17

NOAA Technical Report NMFS 17

Identification and Estimation

of Size From the Beaks of 18

Species of Cephalopods

From the Pacific Ocean

Gary A. Wolff

November 1984

U.S. DEPARTMENT OF COMMERCE

National Oceanic and Atmospheric Administration

National Marine Fisheries Service
NOAA TECHNICAL REPORTS NMFS

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copies may be obtained from: U.S. Department of Commerce, National Technical Information Service, 5285 Port Royal Road,

Springfield, VA 22161.
NOAA Technical Report NMFS 17

Identification and Estimation

of Size From the Beaks of 18

Species of Cephalopods

From the Pacific Ocean

Gary A. Wolff

November 1984

u.s. DEPARTMENT OF COMMERCE

Malcolm Baldrige, Secretary

National Oceanic and Atmospheric Administration

John V. Byrne, Administrator

National Marine Fisheries Service

William G. Gordon, Assistant Administrator for Fisheries
The National Marine Fisheries Service (NMFS) does not approve, recommend or endorse any propriety product or proprietary material mentioned

in this publication. No reference shall be made to NMFS, or to this publication furnished by NMFS, in any advertising or sales promotion which

would indicate or imply that NMFS approves, recommends or endorses any

proprietary product or proprietary material mentioned herein, or which has

as its purpose an intent to cause directly or indirectly the advertised product to be used or purchased because of this NMFS publication.
CONTENTS

Introduction. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1

Methods. . . . .. . . .. . .. . .. . .. .. . .. .. . .. .. . .. . .. .. . .. .. .. 1

Results. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

Key for the upper beak . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

Key for the lower beak . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6

Body weight and mantle length estimates from the beak. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8

Discussion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8

Literature cited. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10

Figures

1. Collection locations of the cephalopods. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11

2. Dimensions measured on the upper and lower beaks. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12

3. Descriptiye characteristics of upper and lower beaks. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13

4. Pigmentation changes with growth in beaks of Onychoteuthis banksii . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14

5. Pigmentation changes with growth in beaks ofAbraliopsis affinis and Abraliopsisfelis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14

6. Pigmentation changes with growth in beaks ofHistioteuthis dofleini . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15

7. Pigmentation changes with growth in beaks ofHistioteuthis heteropsis. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15

8. Pigmentation changes with growth in beaks ofPterygioteuthis giardi and Gonatus onyx . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16

9. Pigmentation changes with growth in beaks ofLeachia danae and Liocranchia reinhardti . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16

10. Pigmentation changes with growth in beaks ofLoligo opalescens . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17

11. Pigmentation changes with growth in beaks of Ornithoteuthis volatilis. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17

12. Pigmentation changes with growth in beaks of Todarodes pacificus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18

13. Pigmentation changes with growth in beaks ofNototodarus hawaiiensis 18

14. Pigmentation changes with growth in beaks ofHyaloteuthis pelagica . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19

15. Pigmentation changes with growth in beaks ofDosidicus gigas. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19

16. Pigmentation changes with growth in beaks of Symplectoteuthis oualaniensis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20

17. Pigmentation changes with growth in beaks ofSymplectoteuthis luminosa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20

18. Pigmentation changes with growth in beaks of Ommastrephes bartramii . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21

19. Upper and lower beaks of Onychoteuthis banksii . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22

20. Upper and lower beaks ofAbraliopsis affinis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23

21. Upper and lower beaks ofHistioteuthis dofleini . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24

22. Upper and lower beaks ofHistioteuthis heteropsis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25

23. Upper and lower beaks ofGonatus onyx. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26

24. Upper and lower beaks ofLeachia danae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27

25. Upper and lower beaks ofAbraliopsisfelis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28

26. Upper and lower beaks ofLoligo opalescens. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29

27. Upper and lower beaks ofLiocranchia reinhardti 30

28. Upper and lower beaks of Pterygioteuthis giardi . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31

29. Upper and lower beaks of Ornithoteuthis volatilis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32

30. Upper and lower beaks of Todarodes pacificus. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33

31. Upper and lower beaks ofNototodarus hawaiiensis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34

32. Upper and lower beaks ofHyaloteuthis pelagica 35

33. Upper and lower beaks ofDosidicus gigas 36

34. Upper and lower beaks of Symplectoteuthis oualaniensis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37

35. Upper and lower beaks of Symplectoteuthis luminosa 38

36. Upper and lower beaks of Ommastrephes bartramii . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39

37. Upper and lower beaks of Thysanoteuthis rhombus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40

38. Upper and lower beak rostral length YS. body weight ofAbraliopsis affinis and Abraliopsisfelis . . . . . . . . . . . . . . . . . . . . . . . . . 41

39. Upper and lower beak rostral length YS. body weight ofPterygioteuthis giardi and Gonatus onyx 42

40. Upper and lower beak rostral length YS. body weight ofLeachia danae and Liocranchia reinhardti . . . . . . . . . . . . . . . . . . . . . . 42

41. Upper and lower beak rostral length YS. body weight ofLoligo opalescens and Onychoteuthis banksii. . . . . . . . . . . . . . . . . . . . . 43

42. Upper and lower beak rostral length YS. body weight ofHistioteuthis dofleini and Histioteuthis heteropsis 43

43. Upper and lower beak rostral length YS. body weight ofNototodarus hawaiiensis and Ommastrephes bartramii . . . . . . . . . . . . . 44

44. Upper and lower beak rostral length YS. body weight ofDosidicus gigas and Symplectoteuthis oualaniensis . . . . . . . . . . . . . . . . 44

45. Upper and lower beak rostral length YS. body weight ofOrnithoteuthis volati/is and Symplectoteuthis luminosa . . . . . . . . . . . . . 45

46. Upper and lower beak rostral length YS. body weight of Todarodes pacificus and Hyaloteuthis pelagica . . . . . . . . . . . . . . . . . . . 45

47. Upper and lower beak rostral length YS. mantle length ofAbraliopsis affinis and Abraliopsisfelis . . . . . . . . . . . . . . . . . . . . . . . . 46

48. Upper and lower beak rostral length YS. mantle length ofPterygioteuthis giardi and Gonatus onyx. . . . . . . . . . . . . . . . . . . . . . . 46

49. Upper and lower beak rostral length YS. mantle length ofLeachia danae and Liocranchia reinhardti . . . . . . . . . . . . . . . . . . . . . 47

iii
50. Upper and lower beak rostral length vs. mantle length ofLoligo opalescens and Onychoteuthis banksii 47

51. Upper and lower beak rostral length vs. mantle length of Histioteuthis dofleini and Histioteuthis heteropsis . . . . . . . . . . . . . . . . 48

52. Upper and lower beak rostral length vs. mantle length ofNototodarus hawaiiensis and Ommastrephes bartramii . . . . . . . . . . . . 48

53. Upper and lower beak rostral length vs. mantle length ofDosidicus gigas and Symplectoteuthis oualaniensis . . . . . . . . . . . . . . . 49

54. Upper and lower beak rostral length vs. mantle length of Ornithoteuthis volatilis and Symplectoteuthis luminosa . . . . . . . . . . . . 49

55. Upper and lower beak rostral length vs. mantle length of Todarodes pacificus and Hyaloteuthis pelagica . . . . . . . . . . . . . . . . . . 50

Tables

I. Upper and lower beak ratio means with standard error of the treatment means. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3

2. Descriptive characteristics of upper and lower beaks . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4

3. Regression equations and r2 values for mantle length and body weight. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9

iv
Identification and Estimation of Size From the Beaks

o! 18 Species of Cephalopods From the Pacific Ocean

GARY A. WOLFf!

ABSTRACT

A method of identifying the beaks and estimating body weight and mantle length of 18 species of cephalopods from the Pacific Ocean is presented. Twenty specimens were selected from each of the following

cephalopod species: Symplectoteuthis oualaniensis, Dosidicus gigas, Ommastrephes bartramii, S. luminosa,

Todarodes pacificus, Nototodarus hawaiiensis, Ornithoteuthis volalilis, Hyaloteuthis pelagica, Onychoteuthis

banksii, Pterygioteuthis giardi, Abraliopsis afJinis, A. felis, Liocranchia reinhard/i, Leachia danae, Histioteuthis

heteropsis, H. dofleini, Gonalus onyx, and Loligo opalescens. Dimensions measured on the upper and lower beak

are converted to ratios and compared individually among the species using an analysis of variance procedure

with Tukey's omega and Duncan's multiple range tests. Significant differences (P =0.05) observed among the

species' beak ratio means and structural characteristics are used to construct artificial keys for the upper and

lower beaks of the 18 species. Upper and lower beak dimensions are used as independent variables in a linear

regression model with mantle length and body weight (log transformed).

INTRODUCTION

The cephalopods are a class of molluscs which contain about

1,000 extant species (Voss 1977). Many of these species are rarely captured in large quantities with conventional sampling gear

since they are generally very adept at avoiding such equipment.

Those cephalopods which are captured are usually only representative of the smaller end of the species' size range.

Cephalopods are regularly captured, however, often in large

quantities and sizes, by many oceanic predators. Confronted by

the limitations imposed by conventional sampling methods for

cephalopods, a number of teuthologists (e.g., Verrill 1879; Joubin

1900; Clarke 1966, 1977; Imber 1978) have used the cephalopods removed from the stomachs of predators to augment sampling of cephalopod populations. Clarke (1977) has discussed the

difference in size range and species composition between netcaught cephalopods and those eaten by a variety of predators.

Predator-collected cephalopods characteristically expand species'

lists and species' size ranges for a given area. The disadvantage of

using cephalopod predators as an alternate sampling method is the

normally poor condition of the cephalopods in the stomachs. In

contrast to other prey such as fish or crustaceans, cephalopods are

usually digested to an unidentifiable condition more rapidly and

completely. Cephalopods have a relatively greater amount of

fleshy tissue directly exposed to the digestive process and a lower

percentage of durable structures which remain after digestion. To

overcome this problem of identification, alternate methods have

been developed to characterize cephalopod prey from the few

durable structures which resist digestion.

The information obtained from different methods of characterizing a cephalopod beak, developed over the last two decades,

has varied widely. Few of these methods enable a specific taxon to

be identified and an associated body weight and length to be

derived from a beak analysis. The result has been that the contribution and importance of cephalopods in predators' diets have

been difficult to accurately estimate. The majority of beak identiITexas A&M University. Environmental Engineering Division. Coliege Station.

TX 77843.

fication studies have used descriptive methoas to separate taxonomic levels of cephalopods. Akimushkin (1955) and Betesheva

and Akimushkin (1955) were the first to use beaks to identify

cephalopods in the stomach contents of cetaceans but neither

described their method of identification. Clarke (1962, 1980)

published two comprehensive studies of cephalopod beak identification keys based on structural features of the beak. Mangold and

Fioroni (1966) separated 18 Mediterranean cephalopod species

on the basis of general beak morphology (6 Octopoda, 12 Teuthoidea). Iverson and Pinkas (1971) and Hotta (1973) published

pictorial guides to cephalopod species from the northeastern and

northwestern Pacific, respectively.

METHODS

An alternate method for identifying cephalopods from beak

characteristics was developed by Wolff (1977) and Wolff and

Wormuth (1979) using beak dimensions. Using this technique a

beak key for eight cephalopod species from the eastern Pacific

was developed (Wolff 1982a) and expanded (Wolff 1982b). The

following presents a cephalopod beak key utilizing beak ratio

comparisons and structural differences among species and the

formulation of equations for estimating body weight and mantle

length using beak dimensions for some species in the Pacific.

The cephalopods for this research were gathered from a variety

of areas (Fig. 1). The species examined were Symplectoteuthis

oualaniensis (S.o.), Dosidicus gigas (D.g.), Ommastrephes bartramii

(0. b.), S. luminosa (S. lum.), Todarodes pacificus (T.pac.),

Nototodarus hawaiiensis (N.haw.), Ornithoteuthis vo/atilis (0. vol.),

Hya/oteuthis pe/agica (H. pel. ), Onychoteuthis banksii (0. bnk.),

Pterygioteuthis giardi (P.gia.), Abraliopsis affinis (A.aff), A. fe/is

(A. fel.), Liocranchia reinhardti (L. rei.), Leachia danae (L.dan.),

Histioteuthis heteropsis (H.het.), H. dofleini (H.dof), Gonatus onyx

(G.ony.), and Loligo opalescens (L.op.). The technique of beak

removal and measurement (Fig. 2) follows that described by

Wolff (1982a, b). The beak dimensions measured on the upper

beak were: Length of the rostrum (RL), rostral tip to inner margin

of wing (RW), length of hood (HL), width of the wing (WW),

wing to crest length (WCL), jaw angle width (JW), and length of
the crest (CL). Dimensions measured on the lower beak were:

Rostral tip to inner posterior corner of the lateral wall (RC),

rostral tip to inner margin of wing (RW), length of the rostrum

(RL), length of the wing (WL), and jaw angle width (JW).

Significant differences among the species' beak ratios were

determined with Tukey's w-procedure and Duncan's new multiple

range test (Steel and Torrie 1960). Combinations of descriptive

characteristics and significant beak ratios are used to identify the

species of cephalopods. Linear regressions were calculated to express the relationship between a beak dimension and the mantle

length and log transformed body weight.

RESULTS

The results of the ANOVA procedure for the beak ratios are

summarized in Table 1. The species' means are ranked by each

beak ratio and the standard error of the treatment mean for each

ratio is given. This table forms the basis for the construction of the

biometric portion of the keys for the upper and lower beaks.

The ratio values in the key represent the midpoints between

species' means. The confidence intervals (Cl) which follow are

derived either from Tukey's method (T) or Duncan's method (D).

When two confidence intervals are given, only the latter

(Duncan's), is significantly different, but both are given for purposes of comparison. Alternate ratios (*) are given at critical

points in the key as well as at the points where species are identified. These alternate ratios are provided for cross reference and

in cases where a specific beak dimension cannot be used (e.g.,

damaged).

Descriptive characteristics of the beak follow those of Clarke

(1980) and Rancurel (1980). The descriptive characteristics are

summarized in Table 2 for each species and are illustrated in

Figure 3. Beak pigmentation patterns at different size ranges are

illustrated in Figures 4 through 18 and referred to in the species

descriptions. Photographs of the species upper and lower beaks

are presented from three different aspects and are also referred to

in the beak key species descriptions (Figs. 19-36). A figure for

Thysanoteuthis rhombus (Fig. 37) is included even though no

measurements were made of the beak. The distinctive shape

should facilitate its identification, however.

Key for the Upper Beak

la. Double anterior-posterior ridge and groove on inner surface of rostrum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 15

1b. Double anterior-posterior ridge and groove absent on

inner surface of rostrum . . . . . . . . . . . . . . . . . . . . . . . . .. 2

2a. Groove at jaw angle. . . . . . . . . . . . . . . . . . . . . . . . . . . .. 4

2b. Groove absent at jaw angle. . . . . . . . . . . . . . . . . . . . . . .. 3

3a. Jaw angle deeply recessed 12

3b. Jaw angle not deeply recessed .. . . . . . . . . . . . . . . . . . .. 5

4a. RLlJW > 1.24 (Cl = 1.35 ± 0.046T) O. banksii

*RLlHL <0.33 (CI = 0.316 ± 0.014T)

*JW/CL <0.184 (Cl = 0.162 ± 0.008T)

Onychoteuthis banksii (Fig. 19). The jaw angle is slightly

recessed and moderately acute; anteriorposterior groove at jaw angle about 1/3 of RL;

two short pigment stripes on inner surface of

crest; wing base inserted about 2/3 down

2

anterior margin of lateral wall; the crest is

moderately curved; the inner margin of the

hood-wing is strongly curved; the outer margin

of the rostrum-hood is strongly curved; pigment

changes with growth are shown in Figure 4.

4b. RLlJW < 1.24 (Cl = 1.128 ± 0.046T) A. affinis

*RLlHL >0.33 (Cl = 0.345 ± 0.014T)

*JW/CL >0.184 (CI = 0.207 ± 0.008T)

Abraliopsis affinis (Fig. 20). The jaw angle is slightly

recessed and roughly square; anterior-posterior

groove at jaw angle about 1/4 of RL or less; two

short pigment stripes on inner surface of crest;

wing base inserted just above base of anterior

margin of lateral wall; the crest is slightly

curved; inner margin of hood-wing is strongly

curved, the outer margin of the rostrum-hood is

moderately curved; pigment changes with

growth are shown in Figure 5.

5a. Double rostral-shoulder edge at jaw angle. . . . . . . . . . .. 6

5b. Single rostral-shoulder edge at jaw angle. . . . . . . . . . . .. 7

6a. RLlJW > 1.281 (CI = 1.355 ± 0.046T; 0.029D) .

....................................... .H. dofleini

*HLlJW >4.078 (Cl = 4.285 ± 0.153T)

*JW/CL <0.194 (Cl = 0.188 ± 0.008T; 0.005D)

Histioteuthis dofleini (Fig. 21). The jaw angle is not

recessed and is obtuse; the shoulder-to-rostral

region of the beak has a double edge; wing base

inserted just above base of interior margin of

lateral wall; the crest is virtually straight; the inner margin of the hood-wing is moderately to

slightly curved; the outer margin of the rostrumhood is moderately curved; pigment changes

with growth are shown in Figure 6.

6b. RL/JW < 1.281 (Cl = 1.207 ± 0.046T) H. heteropsis

*HLlJW <4.078 (CI = 3.872 ± 0.153T)

*JWICL >0.194 (Cl = 0.220 I ± 0.008T; 0.005D)

Histioteuthis heteropsis (Fig. 22). The jaw angle is not

recessed and is obtuse; the shoulder to rostral

region of the beak has a double edge; wing base

inserted about 2/3 down anterior margin of

lateral wall; the crest is slightly curved; the inner

margin of hood wing is moderately to slightly

curved; the outer margin of the rostrum-hood is

moderately curved; pigment changes with

growth are shown in Figure 7.

7a. RLlJW < 1.323 (CI = 1.162 ± 0.046T) 8

*HLlJW <4.261 (CI = 3.846 ± 0.153T)

Wing base inserted 2/3 or more down anterior margin of

lateral wall

7b. RLlJW >1.323 (CI = 1.484 ± 0.046T) G. onyx

*HLlJW >4.261 (CI = 4.676 ± 0.153T)

*JW/CL <0.178 (CI = 0.169 ± 0.008T)

Gonatus onyx (Fig. 23). The jaw angle is not recessed

and is strongly obtuse; wing base inserted about

1/2 down anterior margin of lateral wall; the

crest is virtually straight; the inner margin of

hood-wing is moderately curved; the outer

margin of the rostrum-hood is strongly curved,

particularly in the rostral area; pigment changes

with growth are shown in Figure 8.
Table I.-Ratio means with standard error of the treatment means (Si)'

Species

Si Ratio S. o. S.lum. D. g. O. b. T. pac. N. how. 0. vol. H. pel. A. oj! A.JeI. P. gia. H. hel. H. do! 0. bnk. L. rei. L. dan. G. any. L. opa.

Upper beak

.0118 RLlRW .766 .621 .682 .606 612 .666 .731 .683 .592 .592 .580 .575 .594 .599 .523 .582 .589 .484

.0057 RLlHL .354 .309 .334 .309 .295 .321 .359 .336 .345 .346 .313 .313 .317 .316 .290 .320 .317 .246

.0424 RLlWW 1.506 1.083 1.281 1.111 1.126 1.299 1.658 1.481 1.341 1.264 1.151 1.082 1.156 1.190 0.941 1.198 1.124 0.863

.0062 RLlWCL .358 .314 .354 .319 .296 .329 .382 .341 .306 .304 .287 .310 .331 .271 .261 .302 .327 .211

.0186 RLlJW 1.214 1.265 1.161 1.061 1.119 1.162 1.412 1.162 1.128 1.146 1.042 1.207 1.354 1.349 0.962 1.162 1.483 0.936

.0042 RLlCL .288 .251 .280 .252 238 .260 .290 .265 .234 .237 .226 .243 .253 .218 .211 .235 .250 .176

.0063 RW/HL .463 .498 .491 .509 .481 .482 .492 .492 .583 .568 .542 .543 .540 .528 .557 .550 .539 .509

.0395 RW/WW 1.968 1.740 1.878 1.830 1.823 1.937 2:251 2.147 2.254 2.066 1.979 1.872 1.949 1.980 1.799 2.053 1.906 1.757

.0072 RW/WCL .467 .507 .519 .526 .484 .494 .524 .499 .518 .500 .496 .538 .562 .452 .502 .519 .555 .435

.0388 RW/JW 1.586 2.042 1.705 1.758 1.833 1.751 1.941 1.714 1.916 1.890 1.806 1.103 2.319 2.257 1.850 2.002 2.520 1.954

.0051 RW/CL .376 .404 .411 .416 .389 .391 .398 .389 .396 .390 .391 .422 .431 .364 .405 .405 .425 .365

.0798 HLlWW 4.253 3.498 3.827 3.594 3.788 4.018 4.580 4.370 3.870 3.643 3.660 3.444 3.627 3.756 3.244 3.733 3.539 3.460

.0091 HUWCL 1.010 1.018 1.058 1.033 1.007 1.025 1.065 1.014 0.884 0.881 0.917 0.991 1.042 0.856 0.901 0.945 1.030 0.854

w .0612 HLlJW 3.431 4.104 3.474 3.453 3.811 3.632 3.944 3.479 3.279 3.330 3.332 3.872 4.285 4.277 3.324 3.639 4.676 3.486

.0060 HUCL .813 .812 .837 .817 .808 .811 .808 .791 .677 .688 .721 .777 .798 .689 .728 .736 .789 .718

.0056 WW/WCL .238 .292 .277 .288 .268 .257 .236 .235 .232 .244 .253 .290 .289 .229 .280 .254 .291 .249

.0315 WW/JW 0.811 1.179 0.910 0.966 1.107 0.912 0.877 0.815 0.861 0.926 0.922 1.136 1.195 1.148 1.035 0.983 1.325 1.134

.0046 WW/CL .192 .233 .219 .228 .215 .203 .179 .184 .178 .190 .199 .227 .222 .185 .226 .198 .223 .210

.0729 WCLlJW 3.399 4.032 3.284 3.342 3.791 3.542 3.701 3.345 3.719 3.796 3.641 3.908 4.116 5.014 3.693 3.854 4.544 4.516

.0039 WCLlCL .805 .798 .791 .791 .803 .791 .758 .780 .766 .782 .787 .784 .767 .805 .808 .779 .767 .840

.0031 JW/CL .237 .198 .241 .238 .212 .223 .205 .228 .207 .209 .217 .201 .187 .162 .219 .203 .169 .188

Lower beak

.0124 RC/RW 1.119 1.181 1.232 1.199 1.183 1.169 1.148 1.159 1.209 1.185 1.213 1.200 1.244 1.186 1.142 1.266 1.251 1.235

.0533 RC/RL 2.783 3.071 2.807 2.967 3.220 3.045 2.685 3.035 2.959 3.105 3.424 3.064 3.188 3.223 3.580 3.174 2.907 4.058

.0215 RC/WL 1.755 1.650 1.829 1.700 1.597 1.615 1.724 1.613 1.689 1.618 1.552 1.706 1.751 1.644 1.513 1.792 1.744 1.526

.1252 RC/JW 2.995 4.057 3.357 3.673 3.992 3.519 3.871 3.238 3.852 4.079 3.525 3.741 5.244 3.341 4.402 4.775 8.195 4.025

.0468 RW/RL 2.323 2.599 2.280 2.475 2.721 2.609 2.343 2.618 2.459 2.623 2.828 2.555 2.567 2.722 3.139 2.509 2.330 3.289

.0141 RW/WL 1.465 1.398 1.485 1.418 1.350 1.382 1.501 1.392 1.398 1.365 1.280 1.422 1.406 1.387 1.327 1.416 1.393 1.236

.1054 RW/JW 2.500 3.433 2.727 3.066 3.368 3.016 3.379 2.796 3.179 3.448 2.918 3.121 4.221 2.822 3.867 3.769 6.577 3.258

.0122 RUWL .632 .540 .653 .577 500 .535 .645 .534 .575 .524 .457 .561 .554 .512 .425 .566 .601 .380

.0400 RLlJW 1.077 1.321 1.197 1.243 1.237 1.157 1.439 1.068 1.308 1.316 1.032 1.227 1.653 1.037 1.235 1.506 2.822 0.996

.0830 WUJW 1.709 2.462 1.838 2.168 2.503 2.186 2.265 2.013 2.284 2.527 2.295 2.196 3.023 2.039 2.911 2.671 4.726 2.641
Table 2.-Descriptive characleristics of Ihe beak.

Upper beak

Wing base Double

Innt:r surf:?ce Insertion on Inner margin Rostrum- edge at

of anterior margin Crest of hood shoulder

Jaw angle rostrum-crest of lateral wall curvature hood-wing curvature rostrum -_.._--- u

>

0

e 0

00 c "0 "0 "0

"0 "0 "0 C " .~" " " " E ;;; i: "0 i: "0 i: "0

" e .!'l' u "0 :I

" :I

" :I

" " e > " " " i: " i: " "0

" 0 .0 " u i: '-' u 00 E 0- " C e i: >- :I >- B " ;., :I

e e 00 :g t: c " u 8 :I ;; " ] " ;; 00 >

" :; i: 0 0 E 0 " E >- >- E >- >- " " " ;;; 00 .0 "" 00 "" :! 00 00 c C 0. "" e > " :c .!'l' " " 0 :I :; "" e " '5. " " " .!'l'

"" " c c c

" .!'l' 0 e .E 00 :I 0 0- 00 .0 :! "8 e :! "0 e "8 e " " 0 il 'c 0 "0 ,.., 0 e .0

Q Vi Z 0 0 -< ii: 0 c N :;: 0 E- :;; Z 'c en ::E en en ::E en ::E en -< "'"

S. oualaniensis X X X

D. gigas X

O. banram'-; X

S. luminosa x x

T pacificlls x x x x

N. hawaiiensis x x

H. pelagica x x

O. volatilis x

0. banksii x x

A. affinis x x x

A. felis x

P. giardi x x

L. reinhardti x

L. danae x

L. opalescens

H. dofteini x x x x

H. heteropsis x x

G. onyx x x x x x

Lower beak

Crest-lateral

wall shape

Lateral Ho<>j Hood-wing Rostral upper to inner Crest

Jaw angle wall notch widlh edge view (lOP) fold

"0 "0 :!2 C "0 " ;g u .£ "0 C i: i:

" :I :I

e " " " " e e "0

"0 " il E "" >- E il " " ;g " ..!! 0- C il 0;, 00 e .;;: .0 " -5 .Q 0- " " g 00 .<: c "0 " 0 c -"

" 0 00 "0 -;; " "0 "8 '§ .!'l' e " "0 t: e " " 0 0 C "0 '0 ·z .0 " Z e 0

" " e " :> ii: .0 ::E en Vi en ::E en '" Z z :.: u.. z -< '" Q

"" Z U

S. oualaniensis x

D. gigas

O. banram;; x

S. luminosa x

T pacificlls x

N. hawaiiensis x

H. pelagica x x

0. valotilis x x x

O. banksii

A. affinis x

A. felis

P. giardi x x x x

L. reinhardti x

L. danae x x x x

L. opalescens

H. dofteini x x x

H. heteropsis x x

G. onyx x x X x

4
8a. RLlJW >1.094 (CI = 1.146 ± 0.046T) 9

Jaw angle slightly recessed; crest and rostrum hood not

strongly curved

8b. RLlJW <1.094 (CI= 1.042 ± 0.046T) 10

Jaw angle not recessed and crest straight to slightly

curved or jaw angle slightly recessed and crest strongly

curved

9a. HLlCL>0.712 (CI = 0.737 ± 0.015T) .L. danae

*RLlHL <0.333 (CI = 0.320 ± 0.014T)

*HLlJW >3.484 (CI = 3.639 ± 0.153T)

Leachia danae (Fig. 24). The jaw angle is very slightly

recessed and roughly square; the wing base is inserted just above base of anterior margin of

lateral wall; the crest is slightly curved; the inner

margin of hood-wing is straight to slightly

curved; the outer margin of the rostrum-hood is

moderately curved; pigment changes with

growth are shown in Figure 9.

9b. HLlCL<0.712 (CI = 0.688 ± 0.015T) A.fe/is

*RLlHL >0.333 (CI = 0.346 ± 0.014T)

*HLlJW <3.484 (el = 3.330 ± 0.153T)

Abraliopsis felis (Fig. 25). The jaw angle is slightly

recessed and roughly square; two short pigment

stripes on inner surface of crest; the wing base is

inserted at the base of the anterior margin of the

lateral wall; the crest is virtually straight; the inner margin of the hood-wing is strongly curved;

the outer margin of the rostrum-hood is moderately curved, primarily in the rostral area; pigment changes with' growth are shown in Figure

5.

lOa. RLlCL>0.194(CI=0.211 ±O.OIIT) II

*RLlWCL >0.236 (CI = 0.261 ± 0.016T)

Jaw angle not recessed; crest and rostrum-hood not

strongly curved

lOb. RLlCL <0.194 (CI = 0.177 ± 0.0 lIT) ..... .L. opalescens

*RLlHL <0.268 (CI = 0.246 ± 0.014T)

*HLlJW >3.589 (CI = 3.846 ± 0.153T)

Loligo opalescens (Fig. 26). The jaw angle is moderately

recessed and slightly acute; the wing base is inserted slightly less than 2/3 down the anterior

margin of the lateral wall; the crest is strongly

curved; the inner margin of the hood-wing is

moderately to strongly curved; pigment changes

with growth are shown in Figure 10.

Ila. WCLlCL>0.798 (CI = 0.808 ± O.OIOT) ... .L. reinhardti

*RLlJW <1.002 (CI = 0.963 ± 0.0465T; 0.026D)

*RLlHL <0.301 (CI = 0.290 ± 0.014T; 0.009D)

Liocranchia reinhardti (Fig. 27). The jaw angle is very

slightly recessed and roughly square; the wing

base is inserted 2/3 down the anterior margin of

the lateral wall; the crest is slightly curved; the

inner margin of the hood-wing is straight to

slightly curved; the outer margin of the rostrumhood is moderately curved; pigment changes

with growth are shown in Figure 9.

lIb. WCLlCL<0.798 (CI = 0.788 ± O.OlOT) P. giardi

*RLlJW > 1.002 (CI = 1.042 ± 0.046T; 0.026D)

*RLlHL >0.301 (CI = 0.313 ± 0.014T; 0.009D)

5

Pterygioteuthis giardi (Fig. 28). The jaw angle is not

recessed and is obtuse; the wing base is inserted

just above the base of the anterior margin of the

lateral wall; the crest is straight to slightly

curved; the inner margin of the hood-wing is

moderately curved in the hood region; the outer

margin of the rostrum-hood is moderately to

strongly curved; pigment changes with growth

are shown in Figure 8.

12a. RLlJW < 1.287 (CI = 1.162 ± 0.046T) 13

*RLlWCL <0.362 (CI = 0.341 ± 0.015T)

12b. RLlJW >1.287 (CI = 1.412 ± 0.046T) 0. volatilis

*RLlCL >0.278 (CI = 0.290 ± O.OIIT)

*WCLlCL <0.769 (CI = 0.758 ± O.OIOT)

Ornithoteuthis volatilis (Fig. 29). The jaw angle is deeply

recessed with a wide rostral edge; the wing base

is inserted 1/2 down the anterior margin of the

lateral wall; the crest is straight to slightly

curved; the inner margin of the hood-wing is

moderately curved in the wing region; the outer

margin of the rostrum-hood is strongly curved;

pigment changes with growth are shown in

Figure II.

13a. RLlCL >0.249 (CI = 0.260 ± 0.0 lIT) 14

HLlJW <3.772 (CI = 3.633 ± 0.153T)

Crest not strongly curved

13b. RLlCL <0.249 (CI = 0.238 ± 0.0105T) T. pacificus

*RLlHL <0.3080 (CI = 0.295 ± 0.0142T)

*RLlWCL <0.312 (CI = 0.296 ± 0.0155T)

Todarodes pacificus (Fig. 30). The jaw angle is deeply

recessed with a wide rostral edge; the wing base

is inserted 1/2 down the anterior margin of the

lateral wall; the crest is strongly curved; the inner margin of the hood-wing is moderately

curved; the outer margin of the rostrum-hood is

strongly curved; pigment changes with growth

are shown in Figure 12.

14a. HLlCL >0.801 (CI = 0.811 ± 0.015T; 0.009D)

.................................... .N. hawaiiensis

*WCLlCL >0.785 (CI = 0.791 ± O.OlOT; 0.006D)

Nototodarus hawaiiensis (Fig. 31). The jaw angle is

deeply recessed with a moderately wide rostral

edge; the wing base is inserted slightly more than

112 down anterior margin of lateral wall; the

crest is moderately curved; the inner margin of

the hood-wing is moderately curved; the outer

margin of the rostrum-hood is strongly curved;

pigment changes with growth are shown in

Figure 13.

14b. HLlCL<0.801 (CI = 0.791 ± 0.015T;0.009D) .

.......................................H. pelagica

*WCLlCL <0.785 (CI = 0.780 ± O.OIOT; 0.006D)

Hyaloteuthis pelagica (Fig. 32). The jaw angle is deeply

recessed with a wide rostral edge; the wing base

is inserted slightly more than 112 down the

anterior margin of the lateral wall; the crest is

moderately to slightly curved; the inner margin

of the hood-wing is moderately curved; the outer

margin of the rostrum-hood is strongly curved,
particularly in the rostral region; pigment

changes with growth are shown in Figure 14.

15a. RLlCL >0.266 (CI = 0.280 ± 0.0 liT) 16

*RLlHL >0.322 (CI = 0.335 ± 0.014T)

15b. RLlCL<0.266 (CI = 0.252 ± O.OlIT) 17

*RLlHL <0.322 (CI = 0.309 ± 0.014T)

16a. HLlWCL>1.034 (CI = 1.058 ± 0.030T) D. gigas

*WCLlCL <0.798 (CI = 0.791 ± 0.010T; 0.0060)

*HLlCL >0.825 (CI = 0.837 ± 0.015T; 0.0090)

Dosidicus gigas (Fig. 33). The jaw angle is deeply recessed with a narrow rostral edge; two double ridges

and grooves (two prominent pigment stripes in

juveniles) extend from the inner surface of the

rostrum posteriorly onto the inner surface of the

crest (Fig. 4); the wing base is inserted 1/2 down

anterior margin of lateral wall; the crest is

moderately to slightly curved; the inner margin

of the hood-wing is straight; the outer margin of

the rostrum-hood is strongly curved; pigment

changes with growth are shown in Figure 15.

16b. HLlWCL < 1.034 (CI = 1.010 ± 0.023T) . .S. oualaniensis

*WCLlCL >0.798 (CI = 0.806 ± 0.010T; 0.0060)

*HLlCL <0.825 (CI = 0.813 ± 0.015T; 0.0090)

Symplectoteuthis oualaniensis (Fig. 34). The jaw angle is

deeply recessed with a narrow rostral edge; the

two double ridges and grooves (two prominent

pigment stripes in juveniles) extend from the inner surface of the rostrum posteriorly onto the

inner surface of the crest (Fig. 4); the wing base

is inserted slightly less than 2/3 down the

anterior margin of the lateral wall; the crest is

moderately curved; the inner margin of the

hood-wing is moderately curved; the outer

margin of the rostrum-hood is strongly curved;

pigment changes with growth are shown in

Figure 16.

17a. RLlJW >1.163 (CI = 1.265 ±0.046T) S. luminosa

*HLlJW >3.778 (CI = 4.104 ± 0.153T)

*JW/CL <0.218 (CI = 0.198 ± 0.008T)

Symplectoteuthis luminosa (Fig. 35). The jaw angle is

deeply recessed with a moderately wide rostral

edge; two double ridges and grooves (two prominent pigment stripes in juveniles) extend from

the inner surface of the rostrum posteriorly onto

the inner surface of the crest (Fig. 4); the wing

base is inserted 2/3 down the anterior margin of

the lateral wall; the crest is moderately curved;

the inner margin of the hood-wing is strongly

curved; the outer margin of the rostrum-hood is

strongly curved; pigment changes with growth

are shown in Figure 17.

17b. RLlJW < 1.163 (CI = 1.061 ± 0.046T) 0. bartramii

*HLlJW <3.778 (CI = 3.453 ± 0.153T)

*JW/CL >0.218 (CI = 0.238 ± 0.008T)

Ommastrephes bartramii (Fig. 36). The jaw angle is

deeply recessed with a narrow rostral edge; two

double ridges and grooves (two prominent pigment stripes in juveniles) extend from the inner

surface of the rostrum posteriorly onto the inner

6

surface of the crest (Fig. 4); the wing base is inserted 1/2 down anterior margin of the lateral

wall; the crest is moderately curved; the inner

margin of the hood-wing is slightly curved; the

outer margin of the rostrum-hood is moderately

curved; pigment changes with growth are shown

in Figure 18.

Key for the Lower Beak

1a. Prominent ridge on lateral wall . . . . . . . . . . . . . . . . . . .. 2

1b. Prominent ridge on lateral wall absent 6

2a. Jaw angle visible when viewed from side, rostral edge,

particularly anterior end, strongly curved . . . . . . . . . . .. 3

2b. Jaw angle hidden when viewed from side, rostral edge

slightly curved 4

3a. RLlJW > 1.440 (CI = 1.653 ± 0.1 OOT) H. doj1eini

*RC/JW >4.490 (CI = 5.240 ± 0.313T)

*WLlJW >2.610 (CI = 3.020 ± 0.208T)

Histioteuthis doj1eini (Fig. 21). The jaw angle is recessed

and visible in profile; a knob is present at the

jaw angle; a strong ridge on the lateral wall

extends from beneath the hood to just short of

the inner posterior corner of the lateral wall; the

hood has a deep, narrow notch at the crest; the

hood-wing is moderately wide in profile; the

rostral edge is strongly curved, particularly at the

tip; the crest-lateral wall is moderately broad

from a top view and a strong crest fold is present;

pigment changes with growth are shown in

Figure 6.

3b. RLlJW <1.440 (CI = 1.227 ± O.IOOT) H. heteropsis

*RC/JW <4.490 (CI = 3.740 ± 0.313T)

*WLlJW <2.610 (CI = 2.200 ± 0.208T)

Histioteuthis heteropsis (Fig. 22). The jaw angle is not

recessed and is visible in profile; a weak knob is

present at the jaw angle; a strong ridge extends

from beneath the hood across most of the lateral

wall toward the inner posterior corner; the hood

notch is shallow; the hood-wing width is moderate in profile; the rostral edge is strongly curved;

the crest-lateral wall is broad from a top view

and a weak crest fold is present; pigment

changes with growth are shown in Figure 7.

4a. RLlJW>1.172(CI=1.308±0.100T) 5

*RW/JW >3.000 (CI = 3.180 ± 0.263T; 0.1640)

Hood-wing width is narrow

4b. RLlJW<1.172 (CI = 1.037 ± 0.100T) 0. banksii

*RC/JW <3.595 (CI = 3.340 ± 0.313T; 0.2010)

*RW/JW <3.000 (CI = 2.82 ± 0.263T; 0.1670)

Onychoteuthis banksii (Fig. 19). The jaw angle is not

recessed or visible in profile; a strong, broad

ridge extends from beneath the hood across the

lateral wall towards the inner posterior corner;

the hood width is very shallow; the hood-wing

width is moderate; the rostral edge is slightly

curved toward the tip; the crest-lateral wall is

broad from a top view and a strong crest fold is

present; pigment changes with growth are shown

in Figure 4.
5a. RLlWL >0.549 (CI = 0.574 ± 0.031T) A. affinis

*RWIRL <2.540 (CI = 2.460 ± 0.117T; O.077D)

*WLlJW <2.405 (CI = 2.280 ± 0.208T; 0.125D)

Abraliopsis affinis (Fig. 20). The jaw angle is not recessed or visible in profile; a weak ridge extends

from beneath the hood across the lateral wall

towards the inner posterior corner, being most

prominent beneath and just posterior to the

hood; the hood notch is very shallow; the hoodwing width is narrow in profile; the rostral edge

is slightly curved; the crest-lateral wall is broad

from a top view and a weak crest fold is present;

pigment changes with growth are shown in

Figure 5.

5b. RLlWL <0.549 (CI = 0.524 ± 0.03 IT) A. felis

*RWIRL >2.540 (CI = 2.620 ± 0.117T; 0.077D)

*WLlJW >2.405 (CI = 2.53 ± 0.208T; 0.125D)

Abraliopsis felis (Fig. 25). The jaw angle is not recessed

or visible in profile; a weak ridge extends from

beneath the hood across the lateral wall toward

the inner posterior corner, being most prominent

beneath and just posterior to the hood; the hood

notch is absent; the hood-wing width is narrow

in profile; the rostral edge is slightly curved; the

crest-lateral wall is broad from a top view and a

strong crest fold is present; pigment changes

with growth are shown in Figure 5.

6a. Jaw angle not recessed . . . . . . . . . . . . . . . . . . . . . . . . . .. 7

6b. Jaw angle recessed 14

7a. Strong crest fold present; lateral wall fold present. . . . .. 8

7b. Crest fold absent or weak; lateral wall fold absent ..... 10

8a. RC/RL >2.830 (CI = 2.970 ± 0.133T) 9

*RWIWL < 1.460 (CI = 1.420 ± 0.035T)

Strong crest fold present

8b. RC/RL <2.830 (CI = 2.69 ± 0.133T) O. volatUis

*RLlWL >0.6 I I (CI = 0.645 ± 0.031 T)

*RWIWL > 1.460 (CI = 1.500 ± 0.035T)

Ornithoteuthis volatilis (Fig. 29). The jaw angle is not

recessed or visible in profile; a weak fold extends

across the upper 1/3 of the lateral wall; the hood

notch is shallow; the hood-wing width is wide in

profile; the rostral edge is strongly curved; the

crest-lateral wall is broad from a top view and a

weak crest fold is present; pigment changes with

growth are shown in Figure II.

9a. RLlWL >0.555 (CI = 0.577 ± 0.031T) O. bartramii

*RLlJW >1.155 (CI = 1.243 ± O.IOOT; 0.065D)

*RC/JW >3.450 (CI = 3.670 ± 0.313T; 0.198D)

Ommastrephes bartramii (Fig. 36). The jaw angle is not

recessed and is barely visible in profile; a very

weak fold extends across the upper 1/4 of the

lateral wall; the hood notch is moderately deep;

the hood- wing width is wide in profile; the

rostral edge is strongly curved; the crest-lateral

wall width is moderate and a strong crest fold is

present; pigment changes with growth are shown

in Figure 18.

9b. RLlWL <0.555 (CI = 0.534 ± 0.031 T) H. pelagica

*RLlJW <1.155 (CI = 1.068 ± O.IOOT; 0.065D)

7

Hyaloteuthis pelagica (Fig. 32). The jaw angle is not

recessed and is not visible in profile (just visible

in larger beaks); a weak knob is present at jaw

angle; a very weak fold extends across the upper

1/4 of the lateral wall; the hood notch is shallow;

the hood-wing width is moderate in profile; the

rostral edge is strongly curved; the crest-lateral

wall width is broad and a strong crest fold is

present; pigment changes with growth are shown

in Figure 14.

lOa. RLlJW>1.371(CI=1.506±0.106T) 11

*RLlWL >0.511 (CI = 0.566 ± 0.Q31T)

The crest-lateral wall width is narrow

lOb. RLlJW <1.371 (CI = 1.235 ± O.IOOT) 12

*RLlWL >0.511 (CI = 0.457 ± 0.031 T)

The crest-lateral wall width is moderate to wide

lla. RLlJW >2.164 (CI = 2.822 ± O.IOOT) G. onyx

*RC/JW >6.480 (CI = 8.190 ± 0.313T)

*RWIJW >5.175 (CI = 6.580 ± 0.263T)

Gonatus onyx (Fig. 23). The jaw angle is not recessed

and is visible in profile; a weak knob is present

at the jaw angle; the hood notch is absent; the

hood-wing width is very narrow; the rostral edge

is slightly curved; the crest-lateral wall width is

narrow and the crest is sharp but has no fold;

pigment changes with growth are shown in

Figure 8.

lIb. RLlJW <2.164 (CI = 1.506 ± O.IOOT) L. danae

*RC/JW <6.480 (CI = 4.770 ± 0.313T)

*RW/JW <3.770 (CI = 3.770 ± 0.263T)

Leachia danae (Fig. 24). The jaw angle is not recessed

and is visible in profile; a weak knob is present

at the jaw angle; hood notch is absent; the hoodwing width is moderate; the rostral edge is

strongly curved at the tip; the crest-lateral wall

width is moderately narrow and a weak crest

fold is present; pigment changes with growth are

shown in Figure 9.

12a. RLlJW <1.134 (CI = 1.032 ± O.IOOT) 13

*RW/JW <3.565 (CI = 3.260 ± O.263T)

Rostral edge straight with weak crest fold or rostral edge

strongly curved without crest fold

12b. RLlJW >1.134 (CI = 1.235 ± O.IOOT) L. reinhardti

*RW/JW >3.565 (CI = 3.870 ± 0.263T)

*RW/WL >1.305 (CI = 1.330 ± 0.035T; 0.018D)

Liocranchia reinhardti (Fig. 27). The jaw angle is not

recessed and is visible in profile; a weak knob is

present at the jaw angle; the hood notch is absent; the hood-wing width is moderate; the

rostral edge is straight to slightly curved; the

crest-lateral wall width is broad and the crest is

broad without a crest fold; pigment changes with

growth are shown in Figure 9.

13a. RC/RL >3.740 (CI = 4.058 ± 0.135T) ..... .L. opalescens

*RW/RL >3.060 (CI = 3.290 ± 0.117T)

*RLlWL < 0.418 (CI = 0.380 ± 0.031T)

Loligo opalescens (Fig. 26). The jaw angle is not recessed

and is visible in profile; the hood notch is
shallow; the hood-wing width is moderately

wide; the rostral edge is strongly curved particularly at the tip and is often rough (serrated);

the crest-lateral wall width is broad and the crest

is sharp but has no fold; pigment changes with

growth are shown in Figure 10.

13b. RC/RL<3.740 (Cl = 3.424 ± 0.133T) P. giardi

*RW/RL <3.060 (Cl = 2.830 ± 0.117T)

*RL/WL >0.418 (Cl = 0.457 ± 0.03IT)

Pterygioteuthis giardi (Fig. 28). The jaw angle is not

recessed and is visible in profile; a weak knob is

present at the jaw angle; the hood notch is absent; the hood-wing width is very narrow; the

rostral edge is straight; the crest-lateral wall is

broad and the crest is narrow with a weak fold;

pigment changes with growth are shown in

Figure 8.

14a. RWIRL >2.460 (Cl = 2.600 ± 0.117T) 15

*RL/WL <0.586 (Cl = 0.541 ± 0.031T)

Jaw angle visible, lateral wall fold present and crestlateral wall width broad to moderate or jaw angle not

visible, lateral wall fold present and crest-lateral wall

width moderate

14b. RW/RL<2.460(Cl=2.320±0.117T) 17

*RL/WL >0.586 (Cl = 0.632 ± 0.031T)

Jaw angle not visible, weak lateral wall fold and crestlateral wall width broad or jaw angle visible, wall fold

absent and crest-lateral wall width broad

15a. RC/JW >3.755 (Cl = 3.990 ± 0.313T; 0.20ID) .. , . ,. 16

*WL/JW >2.325 (Cl = 2.460 ± 0.208T; 0.1290)

Hood-wing width moderate; jaw visible with deep hood

notch or jaw angle not visible with shallow hood notch

15b. RC/JW<3.755 (Cl = 3.520 ± 0.313T;0.20ID) ..... ,

............... , . , , N. hawaiiensis

*RC/JW <3.755 (Cl = 3.520 ± 0.263T; 0.1670)

*WL/JW <2.325 (CI = 2.190 ± 0.208T; 0.1320)

Nototodarus hawaiiensis (Fig. 31). The jaw angle is

recessed and is visible in profile; a strong knob is

present at the jaw angle; a weak fold extends

across the upper 113 of the lateral wall; the hood

notch is shallow; the hood-wing width is wide;

the rostral edge is strongly curved; the crestlateral wall width is moderate and a strong crest

fold is present; pigment changes with growth are

shown in Figure 13.

16a. RW/WL>1.375 (CI = 1.400 ± 0.035T; 0.0220) .. , ..

................... , ,S. luminosa

*RL/WL >0.520 (CI = 0.541 ± 0.03IT; 0.0190)

Symplectoteuthis luminosa (Fig. 35). The jaw angle is

recessed and visible in profile; a strong knob is

present at the jaw angle; a weak fold extends

across the upper 1/3 of the lateral wall; the hood

is deeply notched; the hood-wing width is

moderate; the rostral edge is strongly curved,

particularly at the tip; the crest-lateral wall

width is moderately broad and a strong crest fold

is present; pigment changes with growth are

shown in Figure 17.

8

16b. RW/WL<1.375 (CI = 1.305 ± 0.035T; 0.0220) .. , ..

.................. , , .T. pacificus

*RL/WL <0.520 (CI = 0.500 ± 0.031T; 0.0190)

Todarodes pacificus (Fig. 30). The jaw angle is recessed

and is scarcely visible in profile; a strong knob is

present at the jaw angle; a weak fold extends

across the upper 1/4 of the lateral wall; the hood

notch is shallow; the hood-wing width is moderate; the rostral edge is strongly curved; the crestlateral wall width is moderate and a strong crest

fold is present; pigment changes with growth are

shown in Figure 12.

17a. RC/WL>1.792 (Cl = 1.829 ± 0.108T; 0.0320) . .D. gigas

*RL/JW >1.137 (CI = 1.197 ± O.IOOT; 0.0590)

Dosidicus gigas (Fig. 33). The jaw angle is recessed and

visible in profile; a knob is present at the jaw

angle; the hood notch is very deep; the hoodwing width is very wide; the rostral edge is

strongly curved, particularly at the tip; the crestlateral wall width is broad and a strong crest fold

is present; pigment changes with growth are

shown in Figure 15.

17b. RC/WL<1.792 (el = 1.756 ± 0.108T; 0.0320), .....

....................................S. oualaniensis

*RLlJW <1.137 (CI = 1.077 ± O.IOOT; 0.0590)

Symplectoteuthis oualaniensis (Fig. 34). The jaw angle is

recessed and is scarcely visible in profile; a weak

fold extends across the upper 113 of the lateral

wall; the hood notch is deep; the hood-wing

width is moderate; the rostral edge is strongly

curved; the crest-lateral wall width is broad and

a strong crest fold is present; pigment changes

with growth are shown in Figure 16.

Body Weight and Mantle Length Estimates

From the Beak

The equations derived from the regression procedure are given

in Table 3 with their respective r2 values. Although some other

beak dimension regressions resulted in higher r2 values than the

rostral length, it was retained due to its durability and frequent use

as one of the ratio variables in the beak key. The weight and mantle length values were plotted against the rostral length values for

each species (Figs. 38-55) and are referred to in each of the size

estimation equations.

DISCUSSION

The primary use of identifying and estimating the size of

cephalopods from their beaks is in stomach content analyses of

their predators. Since the relationships between dimensions for

the species in this study were established from specimens collected primarily by nets, the beaks were in excellent condition. Beaks

which are removed from a predator's stomach will have been subjected to the possibly damaging processes of ingestion and digestion. As these beaks will ordinarily be in poorer condition than

those used to construct the key, other characteristics of the beak,

in addition to the maximum separation of a species' beak ratio

mean, were considered when the key was constructed. Selection of

a beak dimension was based on the dimension's durability under

mechanical and chemical action, the effect such action would
Table 3.-Regression equations and r' values for ML and body weight, beak dimensions in centimeters.

Upper beak Lower beak

Species Manlle lenglh (mm) r' Body weighI (g) r' Mantle lenglh (mm) r' Body weighl (g) r'

S. oualaniensis ML = - 2.17 + CL 105.2 0.95 In wI = 3.7 + In CL 3.1 0.98 ML = -11.93 + RC 115.4 0.96 In wI = 4.7 + In RC 3.2 0.98

ML = -10.9 + RL 382.2 0.81 In wI = 7.6 + In RL 3.2 0.95 ML= 6.98 + RL 392.5 0.93 In wI = 7.8 + In RL 3.0 0.96

D. gigas ML= 65.8 + CL 86.2 0.95 In wI = 4.3 + In CL 2.23 0.97 ML= 68.0 + WL 207.7 0.95 In wI = 4.97 + In RC 2.3 0.95

ML= 41.1 + RL 346.8 0.87 In wI = 7.3 + In RL 2.54 0.91 ML= 44.2 + RL 357.9 0.84 lnwl=7.4 + In RL 2.48 0.91

L. reinhardti ML=- 5.4 + JW 804.7 0.96 In wI = 7.2 + In JW 2.34 0.88 ML= 0.85 + JW 956.8 0.94 In wI = 7.76 + In JW 2.3 '0.88

ML= - 3.2 + RL 806.9 0.94 In wI = 7.0 + In RL 2.22 0.87 ML= - 1.09 + RL 802.2 0.89 In wI = 6.7 + In RL 2.1 0.80

A. affinis ML= 4.1 + CL 63.7 0.93 In wI = 3.3 + In CL 2.86 0.90 ML= 6.3 + RC 77.7 0.95 In wI = 3.8 + In RC 2.5 0.91

ML= 9.1 + RL 216.1 0.87 In wI = 6.0 + In RL 2.2 0.85 ML= 9.8 + RL 192.8 0.88 In wI = 5.5 + In RL 2.1 0.81

O. banksii ML= -22.1 + CL 127.6 0.92 In wt = 9.4 + In RL 3.8 0.93 ML = -22.5 + RC 177.7 0.93 In wI = 4.7 + In RC 3.5 0.94

ML= -31.0 + RL 641.0 0.87 In wi = 9.4 + In RL 3.8 0.93 ML = -28.9 + RL 610.0 0.95 In wi = 9.1 + In RL 3.7 0.89

P. giardi ML= 2.1 + RW230.9 0.76 In wi = 3.8 + In CL 2.75 0.87 ML= 2.3 + RC 121.9 0.76 In wi = 4.5 + In RC 2.7 0.92

ML= 7.3 + RL 289.8 0.62 In wI = 5.8 + In RL 2.04 0.83 ML= 6.2 + RL 331.6 0.41 In wI = 7.6 + In RL 2.6 0.70

O. bartrami; ML= 42.4 + HL 95.8 0.99 In wI = 3.7 + In CL 2.4 0.98 ML= 44.6 + RC 103.5 0.99 In wI = 4.4 + In RC 2.3 0.99

ML= 51.4 + RL 282.4 0.94 In wi = 6.7 + In RL 2.15 0.96 ML= 52.7 + RL 276.1 0.96 In wi = 6.6 + In RL 2.07 0.98

L. opatescens ML = - 5.7 + CL 153.5 0.94 In wI = 6.0 + In RW 2.25 0.80 ML= 6.0 + RW 240.9 0.87 In wI = 4.4 + In RC 1.95 0.76

ML= 42.2 + RL 542.7 0.79 In wi = 5.7 + In RL 1.21 0.65 ML= 32.4 + RL 607.8 0.74 In wI = 6.0 +lnRLI.4 0.58

S. luminosa ML= 1.27 + CL 10 1.6 0.98 In wI = 3.15 + In CL 3.02 0.99 ML= 0.69 + RC 138.8 0.98 In WI = 4.08 + In RC 3.06 0.99

ML= 9.95 + RL 367.3 0.97 In wi = 6.99 + In RL 2.78 0.98 ML= 11.12 + RL 376.1 0.96 In wI = 7.05 + In RL 2.75 0.98

T. pacifiClls ML= 9.60 + CL 94.8 0.98 In wt = 3.26 + In CL 2.88 0.99 ML= 4.31 + RC 134.4 0.99 In wI = 4.15 + In RC 2.92 0.99

ML= 24.3 + RL 342.6 0.96 In wI = 7.02 + In RL 2.56 0.97 ML= 18.53 + RL 374.4 0.97 In wI = 7.19 + In RL 2.64 0.98

N. hawaiiensis ML= 20.85 + CL 54.1 0.93 In wI = 2.96 + In CL 2.50 0.99 ML= 18.72 + RC 76.6 0.94 In wI = 3.75 + In RC 2.56 0.99

ML= 35.65 + RL 165.9 0.91 In wI = 5.85 + In RL 2.02 0.99 ML= 33.55 + RL 186.1 0.91 In'wI = 6.05' + In RL 2.06 0.98

H. petagica ML= 8.44 + CL 82.7 0.90 In wI = 3.04 + In CL 2.62 0.95 ML= 10.49 + RC 109.4 0.91 In wI = 3.83 + In RC 2.56 0.95

ML= 20.65 + RL 243.1 0.87 In wI = 5.26 + In RL 1.89 0.78 ML= 17.81 + RL 285.5 0.86 In wi = 5.87 + In RL 2.12 0.84

H. heteropsis ML = - 4.94 + CL 59.5 0.95 In wi = 3.84 + In CL 3.22 0.99 ML = - 5.28 + RC 80.5 0.96 In wI = 4.77 + In RC 3.19 0.99

ML= 0.74 + RL 214.92 0.93 In wI = 7.84 + In RL 2.88 0.95 ML= 2.04 + RL 205.7 0.94 In wI = 7.43 + In RL 2.64 0.95

H. dofieini ML= 4.45 + CL 41.0 0.98 In wi = 3.61 + In CL 2.65 0.98 ML= 4.25 + RC 53.6 0.98 In wI = 4.30 + In RC 2.65 0.98

ML= 8.41 +RL 134.4 0.97 In wi = 6.70 + In RL 2.36 0.97 ML= 7.69 + RL 145.5 0.97 In wI = 6.96 + In RL 2.44 0.98

A. feUs ML = - 5.22 + CL 105.2 0.98 In wi = 3.22 + In CL 2.67 0.95 ML = - 5.04 + RC 143.5 0.98 In wI = 4.02 + In RC 2.64 0.93

ML = - 5.05 + RL 442.2 0.94 In wI = 6.95 + In RL 2.63 0.90 ML = - 2.66 + RL 405.5 0.93 In wI = 6.58 + In RL 2.49 0.92

L. danae ML= 19.66 + CL 165.1 0.98 In wI = 2.82 + In CL 2.39 0.97 ML= 20.27 + RC 205.2 0.98 In wI = 3.34 + In RC 2.37 0.97

ML= 20.13 + RL 694.3 0.98 In wI = 6.18 + In RL 2.35 0.96 ML= 18.22 + RL 679.4 0.96 In wI = 6.13 + In RL 2.39 0.95

o. vo/atitis ML= -39.81 +CL 123.9 0.96 In wI = 2.69 + In CL 3.16 0.98 ML = -38.56 + RC 166.2 0.94 In wI = 3.65 + In RC 3.15 0.97

ML = -12.96 + RL 360.4 0.95 In wI = 6.16 + In RL 2.65 0.96 ML = -16.96 + RL 388.1 0.93 In wI = 6.29 + In RL 2.66 0.95

G. onyx ML= 8.28 + CL 58.0 0.81 In wi = 2.30 + In CL 2.42 0.92 ML= 8.07 + RC 76.8 0.84 In wI = 2.96 + In RC 2.42 0.93

ML= 15.22 + RL 181.5 0.71 In wi = 4.69 + In RL 1.93 0.80 ML= 12.82 + RL 190.2 0.72 In wi = 4.99 + In RL 2.13 0.82

have on the accuracy of the beak measurement, and the ability to

separate th,e ratio means at a given confidence level (P = 0.05).

Consequently, small dimensions with easily damaged margins

(e.g., RW, WW, upper beak) were excluded from consideration

when the beak key was constructed, even though they might show

very good separation between species' means when used in a ratio

(e.g., RLlRW, upper beak). Larger dimensions with easily damaged margins (e.g., CL, HL) can still provide a reliable measurement

within the variability of the sample since an eroded margin would

represent less of the overall dimension.

A few of the species in the key have members which were

collected from noncontiguous or disperse areas. The known

distribution of Todarodes pacificus is limited to the northwestern

Pacific and that of Nototodarus hawaiiensis to the area around the

Hawaiian Islands. Some of the specimens of Histioteuthis dofleini,

Hyaloteuthis pelagica, and Liocranchia reinhardti were collected in

the South Atlantic, North Atlantic, Indian Ocean, and the China

Sea. Geographical variations in morphometric characteristics of

cephalopod species with either disjunct or widespread distributions is not uncommon (Young 1972; Wormuth 1976; Wolff

1982a). When the use of the key is restricted to the eastern

Pacific, the beak ratios described for the species identification can

be assumed to be conservative, since the inclusion of measurements made from a few species outside this area can only introduce more variability. This would cause the confidence intervals

for the beak ratios to expand and increase the difficulty in separating species. When this key is used outside the eastern Pacific,

9

the ratio means and confidence intervals are subject to change,

particularly in cephalopod species with disjunct distributions. In

either case, full use should be made of the alternate ratio means,

the beak figures, and the descriptive characteristics, in order to

reduce the misidentification of a cephalopod's beak.

The estimation of the species body weight and mantle length

are based on the upper and lower rostral length of the beak. In a

number of cases, other dimensions, which were more representative of the overall length of the beak (CL, HL, RC), resulted in

more accurate estimations of the cephalopod's size. The rostral

length was retained, however, since it is used in most of the ratios

for species determination and is readily available for size

estimates. The rostral length, additionally, is very durable and is

measurable in all but the most severely damaged beaks. The r2

values of the rostral length regressions, were often within a few

hundredths of the best regression estimates using the crest length

or hood length and represent only a minimal loss in accuracy.

The identification of cephalopod beaks can expand our knowledge of species size and distributional patterns. In addition,

cephalopod beak characteristics can provide useful taxonomic

information. The 21 upper beak ratios and 10 lower beak ratios

provide 31 morphometric characteristics which can be used in

conjunction with other, standard characteristics to aid in structuring taxonomic patterns. For example, there are two forms of

Symplectoteuthis oualaniensis which occur in the Pacific and

Atlantic Oceans (Clarke 1966). One matures at a larger size and

has a distinctive light organ on the dorsal mantle surface while the
other form matures at a smaller size and the dorsal light organ is

absent. The forms are generally accepted to be separate species

(the genus is currently under revision, M. RoeleveldZ). Only two

upper beaks from the small form have been measured and do not

provide an adequate representation. It is noteworthy, however,

that the RLlJW beak ratio mean is 1.11 compared with 1.21 for

the same ratio in the large form. The beaks of the smaller form are

further characterized by a much more extensive pigmentation

than the larger form for a given beak dimension. This characteristic coincides with the maturation at a smaller size since beak

pigmentation is related to maturation (Clarke 1980).

ACKNOWLEDGMENTS

I would like to thank John H. Wormuth, Texas A&M University, and Alan D. Hart, Continental Shelf Associates, Tequesta,

Fla., for critically reviewing the manuscript.

I appreciate the help of a number of individuals who aided in

the completion of this research. Clyde F. E. Roper and Mike

Sweeney, Smithsonian Institution, arranged for the loan of a large

portion of the cephalopod samples. David Au and Benson Lee,

Southwest Fisheries Center, also helped with the location of a

number of the cephalopod samples as did George Snyder, Scripps

Institution of Oceanography, who aided in locating the bulk of the

samples. Nancy Voss, Rosenstiel School of Marine and Atmospheric Science, sent a sample of Cranchiidae on short notice.

Sample location was also aided by Richard Young, University of

Hawaii, Richard Brusca and Janet Haig, Allan Hancock Foundation, and James McLean, Los Angeles County Museum of Natural

History.

Funding for this research was provided by NSF grant

DAR-7924779.

LITERATURE CITED

AKIMUSHKIN, I. I.

1955. Nature of the food of the cachalot. Dok!. Akad. Nauk. SSSR 10J:

1139·1140.

BETESHAVA, E. I., and I. I. AKIMUSHKIN.

1955. Food of the sperm whale (PhysetercQlOdon) in the Kurile Islands region.

Tr. Inst. Okeanol. Akad. Nauk. SSSR 18:86-94.

CLARKE, M. R.

1962. The identification of cephalopod "beaks" and the relationship between

2M. A. Roeleveld, South African Museum, P.O. Box 61, Capetown 8000, South

Africa, pers. commun. June 1981.

10

beak size and total body weight. Bull. Br. Mus. (Nat. Hist.) Zool. 8:422-

480.

1966. A review of the systematics and ecology of oceanic squids. Adv. Mar.

BioI. 4:91-300.

1977. Beaks, nets and numbers. Symp. Zool. Soc. Lond. 38:89-126.

1980. Cephalopoda in the diet of sperm whales of the Southern Hemisphere

and their bearing on sperm whale biology. Discovery Rep. 37, 324 p.

HOTTA, H.

1973. Identification of squids and CUtlie fish in the adjacent waters of Japan,

using the characteristics of beaks. Bull. Sekai Reg. Fish. Res. Lab. 43:133-

147.

IMBER, M. J.

1978. The squid families Cranchiidae and Gonatidae (Cephalopoda: Teuthoideal in the New Zealand region. N.Z. J. Zool. 5:445-484.

IVERSON, I. L. K., and L. PINKAS.

1971. A pictorial guide to beaks of certain eastern Pacific cephalopods.

Calif. Dep. Fish Game, Fish Bull. 152:83-105.

10UBIN, L.

1900. Cephalopodes. Result. Camp. Sci. Prince Albert I, 17: 1- I35.

MANGOLD, K., and P. FIORONI.

1966. Morpho10gie et biometrie des manidibules de quelques cephalopodes

mediterraneens. Vie Milieu (Ser. A) 17: 1139-1196.

RANCUREL, P.

1980. Note pour servir a la connaissance de Symplectoteuthis ouaLaniensis

(Lesson 1830) (Cephalopoda, Oegopsida): Variations ontogeniques du bee

superieur. Cahiers de L'lndo-Pacifique 2(2):217-232.

STEEL, R. G. D., and J. H. TORRIE.

1960. Principles and procedures of statistics. McGraw-Hili Book Co., N.Y.,

481 p.

VERRILL, A. E.

1879. Notice of recent additions to the marine fauna of the eastern coast of

North America, No.3. Am. J. Sci. Ser. 3, 117:239-243.

VOSS, G. L.

1977. Present status and new trends in cephalopod systematics. Symp. Zool.

Soc. Lond. 38:49-60.

WOLFF, G. A.

1977. Morphometry and feeding habits of two ommastrephid squid. M.S.

Thesis, Texas A&M University, College Station, 61 p.

1982a. A beak key for eight eastern tropical Pacific cephalopod species with

relationships between their beak dimensions and size. Fish. Bull., U.S. 80:

357-370.

1982b. A study of feeding relationships in tuna and porpoise through the

application of cephalopod beak analysis. Final Tech. Rep. DAR-7924779.

231 p.

WOLFF, G. A., and J. H. WORMUTH.

1979. Biometric separation of the beaks of two morphologically similar

species of the squid family Ommastrephidae. Bull. Mar. Sci. 29:587-592.

WORMUTH, I. H.

1976. The biogeography and numerical taxonomy of the oegopsid squid

family Ommastrephidae in the Pacific Ocean. Bull. Scripps Inst. Oceanogr.,

Univ. Calif. 23, 90 p.

YOUNG, R. E.

1972. The systematics and areal distribution of pelagic cephalopods from the

seas off Southern California. Smithson. Contrib. Zool. 97, 159 p.
20°

20°

0° 20°

0° 20° .1

60° 20°

.:0'1 I I i 140°

140° 100°

Hdof

Slum

S.lum

180°

Tpuc

1400

I M.,e .

O.b Ob LOP~' H.dof I . Slum I .,..... I H.het Lrer .

ire I I I y": Ovol +-Slum HheL I I I I( ·c .. N h H I Obnk PglO L.op . ow .pe Gony

I Aaff Lrei I L.don I

Hpel (giO S.o O.bnk LreiAaff

I So Dg . Dg' P.glo . Hdof So Sc P.gio

Dg

20° 600 100°

)"I I I I

I I $I,m

;-;;:- I lj i W 0,s, I l-t

6~1 I I I I I I I I II I I I I I I 160°

......

......

Figure I.-Collection locations of the cephalopods: Symplecloleulhis oualaniensis (S.o.). S. luminosa (S.lum.), Dosidicus gigas (D.g.). Ommaslrephes bartramii (O.b.), Todarodes pacific us (T.pac.), Nololodarus

hawaiiensis (N.haw .), Omilholeuthis volalilis (0.vol.), Hyaloleuthis pelagica (H.pel.), Abraliopsis affinis (A.atf.), A. felis (A.fel.), Plerygioleuthis giardi (P.gla.), Hislioleuthis heleropsis (H.het.), H. dofleini (H.

dof.), Onycholeuthis banksii (O.bnk.), Liocranchia reinhardti (L.rei.), Leachia danae (L.dan.), Gonalus onyx (G.ony.), Loligo opalescens (L.op.).
UPPER

side view

top view

LOWER

Figure 2.-Dimensions measured on the upper and lower beaks.

12
UPPER LOWER

(b)

/\7C

/6C\

(a)

6B

'iJ

6A-B

lJ

6A

Figure 3.-Descriptlve characteristics of the upper beak (a) JAW ANGLE: lA-recessed, IB-slightly recessed and acute, IC-not recessed and square (90°), ID-obtuse,

IE-groove, IF-thickened rostral edge, IG-double rostral edge in shoulder region; 2A-ridges and grooves (pigment stripes in juveniles) on inner surface of rostrum,

2B-short pigment stripes on inner surface of rostrum; WING BASE INSERTION: 3A-l/2, 3B-2/3, 3C-just above base, 3D-at base; CREST CURVATURE: 4A-slight,

4B-moderate, 4C-strong; HOOD-WING INNER CURVATURE: SA-straight, SB-moderate, SC-strong; ROSTRUM-HOOD CURVATURE: 6A.moderate, 6B-strong.

Lower beak (b) JAW ANGLE: lA-recessed, IB-not recessed, IC-visible, ID-not visible, IE-knob; LATERAL WALL: 2A-ridge, 2B-weak fold, 2C·strong fold; HOOD

NOTCH: 3A-deep, 3B-shallow, 3C-absent; HOOD-WING WIDTH: 4A-wide, 4B-moderate, 4C·narrow; ROSTRAL EDGE CURVATURE: SA-straight, SB-slight,

SC-strong; CREST-LATERAL WALL WIDTH: 6A-broad, 6B-moderate, 6C-narrow; CREST FOLD: 7A-strong, 7B.weak, 7C-absent.

13
Onychoteuthis banksii

ML-40mm

WT- 3 9

O.5cm

ML- 130mm

WT- 67g

I em

Figure 4.-Pigmentation changes with growth in the beaks of Onycholeulhis banksii.

Abra/iopsis affinis Abra/iopsis feiis

a) ML-20mmL-o~·--"'d

WT-O.6g

ML-36mm

WT-4g

O.5cm

Figure 5.-Pigmentation changes with growth in Ihe beaks of Abraliopsis afJinis (a and b) and Abraliopsis felis (c and d).

14
a)

Histioteuthis

ML- 15mm

WT-2g

O.5em

ML- 84 mm

WT-229g

I em

dof/ein/

Figure 6.-Pigmentation changes with growth in the beaks of His/io/eu/his dofleini.

0)

Histioteuthis heteropsis

ML-24mm

WT-6g

0.5 em

ML-71 mm

WT-IOOg

I em

Figure 7.-Pigmentation changes with growth in the beaks of His/io/eu/his he/eropsis.

15
Pteryqioteuthis qiardi Gonotus onyx

.:;".::

." .:"~ ~~:. '.

a)

ML-18mm

WT-O.4g

ML-30mm

WT-1.2 9

:.:::.: : . ........

&l£f

. : :

..

./// :.~;

..

0.5 em

Figure S.-Pigmentation changes with growth in the beaks of Plerygioleulhis giardi (a and b) and Gonalus onyx (c and d).

Leachia danae Liocranchia reinhardfi

a) ML-65mm~ WT- 0.6g

0.5em

e)

0.5em

ML-125mm

WT- 24g

ML-133mm

WT-6g

I em

Figure 9.-Pigmentation changes with growth in the beaks of Letu:hia danae (a and b) and Liocranchia reinhardli (c and d).

16
a)

Lot/go opatescens

ML- 80mm

WT-12g

b)

ML-153 mm

WT- 33g

lem

Figure IOo-Pigmentation changes with growth in the beaks of Loligo opalescenso

Ornithoteuthis volatllis

ML- 70 mm

WT- 8g

lem

ML- 219 mm

WT- 147g

lem

Figure It.-Pigmentation changes with growth In the beaks of Ornitholeulhis volaliliso

17
Todarodes pacificus

0)

ML-120mm

WT-40 9

I em

ML-234mm

WT-278g

I em

Figure 12.-Pigmentation changes with growth In the beaks of Todarodes pacific"s.

Nototodarus hawaiiensis

ML- 73mm

WT-19g

I em

ML-136mm

WT-130g

lem

Figure 13.-P1gmentation changes with growth In the beaks of No/madarus hawaiiensis.

18
Hya/oteuthis pe/agica

ML- 44mm

WT- 3g

O.5em

ML- 91 mm

WT- 22 g

lem

Figure 14.-Pigmentation changes with growth in the beaks of Hyaloteuthis pelagica.

Oosidicus qlqas

ML-196mm

WT-191

lem

ML- 321 mm

WT- 842 9

Figure IS.-Pigmentation changes with growth in the beaks of Dosidicus gigas.

19
Symp/ectotefJthis ofJ%niensis

ML-130mm

WT-79 9

I em

ML-290mm

WT-927 9

lem

b)

a)

Figure 16.-Pigmentation changes with growth in the beaks of Symplecloleulhis oualaniensis.

Symp/ectoteuthis /uminoso

ML- 32mm

WT- I 9

O.5em

ML-180mm

WT-113

I em

Figure 17.-Plgmentation changes with growth in the beaks of Symplecloleulhis luminosa.

20
Ommasfrephes barframii

ML- S5mm

WT-II g

O.5em

ML-165mm

WT-I/Sg

lem

Figure IS.-Pigmentation changes with growth in the heaks of Ommastrephes bartramii.

21
Onychoteuth;s banks;;

a d

b e

I·I

e

1 em

Figure 19.-The upper (a-c) and lower (d-f) beaks of Onychoteuthis banksii.

22

f
Abraliopsis affinis

a

e f

1 em

Figure 20.-The upper (aoc) and lower (dof) beaks or Abraliopsis affinis.

23
H;st;oteuth;s

a

dofle;n;

_____ ---lf

em

Figure ll.-The upper (a.c) and lower (dol) beaks of Histioteulhis dofleini.

24
Hisfiofeufhis

a

heferopsis

......

d

b e

e

1 em

f

Figure 22.-The upper (aoc) and lower (doC) beaks of HistioteuJhis heteropsis.

25
Gonatus onyx

J a

"'.

'.'

,.".'

b

e

1 em

e

Figure 23.-The upper (a-c) aod lower (d-f) beaks of GOllalUS OIlYX.

26
Leachia

a

danae

'.

d

b e

_. e

1 em

!

/

I .. ·

·

·

·

,

I

/ ........ "

f

Figure 24.-The upper (a-c) aod lower (d-f) beaks of Ltachia danae.

27
Abra/;ops;s

a

fe/ ;5

I

\

\

\

)

,

.,

'~~~~~_.-

d

b e

e

1 em

f

Figure 2S.-The upper (a-c) and lower (d.f) beaks of Abraliopsisfelis.

28
Loligo opa/escens

I.-----'-a I.......- ....Jd

'------ ....11

em

Figure 26.-The upper (a.~) and lower (dol) beaks of Loligo opaltsct"s.

29
Liocranchia reinhardti

\

... " '.-.-

d

b

\

\

e

"-- ... c

1 cm

Figure 27.-Tbe upper (a-c) and lower (d.1) beaks of Liocranchia reinhardti.

30
Pterygioteuthis giardi

_~ e

31

Figure 28.-The upper (a-c) ar<\ lower (d·t) beaks of Pterygioteuthi. giardi.

________f ..... .....e

1 em
____> e

'.

Ornithoteuthis

.'

.' . ........

a

b

em

vo/ati/is

d

~--.I f

Figure 29.-The upper (a-c) and lower (d-t) beaks of Omitholeuthis yolatilis.

32
Todarodes pacificus

1 em

Figure 3O.-The upper (a-c) and lower (d-f) beaks of Todarode. pacificu••

33
Nototodarus hawaiiensis

'- __ ... e

f

Figure 31.-The upper (a-c;' and lower (d-f) beaks of NololodaTUS hawaiieosis.

34
Hya/oteuthis pelagica

I-- ~_~_._ d

b e

e

1 em

f

Figure 32.-The upper (a.c) and lower (d-O beaks of Hyaloteuthis pelagica.

35
Dosidicus • g'gas

I-~~~~ e

1 em

Figure 33.-Tbe upper (aoc) and lower (dof) beaks of Dosidicus gigas.

36
Symplectoteuthis oualaniensis

d

f

1 em.

Figure 34.-The upper (a-c) and lower (d-n beaks of SympleclDleulhis Dualaniensis.

37
Symplectoteuthis luminosa

a

J

.....

e

em

Figure 35.-The upper (a-c) and lower (d·n beaks of Symplecloteulhi. lumino.a.

38
Ommastrephes bartram;;

a

'.

'.

" ... '

d

e

em

Figure 36.-The upper (a.c) and lower (d.f) beaks of Ommastrephes bar/ramii.

39
Thysanoteuthis rhombus

I em

Figure 37.-The upper and lower beak of Thysanoteuthis rhombus (ML = 265 mm, 718 g).

40
ABRALIOPSIS AFFINIS ABRALIOPSIS FELIS

0,275 0.275

0.250 0.250

0.225

/'

,

/ ,

/

/ ,

,

/ , .. /

9-'

,

!lo' a I

,

I

I

I

I

I

0.000...._....-_,..........,._-,.._......_,.........,._...,-_..,..._.,..........,_.....-

0.050

0.025

10 12 14 16 16 20 22 24

WEIGHT (G)

:cI- 0.175

e.!)

Z

W 0.150

..J

..J< 0.125

a::

Ien 0.100

o

a::

.....

::!:

() 0.200

.....

0.225

:c 0.175

le.!)

Z

W

..J

..J

<

a::

Ien

0

a:: 0.075

0.050

0.025

0.000

to 12 14 16 16 20 22 24

WEIGHT (G)

.....

::!: 0.20,)

()

.....

Figure 38.-The upper (U) and lower (L) beak rostral length (RL) versus tbe body weight ofAbra/iopsis ajJinis and Abra/iopsisfe/is [URL, observed 0, predicted

-; LRL, observed /',., predicted ---)_

41
PTERYGIOTEUTHIS GIARDI GONA TUS ONYX

0.125

0.275

0.250

0.225

..... 0.200 0

::I: 0.175

IC'

0.150

...J

...J 0.125

c(

a: 0.100

o

a: 0.075

0.025

0.275

0.250

0.225

0.200

0

..... 0.175

::I:

IC' 0.150

Z

W

...J

...J

c(

0.100

en

oa: 0.075

0.050

0.025

O.OOO ...._ ......._ ...........,~....,._......_ ......._,..........,._......_ ......._,.............

10 12 14 16 16 20 22 24

0.000'1,.._......._ ............_....,._...,...._.,-.....,,-....,._......_ ......._,..........,.

6

WEIGHT (G)

Figure 39.-The upper and lower beak rostral length versus the body weight of Pterygioteathis giardi and Gonatus onyx. Symbols as In Figure 38.

lEACHIA DANAE 1I0CRANCHIA REINHARDT!

0.275

0.250

0.225

O.20G

0

::I: 0.175

IC'

Z 0.150 '" W

...J , ,

...J 0.125

c( , . c

a: /

/ c

I- 0.100 , ,

en , • /

0 ; ! ~/ a: 0.075 Ii ,,' • A,' A

, 0.050

.~' ,

0.025

0.000

0 10 12 14 16 16 20 22 24

WEIGHT (G)

10 12 14 16 16 20 22 24

WEIGHT(G)

c

0.000,'<r-_...,...._.,-_,..........,._......_ ......._ ...........,_....,._......_ ............,.

o

0.075

0.275

0.225

0.250

0.025

0.050

::I:

0.175

Z

0.150

...J

c(a: 0.125

Ien

0 0 • 100

a:

.....

o 0.200 .....

Figure 40.-The upper and lower beak rostral length versus the body weight of uachia danae and Liocranchia ninhardli. Symbols as In Figure 38.

42
LOLIGO OPALESCENS ONYCHOTEUTHIS BANKSII

o 5 10 15 20 25 30 35 40 45 50 55 60 65 70

WEIGHT WEIGHT (G) (G)

0.24 ," • 0.23 , ,. D ,

"/. 0.22 .'/

/

/. ,

D /. 0.21 II

+' ,

, D

/. 0.20 , D , . ,. , • /D

I D 0.19 I •• , U ,

" , , .... O. \6 I . , .. • • .r, D

:I: , D D

• , , 0.17

,

.. e' (!) I . "0' Z 0.16 1 I ... I • W • 1" I ...J 0.15 Dill I , I

• ...J , 1

, O. l~ 1 D < I 1 . D

1 II: 0.13 I

1 .... I

I D en I

1 0.12 , I D 0 ..

I I II: 0.1 L

i I

1

1 D 0.10 1 .,

1 0.09

I

1 O.OB

.I

0.07

0.06

5 10 15 20 CS 30 35 40 45 50 55 60 65 70

0.24

0.23

O. :J2-

0.21

0.20

...... 0.19

U 0.18

:I: 0.17

0.16 (!)

Z 0.15

W

...J 0.14

...J

< 0.13

II: 0.12

en

0 O. t 1

II: 0.10

0.09

0.06

0.07

0.06

0.05

Figure 41.-The upper and lower beak rostral length versus the body weight of Loligo opa/esens and Onychoteuthis banksii. Symbols as in Figure 38.

HISTIOTEUTHIS DOFLEINI HISTIOTEUTHIS HETEROPSIS

0.6' 0.65

0.60 D 0.60

0.55 -- -_.. - 0.55

,-,-

-- 0.50 -- 0.50 ...... ......

U 0.45 0.45 U

:I: 0.4;") :I: 0.40

(!)

Z 0.35 (!) 0.35 ,

Z , w W JY . ...J ,' . 0.30 ...J 0.30 ,

...J ,- D

...J ,, II

< 0.25 < 0.25

II: II: ,

en 0.20 en 0.20

0 0 II: 0.15 II: 0.15

0.10 O. to

0.05 0.05

0.00 0.00

20 40 60 BO 1!)(1 120 140 1&0 IBO 200 220 240 0 20 40 60 BO 100 120 140 160 180 200 220 240

WEIGHT (G) WEIGHT (G)

Figure 42.-The upper and lower beak rostral length versus the body weight of HistioteUlhis dofleini and Histioteuthis heteropsis. Symbols as in Figure 38.

43
0.65

0.60

O.5S

0.50

,... 0.45

(.)

...... 0,40

:I:

ICJ 0.35

Z

W 0.30 ...J

...J

c( 0.25

a:

I- 0.20 (J)

0

a: o. 15

0.10

0.05

0,00

NOTOTODARUS HAWAIIENSIS

/ /

D ,0,./

,'"

/

D / /'.. ..

o /

/

/0

0'

:~ '. lI'

,'. /

, . ,

,

I

,

"

,

'.,

I

,

,

.,' ,

,

20 YO 60 80 100 120 IYO 150 160 200 220 ,"0

WEIGHT (G)

OMMASTREPHES BARTRAM II

U.65

0.50

0.55

0.50

, ,.- , (.) 0.l.I5 , /

,, ,

:I: 0.110 ~'

I- ~~ D

CJ 0 6' • Z 0.35 .' 0

W /

...J ./, • 0.30 , ...J A:I

c( ,• a: 0.25 ,

I- D /

./

(J) 0.20 ,

0 .,,

a: 0

0.15

0.10

0.05

0.00

20 YO 60 60 100 120 1YO 160 180 200 220 2YO

WEIGt-'T (G)

Figure 43.-The upper and lower beak rostral length versus the body weight of N%/odarus hawaiiensis and Ommas/rephes bartramii. Symbols as in Figure 38.

DOSIDICUS GIGAS SYMPlECTOTEUTHIS OUAlANIENSIS

0.'

0.6

a

0.7 a ,... a D/, a (.) 0.6 / . DD ,/ .. /

:I: a / /

I- a A/,:

CJ 0.5 ,/.. Z /

/

W ./

/

...J , O. Y ....

...J /

c( , D ,'.

a: .. I- 0.3 / , (J) ,

0 ,

,

a: ,

0.2

0.1

0.0

100 200 300 YOO 500 600 700 800 900 1000

WEIGHT (G)

Figure 44.-The upper and lower beak rostral length versus the body weight of Dosidicus gigas and Symplecloteulhis oualalliellsis. Symbols as in Figure 38.

44
ORNITHOTEUTHIS VOLATILIS SYMPLECTOTEUTHIS LUMINOSA

'''1 /

/ 0.55

, ,

D

0.60 0.60

D D 0.55 0.55

0.50 0.50 D /

,... , , , , ,... ::E ,~

::E O. Y5 () 0.45 , ,

() ...... /;'/S O. YO J: o. YO

+' J: I- , I- 0.35 C) 0.35 , C) Z ,'11

Z W I

I

W 0.30 ...J 0.30 ,I

...J D I

...J D ,

,

...J 0.25 « 0.25 ,.

I

« a: I

a: lI- 0.20 en 0.20

en 0 0 0.15 a: 0.15

a:

0.10 0.10

0.05 0.05

0.00 0.00

20 YO 60 80 100 J20 1YO 160 180 200 220 2lJO 20 YO 60 80 100 120 I~O 160 180 200 220 2YO

WEIGHT (G) WEIGHT (G)

Figure 4S.-The upper and lower beak rostral length versus the body weight of Ornithoteuthis volalilis and Symplectoteuthis luminosa. Symbols as in Figure 38.

TODARO DES PACIFICUS HY ALOTEUTHIS PELAGIC A 0.9 D 0.275 ,," ,," D " " " / " • 0.8 , , /

,- D , 0.250 , / /

/

, / , , D ,,

D , , 'i 0.225 '"

, ,......, O. 7 ,

, D /

::E , /

, () ,, () D , D , ...... / 0.200 /

/

/

0.6 , J: / D

J: /

l- f/ . I- , ~' C) :6 C) 0.175 / I>

I I> Z /1> D D

Z 0.5 I

W , , I

W , ...J / I O. !SC , ...J , , , ...J ...J I "'" , « ~/C4lP « O.Y ,

I a: 0.125 a: " D. I

l- I l- I , en I

en , I

0 0 .

3 0 I 0,100 , /. a: a: ,

,

I

0.2 0.075

0.050

O. I

0.025

0.0

100 200 300 YOO 500 600 700 800 900 1000 0.000

WEIGHT (G) 10 12 14 16 18 20 22 24

WEIGHT (G)

Figure 46.-The upper and lower beak rostral length versus the body weight of TodDrodes pacificus and Hyaloteuthis pelagica. Symbols as In Figure 38.

45
ABRALIOPSIS AFFINIS ABRALIOPSIS FELIS

"0 60 60 'DO '20 1"0

MANTLE LENGTH (MM)

20

0.325

0,075

O. f'lSO

O. ~50

J: 0.225

I-

"

Z 0.200

W

..J 0.175

..J

a:< 0.150

l-

(/) 0.125

o

a: 0.100

0.025

0.300

,.... 0.275

::E

00.250

.....

"0 60 60 100 120 1"0

/

.:'

..I

./

20

O.OOO-"....,~~,......~.......,C'""""'_.......,,~~..,...,~_.,.......~.......,~~....,...~~

o

0.075

0.275

0.350

0.050

0.300

MANTLE LENGTH (MM)

0.325

0.025

J: 0.225

I-

"

Z 0.200

W

..J 0.175

..J

<a: 0.150

l-

(/) 0.125

o

a: 0.100

::E

0 0 . 250

.....

Figure 47.-The upper and lower beak rostral length versus the mantle length of Abraliopsis affinis and Abraliopsis ft/is. Symbols as in Figure 38.

PTERYGIOTEUTHIS GIARDI GONATUS ONYX

0.3:.0

0.325

0.300

0.275

::E

00.250

.....

i= 0.225

"

Z 0.200

W

..J 0.175

..J

<a: 0.150

l-

(/) 0.125

o

a: 0.100

0.075

0.050

0.025

0.350j

O. 32Si

0.300

..... O. 275

::E

00.250

.....

J: 0.225-

I-

"

Z 0.200

W

..J 0.175

..J

<a: ).150

l-

(/) 0.125

o

a: 0.100

0.075

0.050

0.025

20 "0 60 60 100 120 1"0 20 "0 60 60 100 120 1"0

MANTLE LENGTH (MM) MANTLE LENGTH (MM)

Figure 48.-The upper and lower beak rostral length versus the mantle length of Purygioteuthis giardi and Gonalus onyx. Symbols as in Figure 38.

46
L10CRANCHIA REINHARDTI

40 60 eo lao 120 140

•

MANTLE LENGTH (MM)

20

0.050

0.075

0.325

0.275

0.300

o. ~SO 1

:I: 0.225

C' Z 0.200

W

...I 0.175

...I

«a: 0.150

en 0.125

o

a: 0.100

00.250

......

140

./~

1>,./

LEACHIA DANAE

40 60 eo 100 120

MANTLE LENGTH (MM)

0.350

0.325

0.300

0.275

00.250

......

:I: 0.225

C' Z 0.200

W

...1 0 . 175

...I

« 0.150

a:

en 0.125

0

a: 0.100

0.075

0.050

""'I 0.000 I

0 20

Figure 49.-The upper and lower beak rostral length versus the mantle length of Leachia danae and Liocranchia reinhardti. Symbols as in Figure 38.

LOLIGO OPALESCENS ONYCHOTEUTHIS BANKSII G.350 0.350

0.325 0.325

0.300 0.300

MANTLE LENGTH (MM)

40 eo 100 120 140

MANTLE LENGTH (MM)

0.050

0.025

0.075

",0.275

0 0 . 250 ......

:I: 0.225

C'

Z 0.200

W

...I 0.175

...I

«

a: 0 . 150

en0 0 .

125

a: 0.100

140

•

D

A

20 40 60 eo 100 120

0.275

0.025

:I: 0.<25

" 0.200

Z

W...I 0.175

...I

« 0.150

a:

t; 0.125

o

a: 0.100

0.050

0.075

,....,

o 0.250

Figure 50.-The upper and lower beak rostral length versus the mantle length of Loligo opalenscens and Onychoteuthis banksii. Symbols as In Figure 38.

47
HISTIOTEUTHIS DOFLEINI HISTIOTEUTHIS HETEROPSIS 0.9

c.•

0.7

::c

ICJ 0.5

Z

W

...J 0.4 ...J

<:(

0:

I- 0.3

00

o

0: 0.2

0.1

O.0"l,-;:.....,r-....,._..,.........,._....-_r-....,._..,.........,._....-_r-...,.........,-

25 50 75 100 125 150 175 200 225 250 275 300

MANTLE LENGTH (MM)

o. '501

0.325

0.300

0.275

::EU 0.250

....

::c 0.225

ICJ Z 0.200

W

...J 0.175

...J

<:( 0.1:0

0:

I00 0.125

o

0: 0 . 100

0,075

0.050

20 40 60 eo 100 120

MANTLE LENGTH (MM)

140

Figure 51.-The upper and lower beak rostral length versus the mantle length of Hislioleuthis dofleini and Hislioleulhis heleropsis. Symbols as in Figure 38.

0.9

O••

0.7

'"' :::E

U

...... 0.6

::c

ICJ Z 0.5

W

...J

...J 0.4

<:(

0:

I00 0.3

o

0:

0.2

O. I

NOTOTODARUS HAW AIIENSIS

25 50 75 100 125 150 175 200 225 250 275 ,00

MANTLE LENGTH (MM)

OMMASTREPHES BARTRAM II

0.8

0.7

::E

U

...... 0,6

::c

ICJ Z 0.5

W

...J

...J 0.4

<:(

0:

I00 0.3

o

0:

0.2

•,j

o.al,."~.,........?:,~ ......~.,.........,-....,..,-..,.........,r--.......- .......-r-.....,..~-r 25 50 75 100 125 150 175 200 225 250 275 300

MANTLE LENGTH (MM)

Figure 52.-The upper and lower beak rostral leogth versus the mantle length of Nololodarus hawaiiensis and Ommastnphes bartramii. Symbols as In Figure 38.

48
DOSIDICUS GIGAS 0.9

SYMPLECTOTEUTHIS OUALANIENSIS 0.9

25 5" 75 100 125 150 175 200 225 250 275 300

MANTLE LENGTH (MM)

0.8

0.7

::E

U

"'-" 0.6

J:

I-

(!) 0.5

Z

W

..J

..J O.Y

c(

a:

l-

(/) 0.3

o

a:

0.2

O. I

/'

" ..- .' 0.0 "

0.8

0.7 ,..

::E

U

...., 0.6

J:

I-

(!)

Z 0,5

W

..J

..J O.Y c(

a:

I-

(/)0,3

o

a:

0.2

O. I

c

c

c

c

c

25 50 75 100 1<5 150 175 200 225 250 275 300

MANTLE LENGTH (MM)

Figure 53.-The upper and lower beak rostral length versus the mantle length of Dosidicus gigas and Symplectoteuthis oualaniensis. Symbols as in Figure 38.

o.s

ORNITHOTEUTHIS VOLATILIS 0.9 '

SYMPLECTOTEUTHIS LUMINOSA

0,8 0.8

,,0,7

::E

U

,.;

0.6

J:

I-

(!)

ZO.5

W

..J

..J o. Y c(

a:

l-

(/) o. 3

o

a:

0.2

O. I

c

2S SO 75 100 125 150 175 200 225 250 275 300

MANTLE LENGTH (MM)

25 SO 7S 100 125 150 (75 200 225 250 275 300

MANTLE LENGTH (MM)

Fillure 54.-The upper and lower beak rostral length versus the mantle length of Ornithoteuthis volotitis and Symplectoteuthis luninosa. Symbols as in Figure 38.

49
TODARODES PACIFICUS HY ALOTEUTHIS PELAGICA 0.9

0.9

:I: 0.6

le!)

Z

W 0.5

...J

...J

« 0.4

a:

Ien

00.3

a:

0.2

O. (

/,,";

O. O.,......;.~/.,..........,~....,..~,....",.....,r='-..,..._~_,...,........"~..,,..~~....,,~,.,. 25 50 75 100 125 150 175 200 225 250 275 300

MANTLE LENGTH (MM)

O. }50

0.325

0.30C

..... 0.275

U 0.250

--:I: 0.22'5 le!)

Z 0.200

W

...J 0, ! 75

...J

«

a: 0.150

Ien O. (25

0

a: 0.100

0.075

0.050

0.025

0.000

20 40 60 eo 100 120 140

MANTLE LENGTH (MM)

Figure 55.-The upper and lower beak rostral length versus the mantle length of Todarodes pacificus and Hyaloteuthis pelagica. Symbols as in Figure 38.

50