The temperature ranges for each species in 54 Coryphoenoides ormofus 3000· 3000 rs: 2800 2800 a0 t'" 76-01 JAN. 2600 73-10 JUNE t'!l 2600 ~ ii!!il!!!~!i!mi Z 2400 2400 g; 2200 2200 n = 136 rs:"" 2000 c:: 2000 = 5 ~ 1800 1800 ~ DEPTH DEPTH > 160C 1600 til NO SAMPLES c:: 1400 1400 Z !iZ 1200 Z 1200 C'l t'!l 1000 1000 > Z 800 800 0 8 600 600 ~ ::a 400 400 ;; c:: 200 200 :j 0 Z 0 40 ~O 60 70 80 90 100 40 50 60 70 80 90 100 "J 3000 qH~ftij~ ft 3000 rs: 2800 ~ ~ijfi~ 2800 i!1 i;g 2600 2600 & c:: i!! 2400 2400 !iZ ihi'ri'ior t'!l 2200 75 - 08 SEPT. 2200 2000 2000 74-04 NOV. 1800 n = 145 1800 DEPTH DEPTH 1600 t = 4 1600 n = 92 1400 1400 t = 4 1200 1200 1000 1000 800 800 600 600 400 400 200 200

40 lIO 80 70 80 90 100 40 lIO 60 70 80 90 100 HEADLENGTH ( mm) HEADLENGTH (mm) 01 01 FIGURE 19.-Graph of head length versus depth, by cruise, for Co,"yphaenoidn a"· the lines enclose the range. n = the number of specimens and t = the number matus. The dot is the mean, the rectangle is the 95% confidence interval, and of trawls. FISHERY BULLETIN: VOL. 84, NO.1 en Coryphaenoides armafus z • 73-10 ILl 0 74-04 ~ 3.0 0 u 75-08 ILl I : 76-01 0.._ en ;- 2.0 >C dJ. 0 0 u.. - 0 Oeo 0 ...J 1.0 a:: • 0 ILl 0 • lD 0 ~ :::> z

5.0

0 0 4.0 r. • CD • 0 • t- o X ... 3.0 0 C) 0 >C ILl co ~ 0 ...J 2.0

1.0

1600 1800 2000 2200 2400 2600 2800 3000

DEPTH (m)

FIGURE 20.-The distribution of log transformed (log (x+ 1» abundance and weight of C&ryphaenWJ.es anna/us at each station plotted against depth. the present study showed some overlap, but the the deepest trawl of that cruise did not encompass temperatures at which the population modes were the entire range of C. carapinus. Similarly, the found were fairly discrete except for Nezumia (1.6­ minimal temperatures for C. armatus may not be qualis, Nezumia bairdii, and Coryphaenoides representative. rupestris. In Figure 6 the relationship of species with Competition Among Macrourids temperature is more clearly defined. The minimum temperature of each species remained fairly constant Competition among macrourids in the Norfolk as did the maximum and modal temperature for Canyon region is probably minimal because the those species in which there was no indication of species differ in body size and feeding strategies or, seasonal migratory patterns (Coelorinchus car­ iffeeding strategies are similar, the species have dif­ minatus, Coryphaenoides carapinus, C. armatus). ferent distributions with temperature and depth. The 3.5°C minimum temperature found for C. Close congeners such as Nezumia bairdii and N. carapinus in June was probably not accurate since (1.6qualis might be expected to occupy similar depth 56 MIDDLETON and MUSICK: ABUNDANCE AND DISTRIBUTION OF MACROURIDAE

Co,yphoenoides o,_Ius N. bairdii, and CO'ryphaenoides ?'Upestris dominated

6 (Fig. 22). In depths of over 2,000 m the numerical dominance of C. arma.tus was evident. Some of the minor inflections can be attributed to the contagious 4 IEI3-l distributions displayed by these fishes. The graph of macrourid biomass (Fig. 23), as per­ IE$31 cent of the catch, was similar to that for numerical I E$3 I abundance except for a shift in biomass from 800 m to below 1,000 m between January and June. This ... I I I co was probably because of the seasonal movement of :;;c 01---.--..----.--r---.---.-..., the larger macrourid Coryphaenoides 1"1Lpestris. Be­ tween about 1,400 and 2,200 m, macrourids made up a very small portion of the biomass, although their percent by number was comparable with lesser depths. The dominant macrourid in this area, C. I Q::JI carapinus, was small. and Antimora rostrata, a large morid, was the most abundant member of the 4 c:t=:J I benthic fish community from 1,300 to 2,500 m (Wen­ ~ I e:::t::::J ner and Musick 1977). In depths >2,200 m the biomass of C. armatus steeply increased with depth, 1Et3 I until it was the predominant member of the benthic /-EI3I community.

O~_r-.,....---..-.,....--,--T:"'"--, All the macrourid species, with the exception of 40 50 60 70 80 90 100 110 C. rupestris, maintained a fairly constant numerical HEADLENGTH distribution from cruise to cruise. There was ap­

FIGURE 21.-The gonadal maturity stages plotted against head parent variability for C. ca.ru;pinus and C. a.rmatus, length for Coryphaenoides a·1°matus. but this was due to the small number of samples from deeper areas. Distribution of macrourids as the percent of catch revealed a gradual replacement of and temperature ranges; however, the N. aequalis species with depth, and the predominance of C. ar­ in this area were at the northern limit of their matus in depths >2,500 m. geographic range, occurred in small numbers, and may have been in direct competition with N. bair­ Macrourids made up a major numerical portion of dii. Although C. rupestris also occupied the lower the benthic fish community from 300 m to the section of the two Nezumia spp. temperature and deepest station at 3.083 m. Macrourids were also depth regimes, direct competition was probably a main component of the biomass of the commu­ low because of their dissimilarity in mouth size and nities from 300 to 3.083 m, excluding the 1,300­ morphology and related differences in diet 2,500 m range where the morid, A. rostrata, (Podrazhanskaya 1971; Geistdoerfer 1975; McLellan dominated. 1977). Although Macrouridae is a dominant family in the Norfolk Canyon area, the potential for a fishery is Abundance and Density of essentially nonexistent. Cor1jpha.enoides rupestris is the Family Macrouridae the only species which attains an appreciable size in the mid-Atlantic area; a modal length of 46 cm In the study area the abundance of macrourids, TL. However, this size is much smaller than typically in water shallower than 2,000 m, was fairly constant found in the North Atlantic and the density of with respect to other bottom fishes. The average per­ organisms is generally low (normally <0.86 in­ cent of macrourids by number in each cruise was dividuals/l002). In addition, C. r1tpestris demon­ 16.6% in cruise 73-10 (June), 15.0% in 74-04 strates a tropical submergence, being found deeper (December), 14.6% in 75-08 (September), and 18% in lower latitudes. The depth range of this species in 76-01 (January). The major peaks of abundance in the Norfolk Canyon area (578-1,698 m), combined were found between 300 and 400 m, where Coelorin­ with smaller size and lower density of organisms, in­ chus Co carminatus was present, and around 800 m dicate that a commercial fishery would not be where the complex comprised of NeZ1tmia aequalis, economically feasible. 57 FISHERY IlULLETIN: VOL. 84. NO.1

0 JAN. JUNE 2 76-01 73-10 4 6 • 10 12 14 16

18 - 20 22 24 - 26 - 0 28 Q 30 0 SEPT. _ 74-04 NOV. 2 75-08 E 4 -:r I- 6 IL I..t 8 C 10 12 - 14 - 16 18 20 - 22 24 - 26 28 - 30 50 100 o, , , PERCENT

FIGURE 22.-Depth versus relative abundance (as percent. by number. of total capture) for the family Macrouridae, by individual cruise.

Comparison With Other Studies yon area the dominant macrourids (4 species) con­ tributed about 31% to the total catch. The comparison of this study with others in the Merrett and Marshall (1981) remarked on the high North Atlantic lends support to Marshall and diversity (and apparent resource partitioning) of Iwamoto's (1973) hypothesis that the greatest diver­ macrourids from a tropical upwelling area off north­ sity of macrourids is in the bathyal tropical regions. west Mrica and reported 26 species from there. They The number of macrourid species declines from found 18 species on the slope (~1,600 m), including tropical to boreal regions. Marshall and Iwamoto four species of Nezumia. Bathygadine macrourids (1973) reported 32 macrourid species from the Carib­ were important off Mrica but virtually absent in our bean and Gulf of Mexico, but only 22 species were study area. Thus macrourid diversity is probably captured during our study (Thble 2). Bullis and highest on the continental slope in the tropics, par­ Struhsaker (1970) found that Macrouridae was one ticularly in areas of higher productivity. In addition, of the dominant families on the western Caribbean high diversity is manifested there at several tax­ slope between 201 and 400 fathoms (368-732 m). The onomic levels, from the species to the subfamily. deepest stratum sampled was 451-500 fathoms Haedrich etal. (1975) reported the capture of 121 (825-914 m), and macrourids (9 species) comprised macrourid specimens (3 species) in 29 trawls off about 67% of the individuals captured within these Southern New England. Their trawl depths ranged depths. Within the same depths in the Norfolk Can- from 141 to 1,928 m. Their findings were similar to 58 MIDDLETON and MUSICK: ABUNDANCE AND DISTRIBUTION OF MACROURIDAE

0 JAN. JUNE 2 76- 01 73-10 4 6 8 10 12 14 16 18 20 22 0 24 0 26 .. 28

E 30 0 NOV. 75-08 SEPT. 74-04 :z: 2 l- lL 4 lIJ C 6 8

10 12 14 16 18 20 22 24 26

28 30 0 50, IllO, I PERCENT

FIGURE 23.-Depth versus relative abundance (as percent, by biomass, of total capture) for the family Macrouridae, by individual cruise.

TABLE 2.-Species captured during study, with total number and total weight.

Total Total Total weight Total weight Species number (g) Species number (g) Coe!orinchus c. carminatus 1,827 38,597 Coryphaenoides colon' 1 20 Coe/orinchus caribbaeus' 10 419 Coryphaenoides Iepto/ep/s 12 4,922 Coe/orinchus occa' 1 2 ventrifossa occidenta/is 60 1,449 Nezumia aequa/is 285 4,041 V8ntrifossa macropogon 1 8 Nezumia bairdii 2,222 72,865 Hymenocepha/us graci/is' 1 1 Nezumia /ongebarbatus> 12 1,299 Hymenocepha/us ita/icus' 1 12 Nezumia sc/erorhyncus 1 8 Bathygadus favosus 2 Nezumia cyrano' 1 Bathygadus macrops' 1 22 Coryphaenoides rupestris 7,120 1,229,304 Sphagemacrurus granadas' 4 30 Coryphaenoides carap/nus 213 4,703 Macrourus bergiax3 2 4,470 Coryphaenoides armatus 391 120,456 Gadomus dispar' 1

1Range extension from the Gulf 01 Mexico-Caribbean area. 'A1so reported by Haedrich and Polloni (1974). 'Range extension from Boreal Northwest Atlantic.

59 FISHERY BULLETIN: VOL. 84. NO. I those in the present study within the 350-1,100 m better adapted to use the constant abundant food depth interval. Respectively, the family Macrouridae supply there. This speculation is not supported by accounted for 21% and 22.4% ofthe fishes captured data from the southern Sargasso Sea and Bahamas in these depth intervals. (Musick and Sulak unpubI. data), a tropical region Haedrich and Krefft (1978) studied the fish fauna quite low in productivity, in which large rat tails, such in the Denmark Straitand Irminger Sea. In the five as C. armatus, are also very rare. The virtuaI absence fish assemblages that they reported, macrourids of C. armatus from tropical abyssal areas may be were abundant in the 2,026-2,058 m assemblage due instead to some restriction on the life history (22.4%) and very dominant in the 763-1,503 m of the species. Musick and Sulak (1979) have sug­ (48.3%) and 493-975 m (55.4%) assemblages. gested that this species (along with some other large Macrourids were conspicuously absent from their species of predator/scavenger such as C. rupestris group three assemblage, although it was well within and AntimoraTostrata) may migrate to boreal areas macrourid depth and temperature range (280-776 m, to spawn. The tropics may be too far removed from 1.4Q-7.4 Q C). An interesting aspect of Haedrich and such spawning areas for individuals to successfully Krefft's (1978) study was evident in their group two return. assemblage Coryphaenoides rupestris was the highly dominant fish (48.3%) in this group, and the ACKNOWLEDGMENTS temperature range of this group (3.9'!.5.6 Q C) corre­ sponded closely to the temperature range we found We wish to thank all colleagues formerly or pres­ for C. rupest?"is in the present study (3.7 Q-5.7 Q C). ently with the Virginia Institute of Marine Science Pearcy et aI. (1982) summarized data on deep-sea for their enthusiastic participation in the deep-sea benthic fishes collected over several years offOregon program, and particularly to Charles Wenner, (Day and Pearcy 1968; Pearcy and Ambler 1974). Richard Carpenter, Douglas Markle, George Iwamoto and Stein (1974) reported 11 species of Sedberry, and Kenneth Sulak. Daniel Cohen of the macrourids from the northeast Pacific and Pearcy Los Angeles County Museum of Natural History et aI. (1982) recorded 8 of these off Oregon. A com­ kindly contributed cogent comments on early stages parison of these data with ours shows that the of this manuscript. Drafts and final copy of this greatest contrast in the two areas is on the upper report were prepared by the Virginia Institute of and middle slope (500-1,000 m) where five common Marine Science Report Center. species are regularly encountered in the western Atlantic (Coelorinchus c. carminatus, Nemmia bair­ dii, C. aequalis, Coryphaenoides rupestris, and Ven­ LITERATURE CITED trifossa occidentalis), but Pearcy et al. (1982) record­ BEVERrON, R. J. H., AND S. J. HOLT. ed no macrourid as common. This faunal difference 1957. On the dynamics of exploited fish populations. Fish. may be due to the high density off Oregon of scor­ Invest. Minist. Agric. Fish. Food (G.R) Ser. II 19, 533 P. paeniform and lycodine fishes, many of which may BIGELOW, H. R, AND W. C. SCHROEDER. fill niches on the upper slope occupied by macrourids 1953. Fishes of the Gulf of Maine. U.S. Fish Wildl. Serv., elsewhere. The macrourid fauna in depths >2,000 m Fish. Bull. 53, 557 p. BULLIS, H. R., JR., AND P. T. STRUHSAKER. have many similarities to our study. Coryphaenoides 1970. Fish fauna of the Western Caribbean upper slope. Q. armatus becomes increasingly dominant below this J. Fla. Acad. Sci. 33(1):43-76. depth and often is the only species captured deeper CHRISTENSEN, J. M. than 3,000 m in both areas (see also Musick and 1964. Burning ofotoliths, a technique for age determination Sulak 1979). Among other macrourid species Cory­ of soles and other fish. J. Cons. Int. Explor. Mer 29:73-81. DAY, D. S., AND W. G. PEARCY. phaenoides leptolepis is usually second or third in 1968. Species associations of benthic fishes on the continen· abundance at abyssal depths in both regions (Musick tal shelf and slope off Oregon. J. Fish. Res. Board C8lL and Sulak 1979). 25:2665-2675. This distribution pattern is very different from FARRON, G. P. 1924. Seventh report on the fishes of the Irish Atlantic Slope. that reported for the continental rise in the tropics The macrourid fishes (Corgphae'llQididlle). Proc. R. Ir. Acad, off west Africa (Merrett and Marshall 1981) where 36 Sect. B. C. armatus and other large rat tails were very rare. GATIEN, M. G. Marshall and Merrett (1981) speculated that the rari­ 1976. A study in the slope water region south of Halifax. J. ty of large predatory scavengers in the upwelling Fish. Res. Board Can. 33:2213-2217. GEISTDOEFER, P. area they studied might be because of the com­ 1975. Eoologie alimentaie des Macrouridae. 'Ieleosteens Gadi· petitively superior fishes of small size which were formes. Ph.D. Thesis. Univ. Paris. Paris, France, 315 p.

60 MIDDLETON and MUSICK: ABUNDANCE AND DISTRIBUTION OF MACROURIDAE

GILBERr, C. H., AND C. L. HUBBS. Atlantic Mem. Sears Found. Mar. Res., No.1, part 6, p. 1920. The macrourid fishes of the Phillippine islands and the 496-662. East Indies. U.S. Natl. Mus. Bull. 100, 1(pt. 7):369-582. MAcPHERSON, E. GoODE, G. R, AND T. H. BEAN. 1979. Ecological overlap between macrourids in the western 1885. Descriptions of new fishes obtained by the United States Mediterranean Sea. Mar. BioI. (Berl.) 53:149-159. Fish Commission mainly from deep water off the Atlantic McEACHRAN, J. D., AND J. DAVIS. and Gulf Coast. Proc. U.S. Natl. Mus. 8:589-605. 1970. Age and growth of the striped sea-robin. Trans. Am. GoRDON, J. D. M. Fish. Soc 99:343-352. 1979. Phenology of deep-sea anacanthine fishes. In P. J. McLELLAN, T. Miller (editor~ Fish phenology: anabolic adaptiveness in tele­ 1977. Feeding strategies of the macrourids. Deep-Sea Res. osts, p. 327-359. Symp. Zool. Soc. Lond., Acad. Press, N.Y. 24:1019-1036. GRIGOR'EV; G. V. MERRETT, N. R. 1972. Orazmnozhenii tuporylogo rnakrurusa severnoi Atlan­ 1978. On the identity and pelagic occurrence of larval and tiki (Reproduction of Macrurus rupestriB (Gunner) of the juvenile stages of rattail fishes (family Macrouridae) from northern Atlantic). Tr. Polyarn. Nauchno-Issled. Proektn. 60 oN, 200W and 53°N, 20OW. Deep-Sea Res. 25:147-160. Inst. Morsk. Rybn. Khoz. Okeanogr. (PINRO) 289:107­ MERRETT, N. E., AND N. R MARSHALL. 115. (Engl. transl., Fish. Res. Board Can., Transl. Ser. 2529.) 1981. Observations on the biology of deep-sea bottom-living GUNNERUS, J. E. fishes collected off northwest Africa (08'!.27°N). Prog. 1765. Efterretning OM berglaxen, en rar norsk fishk, som Oceanogr. 9:185-244. kunde kaldes: Coryphaenoides rupestriB. K. Nor. Vidensk. MUSICK, J. A. Selsk. Skr. Trondhjem 3(4):40-58. 1969. The comparative biology of two American Atlantic GUNTHER, A. hakes, Urophycis MUSS and U. tenn"is (Pisces: Gadidae). 1887. Report on the deep-sea fishes collected by H.M.S. Ph.D. Thesis, Harvard Univ., Cambridge, 150 p. Challenger during the years 1873-76. Report on the scien­ MUSICK, J. A., AND K. J. SULAK. tific results of the voyage of H.M.S. ChaUenger. Zoology 1979. Characterization of the demersal fish community of a 22:1-335. deep-sea radioactive dump site. Contract Report submitted HAEDRICH, R. L., AND N. R. HENDERSON. to the U.s. Environmental Agency. Va. Inst. Mar. Sci., 61 P. 1974. Pelagic food of Coryphaenoides a7'lnatus, a deep ben­ MUSICK, J. A., C. A. WENNER, AND G. R. SEDBERRY. thic rattail. Deep-Sea Res. 21:739-744. 1975. Archibenthic and abyssobenthic fishes. I" May 1974 HAEDRICH, R. L., AND G. KREFFT. baseline investigations of deep water dumpsite 106 NOAA 1978. Distribution of bottom fishes in the Denmark Strait and evaluation. Rep. 75-1, 388 P. Irminger. Deep-Sea Res. 25:705-720. NODZYNSKI, J., AND C. ZUKOWSKI. HAEDRICH, R. L., AND P. T. POLLONI. 1971. Biological and technological characteristics of grenadier 1976. A contribution to the life history of a small rattail fish, family fishes (Macrouridae) of the Northwestern Atlantic Coryphaenoides carapi7l.us. Bull. South. Calif. Acad. Sci. Studia I Materially Series D, No. 5, p. 1-45. 75:203-211. OKAMURO, O. HAEDRICH, R. L., G. T. RoWE, AND P. T. POLLONI. 1970. Fauna Japonica. Macrourina (Pisces). Acad. Press, 1975. Zonation and faunal composition of epibenthic popula­ 'Ibkyo, 179 p. tions on the continental slope south of New England. J. Mar. PEARCY, W. G. Res. 33:191-212. 1975. Pelagic capture of abyssobenthic macrourid fish. Deep­ IWAMOTO, T. Sea Res. 23:1065-1066. 1970. The RN "Pillsbury" deep sea biological expedition to PEARCY, W. G., AND J. W. AMBLER. the Gulf of Guinea, 1964-65. Macrourid fishes of the Gulf 1974. Food habits of deep-sea macrourid fishes off the Oregon of Guinea. Stud. Trop. Oceanogr. Miami 4:316-431. Coast. Deep-Sea Res. 21:745-759. IWAMOTO, T., AND D. L. STEIN. PEARCY, W. G., D. STEIN, AND R. CARNEY. 1973. A systematic review of the rattail fishes (Macrouridae: 1982. The deep-sea benthic fish fauna of the northeastern Gadiformes) from Oregon and adjacent waters. Occas. Pap. Pacific Ocean on Cascadia and Thfts Abyssal Plains and ad­ Calif. Acad. Sci. No. Ill, 79 P. joining continental slopes. BioI. Oceanogr. 1:375-428. KOBLENTZ-MISHKE, Q J., V. V. VOLKOVINSK~ AND J. G. KABANOVA. PHLEGER, G. F. 1970. Plankton primary production of the world ocean. In 1971. Biology of macrourid fishes. Am. Zool. 11:419-423. W. S. Wooster (editor~ Scientific exploration of the South POKDRAZHANSKAYA, S. G. Pacific, p. 183-193. Natl. Acad. Sci., Wash., D.C. 1967. Feeding of Macrurus rupestriB in the Iceland area. KULIKOVA, E. R Ann. BioI. 24:197-198. 1957. Growth and age of deep water fishes. In R N. Nikitin 1971. Feeding and migrations of the roundnose grenadier, (editor), Marine biology, p. 284-290. 'fr. Inst. Okeanol. Akad. MlU.'TI}lt/'"UB rupestriB, in the Northwest Atlantic and Icelandic Nauk. SSSR 20. [Engl. trans., Am. Inst. BioI. Sci. 1959.] waters. Int. Comm. Northwest Atl. Fish., Redb. Part III, P. LEIM, A. H., AND W. R SCO'IT. 115-123. 1966. Fishes of the Atlantic Coast of Canada. Fish. Res. POLLONI, P., R. HAEDRICH, G. RoWE, AND C. H. CLIFFORD. Board Can., Bull. 155, 485 p. 1979. The size-depth relationship in deep ocean animals. Int. MARSHALL, N. R Rev. Gesamten Hydrobiol. 64:39-46. 1965. Systematic and biological studies of the Macrourid RANNou, M. fishes (Anacanthini!feleostii). Deep-Sea Res. 12:299-322. 1975. Donnees nouvelles sur l-activite reproduetrice cyclique 1971. Explorations in the life of fishes. Harv. Univ. Press, des poissions benthiques bathyaux et abbyssaux. Acad. Sci. Camb., 204 p. Paris, T281, Ser. D, p. 1023-1025. MARSHALL, N. R, AND T. IWAMOTO. 1976. Age et croissance d'un poisson bathyal: Nezumia 1973. Family Macrouridae. In Fishes of the western North sclerorkynchll.B (Macrouridae Gadiformes) de la mer 61 FISHERY BULLETIN: VOL. 84. NO. 1

d~lboran. Cah. BioI. Mar. 17:413-421. biological research. W. H. Freeman and Co., San Franc., 776 RANNOU, M., AND C. 'l'HIRIOT-QmEvREuAX. P. 1975. Structure des otolithes D'un Macrouridae (poisson Gadi­ STANEK, E. forme) Bathyal. Etude au microscope Electronique a 1971. Badania zasobow rybnych wod Labradora i Nowej Balayage. Ann. Inst. Oceanogr., Paris 51:195-201. Fundlandii (Studies on the fish stocks in the waters off SAVVATIMSKII, P. I. Labrador and Newfoundland). Pro Morsk. Inst. Rybackiego, 1971. Studies of the age and growth of roundnose grenadier p. 121-146. (E ngl. trans., Fish. Res. Board Can., Transl. Ser. (MacroltT'lI,s rupestris) in the North Atlantic 1967-1970. Int. 2754.) Comm. Northwest Atl. Fish., Redb. Part III, p. 125-138. WEBSTER, F. 1972. 0 vozraste tuporylogo makrurusa severo-zapadnoi 1969. Vertical profiles of horiwntal ocean currents. Deep­ Atlantiki i vozmozhnom vliyanii promysla no ego chislennost' Sea Res. 16:85-98. (The age of the rock grenadier in the northwest Atlantic and WENNER, C. A., AND J. A. MUSICK. a possible influence of fisheries on its population numbers). 1979. Biology of the morid fish Anti1llora roBtrata in the 'It. Polyarn. Nauchno-Issled. Proektn. Inst. Morsk. Rybn. western North Atlantic. J. Fish. Res. Board Can. 34:2362­ Khoz. Okeanogr. (PINROj 28:116-127. (Engl. transl., Fish. 2368. Res. Board Can., Transl. Ser. 2491.) WORTHINGTON, L. V. SMITH, K. L., JR. 1964. Anomalous conditions in the slope water area in 1978. Metabolism of the abyssopelagic rattail C~dRs 1959. J. Fish Res. Board Can. 21:327-333. armattUl measured ·in situ. Nature (Lond.) 274:362-364. ZAKHAROV, G. P., AND I. D. MOKANU. SMITH, K. L., JR., G. A. WHITE, M. R LAVER, R. R. MCCONNAUGHEY, 1970. Distribution and biological characteristics ofMacrourus AND J. P. MEADOR. rupestris of the Davis Strait in August-September 1969. 1979. Free vehicle capture of abyssopelagic animals. Deep­ Reports of Pinro Marine Expeditionary Investigations, 3rd Sea Res. 26A:57-64. cruise of RN Perseus Ill. SOKAL, R. R., AND F. J. RoHLF. 1969. Biometry: The principles and practice of statistics in

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