A Complete Description of Sepia mira (Cotton 1932) 
(Cephalopoda: Sepiidae) from Eastern Australia 
Amanda Reid 
(Communicated by M.L. Augee) 
140 Napoleon Street, Eltham, Victoria 3095 
Reid, A. (1998). A complete description of Sepia mira (Cotton 1932) (Cephalopoda: 
Sepiidae) from Eastern Australia. Proceedings of the Linnean Society of New South 
Wales 119, 155-171. 
Sepia mira (Cotton 1932) is described on the basis of five specimens trawled between 
20-72 m off northern New South Wales. The species was known previously only from cuttle- 
bones collected from a number of eastern Australian localities. This complete description 
confirms the status of S. mira as valid. 
Manuscript received 20 July 1997, accepted for publication 17 September 1997. 
KEY WORDS: Sepia mira, Sepiidae, doratosepion, description, eastern Australia. 
INTRODUCTION 
Sepia mira was described by Cotton (1932) on the basis of a cuttlebone collected 
on Norfh-West Islet in the Capricorn Group of Islands off Gladstone, Queensland. Since 
that time, bones have been collected from additional localities in New South Wales 
(NSW) and Queensland, but whole animals had remained elusive. In 1995 and 1996, 
Ken Graham from NSW Fisheries collected some representatives of this species near the 
Clarence River mouth in northern New South Wales, enabling the species to be fully 
described here. The specimens were collected as part of a NSW Fisheries survey of the 
Newcastle and Clarence River prawn grounds. Other sepiids collected within the range 
of S. mira were: S. apama Gray 1849; S. limata (Iredale 1926) (see endnote); S. opipara 
(Iredale 1926); S. plangon Gray 1849 and S. rozella (Iredale 1926), with the latter two 
species occurring in the greatest abundance (Graham and Wood 1997). 
Cotton (1932) decided that Sepia mira was sufficiently distinctive to place in a new 
genus, Tenuisepia. Subsequent workers (with the notable exception of Iredale and 
McMichael (1962) who placed the species in its own subfamily) have generally conclud- 
ed that distinct generic status is not justified. Until a complete revision of the sepiids is 
undertaken, the classification followed here is that proposed in Khromov et al. (in press) 
which recognises only three genera within the Sepiidae with the present species designat- 
ed to the genus Sepia. 
This complete description confirms the status of S. mira as valid. 
MATERIALS AND METHODS 
This work was based on museum material. All material studied is listed in the 
Material Examined section. Institutional acronyms used are: AM — Australian Museum, 
Sydney, Australia; MV — Museum of Victoria, Melbourne, Australia and SAM — South 
Australian Museum, Adelaide, Australia. Other abbreviations: coll. — collected, F — 
female, FV — Fisheries Vessel, Is. — Island, J — juvenile, m — meters, M — male, mm 
— millimeters. 
Proc. Linn. Soc. n.s.w., 119. 1998 
156 complete description of sepia mira 
Table 1 
Description of measurements and counts. Definitions largely follow Roper and Voss (1983). New or modified 
definitions are indicated by an asterisk (*). Indices (shown in square brackets) are calculated by dividing each 
measure by mantle length or, for cuttlebone characters, cuttlebone length (unless otherwise specified). 
Arm Length — AL: length of each designated (i.e. 1,2 etc.) arm measured from first basal (proximal-most) 
sucker to distal tip of arm (Arm 1, dorsal; 2, dorso-lateral; 3, ventro-lateral; 4, ventral) [ALI]. 
Anterior Mantle to Head length * — AMH: dorsal length of mantle measured from anterior-most point of man- 
tle to intersection of transverse line joining dorso-lateral mantle margin [AMHI]. 
Arm Sucker Count * — ASC: total number of suckers on each designated arm (e.g. ASC2). 
Arm Sucker diameter — AS: diameter of largest sucker on each designated (i.e. 1,2 etc.) arm [ASIn]. Suckers 
on left ventral hectocotylised arms are differentiated as follows: 
Arm Sucker left 4 * — AS14: diameter of largest sucker on left ventral arm of male [ASInI4]. 
Cuttlebone Breadth — CbB: greatest dorso-ventral breadth of cuttlebone. [CbBI] 
Cuttlebone Length — CbL: dorsal length of cuttlebone along midline, including spine. 
Cuttlebone Width — CbW: greatest width of cuttlebone [CbWI]. 
Club Length — C1L: length of tentacular club measured from proximal-most basal suckers (carpus) to distal tip 
ofclub[ClLI]. 
Club Row Count — CIRC: number of suckers in transverse rows on tentacular club. 
Club Sucker diameter — CIS: diameter of largest sucker on tentacular club [C1SI]. 
Club Sucker dorsal * — CISd: diameter of largest tentacular club sucker in dorsal-most (closest to swimming 
keel) longitudinal row [CISId]. 
Club Sucker ventral * — CISv: diameter of largest tentacular club sucker in ventral-most (opposite swimming 
keel) longitudinal row [CISIv]. 
Eye Diameter — ED: diameter of eye [EDI]. 
Egg Length * — EgL: length of egg [EgLI]. 
Free Funnel length — FFu: the length of the funnel from the anterior funnel opening to the point of its dorsal 
attachment to the head [FFul]. 
Fin Insertion anterior * — Fla: anterior origin of fin measured from mantle margin to anterior-most junction of 
fin and mantle [Flla]. 
Fin Insertion posterior* — FIp: measured between posterior junctions of fins with mantle [Flip]. 
Funnel Length — FuL: the length of the funnel from the anterior funnel opening to the posterior margin mea- 
sured along the ventral midline [FuLI]. 
Fin Width — FW: greatest width of single fin [FWI]. 
Gill Lamellae Count — GiLC: number of lamellae on outer demibranch including the terminal lamella. 
Gill Length * — GiL: length of gill [GiLI]. 
Head Length — HL: dorsal length of head measured from point of fusion of dorsal arms to anterior tip of 
nuchal cartilage [HLI]. 
Head Width — HW: greatest width of head at level of eyes [HWI]. 
Loculus Length * — LoL: length of the last loculus (ventral anterior smooth zone of the cuttlebone) [LoLI]. 
Mantle Length — ML: dorsal mantle length. Measured from anterior-most point of mantle to posterior apex of 
mantle. 
Mantle Width — MW: greatest straight-line ventral width of mantle [MWI], 
Spine Length * — SL: length of spine [SLI]. 
Spermatophore Length — SpL: length of spermatophore [SpLI]. 
Spermatophore Width — SpW: greatest width of spermatophore Spermatophore width index is expressed as a 
percentage of spermatophore length [SpWI]. 
Striated Zone length — StZ: length of striated zone of cuttlebone [StZI). 
Transverse Row Count — TrRC: number of suckers in longitudinal series on tentacular club [counted from 
proximal-most basal suckers (carpus) to distal tip of club]. 
Ventral Mantle Length — VML: length of ventral mantle measured from anterior mantle margin at ventral 
midline, to posterior apex of mantle [VMLI]. 
Prcx:. Linn. Soc. n.s.w., 119. 1998 
A. REID 
157 
OGO 
DDD 
Figure 1. Measurements and Terminology: (a) whole animal dorsal view (for abbreviations and definitions see 
Table 1); (b) tentacular club (C — carpus, D — dorsal, PM — protective membranes, SK — swimming keel, V- 
ventral). The number of suckers intersected in an oblique transverse line across the club, shown as a hatched line 
on this figure, is the Club Row Count (CIRC) [In the example illustrated CIRC = 4]; (c) upper beak (C — crest, 
H — hood, R — rostrum); (d) lower beak (C — crest, H — hood, L — lateral wall, W — wing); (e) arm sucker 
rim (I — inner ring, INF — infundibulum, PO — polygonal process, P — peg); (f) radula (R — rhachidian 
teeth, LI — first lateral teeth, L2 — second lateral teeth, M — marginal teeth); (g) cuttlebone, ventral view (for 
abbreviations and definitions, see Table 1). [(a) and (g) modified from Roper and Voss (1983) Figure 1]. 
158 COMPLETE DESCRIPTION OF SEPIA MIRA 
Table 2 
Measurements (mm), counts and indices of 3 male and 2 female Sepia mira (Cotton). M — male, F — female. 
Museum 
Reg. No. 
MV 
F80995 
M 
MV 
F80996 
M 
AM 
C306764 
M 
MV 
F80997 
F 
AM 
C306387 
F 
ML 
38.1 
42.7 
44.0 
43.0 
43.9 
MWI 
43.8 
39.3 
43.2 
46.7 
45.6 
AMHI 
11.0 
12.6 
12.3 
9.3 
10.9 
VMLI 
85.3 
84.3 
83.6 
90.0 
80.6 
FWI 
6.3 
8.4 
8.4 
9.3 
7.5 
Flla 
6.3 
2.8 
5.7 
5.1 
4.6 
Flip 
4.7 
3.3 
4.5 
12.3 
9.6 
FuLI 
26.1 
29.5 
25.7 
29.1 
28.5 
FFul 
11.8 
10.5 
14.8 
9.3 
9.1 
HLI 
27.1 
23.0 
22.3 
22.6 
28.5 
HWI 
31.5 
32.3 
30.7 
34.7 
31.9 
EDI 
12.6 
10.8 
10.5 
10.5 
8.0 
ALII 
31.5 
31.6 
25.0 
34.9 
27.3 
ALI2 
30.2 
29.3 
36.4 
32.6 
31.9 
ALI3 
31.5 
25.8 
31.8 
31.4 
37.6 
ALI4 
39.4 
37.5 
38.6 
39.5 
36.4 
ASInl 
0.84 
1.03 
1.02 
1.12 
1.03 
ASIn2 
1.05 
1.03 
1.14 
1.05 
1.03 
ASIn3 
1.05 
1.17 
1.25 
1.12 
1.25 
ASIn4 
1.18 
0.94 
1.18 
1.28 
1.25 
ASCI 
80 
78 
90 
90 
82 
ASC2 
60 
82 
95 
92 
82 
ASC3 
60 
78 
110 
103 
98 
ASC4 
92 
122 
104 
116 
110 
ASInl4 
1.05 
0.70 
1.14 
- 
- 
CILI 
11.8 
11.2 
10.0 
10.0 
- 
CIRC 
3 
3 
3 
3 
- 
TrRC 
16 
13 
17 
20 
- 
C1SI 
1.34 
1.41 
1.36 
0.93 
- 
CISId 
0.63 
0.63 
0.80 
0.56 
- 
CISIv 
0.39 
0.35 
0.34 
0.37 
- 
GiLC 
- 
26 
- 
24 
- 
GiLI 
28.3 
28.3 
34.1 
30.0 
27.3 
SpLI 
7.9 
6.6 
6.6 
- 
- 
SpWI 
5.33 
5.71 
4.14 
- 
- 
EgDI 
- 
- 
- 
5.8 
7.3 
CbL 
39.9 
47.9 
44.0 
51.4 
45.1 
CbWI 
22.8 
22.1 
22.7 
23.7 
22.0 
CbBI 
7.0 
7.3 
7.9 
5.8 
7.3 
SLI 
3.9 
3.4 
2.7 
4.5 
3.3 
StZI 
69.6 
63.9 
66.6 
72.8 
67.2 
LoLI 
21.0 
28.8 
30.5 
20.2 
30.8 
LoL/StZ(%) 
30.2 
45.1 
45.7 
27.1 
45.9 
Proc. Linn. Srx\ n.s.w., 119. 1998 

A. 
REID 
159 
Table 3 
Sepia mira 
(Cotton); ranges of arm length indices (ALI), 
arm sucker diameter indices (ASIn) and arm 
sucker counts 
(ASC) of 3 mature males and 2 
mature females 
. min. : 
= minimum, max. = maximum, 
SI) 
= standard deviation. 
Males 
Females 
min. 
mean 
max. 
SD 
min. 
mean 
max. 
SD 
ALII 
25.0 
29.4 
31.6 
3.8 
27.3 
31.1 
34.9 
5.3 
ALI2 
29.3 
31.9 
36.4 
3.9 
31.9 
32.2 
32.6 
0.5 
ALB 
25.8 
29.7 
31.8 
3.4 
31.4 
34.5 
37.6 
4.4 
ALI4 
37.5 
38.5 
39.4 
1.0 
36.4 
38.0 
39.5 
2.2 
ASInl 
0.84 
0.96 
1.03 
0.11 
1.03 
1.07 
1.12 
0.06 
ASIn2 
1.03 
1.07 
1.14 
0.06 
1.03 
1.04 
1.05 
0.02 
ASIn3 
1.05 
1.16 
1.25 
0.10 
1.12 
1.18 
1.25 
0.10 
ASIn4 
0.94 
1.10 
1.18 
0.14 
1.25 
1.27 
1.28 
0.02 
ASCI 
78 
83 
90 
6 
82 
86 
90 
6 
ASC2 
60 
79 
95 
18 
82 
87 
92 
7 
ASC3 
60 
83 
110 
25 
98 
101 
103 
4 
ASC4 
92 
106 
122 
15 
110 
113 
116 
4 
Measurements and indices used throughout this paper are primarily those given in 
Roper and Voss (1983), using dorsal mantle length (ML) as a size standard. Some addi- 
tional measurements are used, and these with the definitions listed by Roper and Voss 
(1983) are given in Table 1. Parts of the club and arm sucker rims are described using the 
terminology of Nixon and Dilly (1977), while nomenclature for the radula follows Nixon 
(1995). Beaks were described following Clarke (1986). Diagrammatic illustrations of 
measurements and terminology used for key structures are shown in Fig. 1 . 
Measurements were made either using dial callipers, or an eyepiece micrometer 
attached to a stereo microscope. All measurements are expressed in millimetres (mm). 
Measurements and counts for individual specimens are shown in Table 2. Ranges of arm 
length indices, arm sucker diameter indices, and arm sucker counts are presented in 
Table 3; ranges for all other characters appear in the text. In species descriptions and 
tables, the range of values for each character is expressed as: minimum — mean — max- 
imum (standard deviation, SD). Values for each sex are given separately. 
Measurements for structures which were clearly distorted or broken were not 
attempted, and these, in addition to missing values, appear as a dash (-) in the tables. 
Ranges for specific character traits given with the species description do not, therefore, 
always refer to the total number of specimens examined. 
For examination of arm and club sucker rims, suckers were removed from the mid- 
dle of designated arms and the tentacular club, mounted in glycerine jelly and viewed 
using a compound microscope. Radulae and beaks were dissected from the buccal mass, 
and soaked for approximately 30 minutes in a warm, saturated potassium hydroxide solu- 
tion, then radulae were cleaned using forceps and a fine brush. Radulae were also mount- 
ed in glycerine jelly, and the new, unused portion was examined. 
The species description was generated by the DELTA (Description Language for 
Taxonomy) system (Dallwitz 1980; Dallwitz et al. 1993; Partridge et al. 1993). 
Proc. Linn. Soc. n.s.w., 119. 1998 
1 60 COMPLETE DESCRIPTION OF SEPIA MIRA 
Sepia mira (Cotton 1932) 
(Figures 2-9, Tables 2 & 3) 
Synonymy 
Tenuisepia mira Cotton 1932: 546-547, figs 7-9.-Iredale 1954: 75, pi. V, figs 7 & 
8.-Iredale & McMichael 1962: 99. 
Sepia mira.- Adam & Rees 1966: 87.-Nesis 1982: 119, fig. 24X.-Lu & Phillips 
1985: 25. 
Material examined 
Holotxpe: Australia: Queensland, North- West Is. — cuttlebone (55 mm CbL), 
23°18'S, 151°42'E, coll. W.J. Kimber, (SAM D10507). Additional Material: Australia: 
Queensland — 5 cuttlebones (41.5-42.5 CbL), Lizard Is., Coconut Beach, 14°40'S 
145°28'E, 10 Oct 1982, coll. R. Burn, (MV F80999); cuttlebone (37.3 CbL), Heron Is., 
23°26'S 151°55'E, 1988, coll. A. Reid & P. Ward, (MV F80998). New South Wales — 
IF (43.0 mm ML), Clarence R., 29°19'S 153°29'E — 29°20'S 153°29'E, 49^18 m, 19 
Apr 1996, coll. K. Graham on FV "Kapala" K960638, (MV F80997); 1M (44.0 mm 
ML), Off Iluka, 29°20'S 153°25'E — 29°19'S 153°25'E, 37-34 m, 29 May 1995, coll. 
K.J. Graham on FV "Kapala" K950620, (AM C306764); 1M (42.7 mm ML), Clarence 
R., 29°22'S 153°23'E — 29°23'S 153°23'E, 28-20 m, 31 Aug 1995, coll. K. Graham on 
FV "Kapala" K951131, (MV F80996); 1M (38.1 mm ML), Clarence R., 29°50'S 
153°20'E — 29°48'S 153°20'E, 37-33 m, 10 Apr 1996, coll. K. Graham on FV 
"Kapala", K960602 (MV F80995); IF (43.9 mm ML), off Wooli, 29°49'S 153°27'E — 
29°48'S 153°27'E, 72-69 m, 18 May 1995, coll. K. Graham on FV "Kapala", K950604 
(AM C306387); cuttlebone (48 mm CbL), Trial Bay, 30°53'S 153°04'E, coll. M. Ward, 
(AMC133312). 
Diagnosis 
Cuttlebone oblong; spine with ventral keel; striated zone convex; anterior striae 
slightly convex to straight; sulcus very shallow, narrow; last loculus very short; inner cone 
limbs straplike anteriorly, narrower, thicker posteriorly; outer cone posterior lateral limbs 
flared ventro-laterally. Arm suckers tetraserial. Tentacular club suckers differ in size, ?>-A 
enlarged suckers toward posterior end of club; 3 suckers in transverse rows; dorsal and 
ventral protective membranes not fused at base of club; swimming keel extends beyond 
carpus. [Modified from the original description by Cotton (1932, p. 546)]. 
Description 
Counts and indices for individual specimens are given in Table 2; ranges for arm 
length indices, arm sucker diameter indices and arm sucker counts are shown in Table 3. 
Small to moderate-sized species (Fig. 2); ML males 38.1- 41.6 -44.0 (SD, 3.1), 
females 43.0-43.5-43.9 (SD, 0.6). Mantle oblong; MWI males 39.3-42.1-43.8 (SD, 
2.4). females 45.6-4(x2-46.7 (SD, 0.8); dorsal anterior margin triangular, acute; extend- 
ing to level of posterior margin of eyes; AMHI males 11.0- 12.0 -12.6 (SD, 0.8), females 
9.3- 10. 1 -10.9 (SD, 1.2). Ventral mantle margin weakly convex; VMLI males 
83.6-84.4-85.3 (SD,0.8), females 80.6-85.3-90.0 (SD, 6.6). Fins widest in posterior 
third; FWI males 6.3-7J-8.4 (SD,1.2), females 7.5-84-9.3 (SD,1.3); anterior origin 
posterior to mantle margin; Flla males 2.8^.9-6.3 (SD, 1.9), females 4.6-4.9-5.1 (SD, 
0.3); rounded posteriorly; narrow gap between fins; Flip males 3.3—4^2-^4.7 (SD, 0.8), 
females 9.6- 10.9 -12.3 (SD, 2.0). Funnel long, broad-based; extends to level of anterior 
Proc. Linn. Soc. n.s.w.. 119. 1998 
A. REID 
161 
imiiHii!lll|U : 
CIV! 
Figure 2. Sepia mira: left, cuttlebone dorsal view (posterior end at top); right, freshly caught animal dorsal 
view, cuttlebone removed, male MV F80996, 42.7 mm ML [photo K. Graham]. 
rim of eye; FuLI males 25.7-27.1-29.5 (SD, 2.1), females 28.5-28.8-29.1 (SD, 0.4). 
Funnel free portion approximately one-third funnel length; FFul males 10.5- 12.4 -14.8 
(SD, 2.2), females 9.1-9.2-9.3 (SD, 0.1). Funnel organ dorsal elements inverted V-shape 
with small anterior papilla; ventral elements oval (Fig. 3a). Mantle-locking cartilage 
curved, with semicircular ridge; funnel-locking cartilage with depression which corre- 
sponds to ridge (Fig. 3b). Head short (Fig. 2); HLI males 22.3-24.1-27.1 (SD, 2.6), 
females 22.6- 25.5 -28.5 (SD, 4.2); slender, narrower than mantle; HWI males 
30.7-31.5-32.3 (SD, 0.8), females 31.9-33.3-34.7 (SD, 2.0). Eyes moderate size; EDI 
males 10.5-11.3-12.6 (SD, 1.2), females 8.0-9^-10.5 (SD, 1.8); ventral eyelids present 
(anterior half of eye only). 
Male arms 4 slightly longer than other arms, arms 1-3 all similar length, not elon- 
gate (Table 3). Female arm lengths subequal (Table 3). ALI of longest arms in males 
(ALI4) 37.5-38.1-38.6 (SD, 0.8), females (ALI4) 36.4-38.0-39.5 (SD, 2.2). Protective 
membranes (both sexes) narrow; normal, not thickened. Distal arm tips (both sexes) not 
markedly attenuate. Arm suckers tetraserial in both sexes. Suckers in males normal in 
size (not greatly enlarged); similar to female arm suckers in size (Table 3). Chitinous 
Proc. Linn. Soc. n.s.w., 119. 1998 
162 
COMPLETE DESCRIPTION OF SEPIA MIRA 
Figure 3. Sepia mira: (a) funnel organ, male, MV F80996, 42.7 mm ML, scale bar 2 mm; (b) funnel locking 
cartilage (left), and mantle locking cartilage (right), female, MV F80997, 43 mm ML, scale bar 2 mm; (c) suck- 
er rim arm 2, scale bar 50 fim, arrow indicates elongate teeth on distal margin of inner ring; (d) enlargement of 
sucker rim arm 2, scale bar 50 /jm; (e) hectocotylus, left arm 4, scale bar 2 mm; (f) right arm 4, scale bar 2 mm. 
(a-b & e-f, male MV F80995, 38. 1 mm ML). 
rims of arm suckers with elongate rectangular teeth on distal half of inner ring, teeth 
much smaller on proximal half of ring; infundibulum with 4-5 rows of hexagonal 
processes, inner 2-3 rows with elongate rounded pegs on distal half, becoming smaller 
towards periphery of sucker, shorter pegs on proximal half as for inner ring; peripheral 
sucker rim processes radially arranged, elongate, without pegs (Figs 3c and 3d). Sucker 
counts range from 60-122; females with higher average counts than males (Table 3). 
Hectocotylus present, left ventral arm modified (see remarks); sucker size slightly 
reduced proximally; suckers equal in size across rows, maximum sucker diameters: 
ASInl4 0.70-0.96-1.14 (SD, 0.23) (compare with right ventral arm, Table 3). Oral sur- 
face of modified region wide, fleshy, with transversely grooved ridges (Fig. 3e, compare 
with Fig. 3f); without distinct median furrow. 
Proc. Linn. Soc n.s.w., 119. 1998 
A. REID 
163 
Figure 4. Sepia mira: (a) tentacular club, oral view; (b) tentacular club, oral view, dorsal protective membrane 
pinned back to show sucker arrangement; (c) tentacular club, aboral view, (a-c, female, MV F80997, 43.0 mm 
ML, scale bars 2 mm). 
Tentacular club similar length in males and females; C1LI males 10.0- 11.0 -11.8 
(SD, 0.9), female 10. Club crescent-shaped; sucker-bearing face flattened (in all speci- 
mens examined, posterior half to two thirds of the club is folded toward dorsal side, ven- 
tral protective membrane covering posterior suckers, Fig. 4a). Club with 3 suckers in 
transverse rows in both sexes; 13-20 suckers in longitudinal series, TrRC males 
13-15-17 (SD, 2), female 20. Suckers differ markedly in size, 3-4 enlarged suckers 
towards posterior end of club (Fig. 4b); C1SI males 1.34-1.37-1.41 (SD, 0.03), female 
0.93; dorsal and ventral marginal longitudinal series of suckers differ slightly in size; 
dorsal marginal longitudinal series of suckers slightly larger than those in ventral margin- 
al series; CISId males 0.63-0.69-0.80 (SD, 0.09), female 0.56; CISIv males 
0.34- 0.36 -0.39 (SD, 0.03), female 0.37. Sucker dentition: half inner ring circumference 
in both sexes with elongate rectangular teeth, remaining half with blunt projections; 
infundibulum with 3^4- rows of hexagonal processes, innermost with elongate rounded 
pegs, pegs smaller towards periphery of sucker; at periphery, processes smaller, elongate- 
rectangular, without pegs (Figs 5a and 5b) (similar to arm suckers). Swimming keel of 
club extends well beyond carpus. Dorsal and ventral protective membranes not fused at 
base of club; joined to stalk; dorsal and ventral membranes same length, terminate at 
posterior end of carpus (Figs 4a and 4b); approximately equal width (Fig. 4c); dorsal 
membrane forms shallow cleft at junction with stalk. 
Gills with 24-26 lamellae per demibranch; GiLC in only intact male 26, 24 in only 
intact female. Gill length: GiLI males 28.3-30.3-34.1 (SD, 3.3), females 27.3-28.7-30.0 
(SD, 1.9). 
Buccal membrane without suckers. Upper beak (Fig. 5c) rostrum sharply pointed, 
long, length greater than width, cutting edge slightly curved; hood high above crest poste- 
riorly; crest curved, lateral wall shallowly indented posteriorly; wings narrow and short; 
jaw angle approximately 90 degrees, slightly acute; hood dark brown, lighter on ventral 
Proc. Linn. Soc. n.s.w., 119. 1998 
164 
COMPLETE DESCRIPTION OF SEPIA MIRA 
DG 
Figure 5. Sepia mira: (a) club sucker rim, large club sucker, scale bar 100 /ym; (b) enlargements of large club 
sucker rim, scale bar 50 fjm; (c) upper beak, side view; (d) lower beak, side view; (e) lower beak, ventral view; 
(f) radula, male, MV F80996, 42.7 mm ML, scale bar 200 fim; (g) digestive tract, dorsal view, scale bar 500 
/urn (A — anus; BM — buccal mass; C — caecum; DG — digestive gland; IS — ink sac; P — pancreas; S — 
stomach; SG — salivary gland), (a-b & g, male MV F80995, 38.1 mm ML; c-e, female, MV F80997, 43.0 mm 
ML. scale bars 3 mm). 
margin, crest only slightly pigmented. Lower beak (Fig. 5d) rostrum protrudes only slight- 
ly, cutting edge curved; hood low on crest; crest straight, no indentation on lateral wall 
edge; hood and wings, width narrow; hood notch deep, broad (Fig. 5e); wings widely 
spaced; crest wide; rostrum pigmented dark brown, wings and crest only slightly pigment- 
ed. Radula homodont; rhachidian teeth with narrow, truncate bases, slender, triangular, 
sides slightly concave (Fig. 5f); first lateral teeth similar length and width to rhachidian 
teeth, asymmetrical with mesocone slightly displaced toward centre of radula; second lat- 
erals longer than first, distinctly curved on lateral margin, with broad heels; marginal teeth 
elongate with long tapered and curved mesocone (Fig. 4f). Digestive tract: (Fig. 5g) 
paired salivary glands approximately one-third length of buccal mass; paired digestive 
Prcx:. Linn. Soc. n.s.w., 119. 1998 
A. REID 
165 
Figure 6. Sepia mira: (a) male reproductive tract (testis not shown; spermatophore storage sac containing sper- 
matophores), scale bar 2 mm (AAG — appendix of accessory gland; AG — accessory gland; CC — ciliated canal: 
DDC — distal deferent canal; GO — genital orifice; MG — mucilaginous gland; SG — spermatophoric gland; SSS — 
spermatophore storage sac; VD — vas deferens); (b) spermatophore, scale bar 300 ^m; (c) spermatophore, oral end. 
scale bar 100 /jm; (d) female reproductive tract MV F80997, 43.0 mm ML, scale bar 3 mm (ANG — accessory nida- 
mental gland; GO — genital opening; NG — nidamental gland; O — ovary), (a-c male MV F80995, 38.1 mm ML). 
Proc. Linn. Soc. n.s.w., 119. 1998 
166 
COMPLETE DESCRIPTION OF SEPIA MIRA 
Figure 7 Sepia mira: fa) cuttlebone, dorsal view, holotype D 1 0507, 55.0 mm CbL, scale bar 1 cm; (b) cuttle- 
bone, ventral view, same specimen, scale bar 1 cm | photo C. Rowley]; (c) posterior end of cuttlebone, ventral 
view, AM C306764, 44.0 mm CbL, scale bar 2mm. 
Proc. Linn. Soc. n.s.w., 119. 1998 
A. REID 167 
glands large, located close together, with narrow, elongate triangular lobes posteriorly, 
ducts (not shown in figure) connect digestive glands near midline with caecum, ducts with 
branched attached pancreatic tissue; oesophagus runs dorsally along median junction of 
digestive glands, joins sac-like stomach immediately posterior to digestive glands; caecum 
disc-like, grooved in a blunt V-shape anteriorly, surface lining finely pleated; intestine 
undifferentiated; ink sac very large, elongate; anal flaps well developed. 
Male reproductive tract: testis on left posterior side of viscero-pericardial coelom; 
at distal end, convoluted vas deferens opens into broad, cone-shaped mucilaginous gland, 
then narrower, curved, spermatophoric gland (Fig. 6a). Close to junction with lobe- 
shaped accessory gland and gland appendix, delicate ciliated canal joins spermatophoric 
gland; distal deferent canal connects appendix of accessory gland to spermatophore stor- 
age sac; genital orifice opens dorsal to left gill in anterior end of mantle cavity. 
Spermatophores: cement body bipartite (Figs 6b and 6c); aboral end rounded, bulbous, 
connects to sperm reservoir via narrow duct; oral end of cement body cone-shaped, 
approximately twice length of, and slightly narrower than aboral end, connects to aboral 
end via long neck, tapers toward oral extremity of cement body; middle tunic com- 
mences along aboral part of cement body; ejaculatory apparatus coiled, extends into oral 
dilation of spermatophore. Spermatophores 2.8-3.0 mm long, 0.12-0.16 mm wide; SpLI 
6.6-LQ-7.9 (SD, 0.8); SpWI 4.14-5.06-5.71 (SD, 0.82). Smallest male with well devel- 
oped spermatophores in Needham's sac is 38.1 mm ML. 
Female reproductive tract: ovary hangs medially from dorsal wall of posterior vis- 
cera pericardial coelom. Oviduct thin-walled, continuous with body cavity; distally with 
thickened, glandular walls (oviducal glands). Nidamental glands in mature animals occu- 
py large portion of ventral side of mantle cavity. Accessory nidamental glands anterior to 
nidamental glands (Fig. 6d). Eggs oval; 2.5-3.2 mm long; EgLI 5.8-6.6-7.3 (SD, 1.0). 
Smallest female with eggs in ovaries is 43.0 mm ML. 
Subdermal cartilaginous layer between cuttlebone and skin absent. Cuttlebone 
length approximately equal to mantle length; outline oblong (Figs 2, 7 and 8); CbL males 
39.9-43.9-47.9 (SD, 4.0), females 45.1-48.3-51.4 (SD, 4.5); CbWI males 
22.1-22.6-22.8 (SD, 0.4), females 22.0-22.8-23.7 (SD, 1.3); not strongly convex in lat- 
eral view; CbBI males 7.0-7J-7.3 (SD, 0.2), females 5.8-7^-7.0 (SD, 1.1). Bone blunt- 
ly rounded anteriorly; bluntly rounded posteriorly; not strongly recurved ventrally. 
Dorsal surface pinkish laterally (see remarks); evenly convex; granulose, calcified dor- 
sally, particularly posteriorly (not as pronounced on flared outer cone). Dorsal median rib 
absent; lateral ribs absent. Chitin borders lateral and anterior margins of cuttlebone. 
Spine present, long; SLI males 2.7-33-3.6 (SD, 0.5), females 3.3-3j6-3.9 (SD, 0.4); 
straight, directed dorsally; with ventral keel (Fig. 8b); fine, radiating ribs between outer 
cone and spine (Fig. 7c). Dorso-posterior end of cuttlebone without median longitudinal 
ridge anterior to spine. Striated zone convex (with narrow, flat margins, Fig. 8b); StZI 
males 63.9-66.7-69.6 (SD, 2.9), females 67.2-70.0-72.8 (SD, 3.9). Last loculus convex 
(strongly convex immediately anterior to striated zone, becoming flat posteriorly); LoLI 
males 21.0-26.8-30.5 (SD, 5.0), females 20.2-25.5-30.8 (SD, 7.5); at midline half 
length of striated zone (approximately), LoL/StZ(%) males 30.2-40.3-45.7 (SD, 8.8), 
females 27.1- 36.5 -45.9 (SD, 13.3); loculus extends posteriorly as narrow margin on 
each side of striated zone. Sulcus extends entire length of cuttlebone; shallow, narrow 
(indistinct, visible only in large specimens); not flanked by rounded ribs. Last loculus 
with shallow median indentation, not very pronounced. Anterior striae broad inverted U- 
shape to straight (Fig. 8a). Limbs of inner cone extend anteriorly to approximately 
halfway along striated zone. Inner cone lateral limbs overlie calcareous striated zone 
anteriorly, extreme anterior tips bordered, separated from outer cone by striated zone. 
Inner cone limbs strap-like anteriorly, narrower posteriorly; raised, separated from striat- 
ed zone to form rounded ledge posteriorly; thickened (Fig. 7c). Inner cone without irreg- 
ular calcareous ribs posteriorly. Outer cone calcified; narrow anteriorly, broadens posteri- 
Proc. Linn. Soc. n.s.w., 119. 1998 
COMPLETE DESCRIPTION OF SEPIA MIRA 
U; f 
&* ^v 
[jjnr" »^*"> >f '«^h t . 
II 
^£ 
Figure 8. Sepia mira: (a) cuttlebone, ventral view, MV F80998, 37.3 mm CbL, scale bar 5 mm; (b) posterior 
end of cuttlebone, ventral view, same specimen, scale bar 3 mm. 
orly; lateral limbs flared ventro-laterally; limbs continue as narrow ledge ventral to spine 
(swollen slightly to cover posterior end of inner cone). 
Body papillae present (visible on a single specimen only); dorsal mantle with scat- 
tered short longitudinal ridges concentrated mid-dorsally and in a single row close to fins; 
ventral mantle without ridges. Head and arm papillae absent. Colour: head, arms and dor- 
sal mantle reddish brown (some dull orange spots below chromatophores); paired dorsal 
eye spots absent. Fins pale; without markings at base. Body without ventral pigment. 
Distinctive pigment spots on ventral side of club, spots in radiating rows extending from 
central region towards ventral margin (Fig. 4c). Mantle ridges orange-pink in colour. 
Type locality 
Queensland. North-West Islet, Capricorn Group, 23°18'S 151°42'E. 
Linn. Soc. n.s.w.. H9. 1998 
A. REID 169 
Type 
Holotype by original designation. SAM: D10507. Cuttlebone only, 55 mm CbL. 
Distribution 
Queensland, Lizard Is., 14°40'S 145°28'E — Trial Bay, 30°53'S 153°04'E (cuttle- 
bones); New South Wales, Clarence R., 29°19'S 153°29'E — off Wooli, 29°49'S 
153°27'E (entire animals) (Fig. 9); depth range 72-20 m. 
Remarks 
Contrary to Cotton's (1932) original description, a very faint sulcus was seen in 
large specimens examined in this study. As Cotton (1932) described, the sulcus cannot be 
seen on the type specimen (Fig. 7b), possibly due to wear on the ventral surface of this 
beach-collected cuttlebone. In other respects, the cuttlebones examined here do not differ 
from the original description. Adam and Rees (1966) report the absence of a keel on the 
spine, but a ventral keel is clearly present in this species. The dorso-lateral sides of the 
cuttlebone are pinkish in freshly preserved animals, but this colour is lost in specimens 
that have been stored in ethanol. 
Of the three male specimens collected, the spermatophores were well developed in 
one specimen only (MV F80995). The modification of the left ventral arm could be seen 
only in this specimen. The arms of this animal are not in very good condition, so the 
details of the modification of the hectocotylus needs to be confirmed when more material 
is available. In particular, as a number of suckers are missing from this specimen, it is 
difficult to determine the relative sizes of the suckers in the modified region. 
It is not possible to ascertain whether the folding of the posterior half of the tentac- 
ular club, seen in all specimens, is an artefact of preservation, or if this is characteristic 
for the species. It has not been observed in other species of sepiids examined by the 
author. 
The five animals upon which the above description is based were collected from a 
narrow geographic range off the northeastern coast of New South Wales (Fig. 9). 
Cuttlebones have been collected at considerable distances north of this region, suggest- 
ing that the distribution of this species may be more extensive than that indicated from 
the complete specimens, though the bones may have drifted some distance to these loca- 
tions. The small size of the species possibly accounts for the absence of material from 
museum collections until now. It may be escaping trawl nets, or be of insufficient interest 
to most collectors to be retained. 
Determination of the relationships of Sepia mira to other species is at present diffi- 
cult owing to the general lack of knowledge about the phylogeny of the sepiids. 
Khromov et al. (in press) have suggested that S. mira belongs to the 'doratosepion' 
species complex. This name is based on Rochebrune's (1884) generic name 
Doratosepion, though the validity of this genus has generally been rejected. The 
' doratosepion ' species complex (which may or may not be monophyletic) includes those 
species with an elongated body, short arms with biserial suckers, short tentacular clubs 
with unequal suckers, and a narrow elongate cuttlebone with two posterior 'wings' and a 
spine (Roeleveld 1972). In addition to the presence of biserial suckers on some or all of 
the arms, doratosepia often have modified arms in one or both sexes, some pairs may be 
elongate, tapering to a very narrow threadlike tip, or differ in sucker arrangement to the 
remaining arms. Most members of the group show sexual dimorphism. While sharing 
with members of this complex an elongated body, narrow bone, U-shaped inner cone, 
and unequal club suckers, S. mira does not have any distinctive modification of the arms 
(though they are short) or arm suckers, and little sexual dimorphism is evident. The outer 
Proc. Linn. Soc. n.s.w., 119. 1998 
170 
COMPLETE DESCRIPTION OF SEPIA MIRA 
Figure 9. Distribution of Sepia mira. Triangles indicate collection localities for cuttlebones only, circles indi- 
cate whole animals. The open triangle shows the type locality. 
cone in S. mira has been described as consisting of two long 'wings' (Adam and Rees 
1966), though, as these authors acknowledge, they do not form a recurved 'cup-like' 
expansion as is often seen in other narrow-boned species. The outer cone 'wings' are 
usually shorter in these species than in S. mira, though the outer cone 'wings' of S. elon- 
gata d'Orbigny 1845 from the Red Sea (figured in Adam and Rees (1966), PI. 21, Fig. 
132) are not dissimilar in appearance to those of S. mira in ventral view. Many species 
with broad cuttlebones, and therefore not included within the 'doratosepion' species 
complex, also have a ventro-laterally flared posterior outer cone. 
According to Cotton (1932), ,S'. mira shows some affinity to a species described by 
Iredale (1926) as Decorisepia, type species S. rex. Though not stated in the original 
description, the reason for this may be that S. mira and S. rex both have a very narrow 
inner cone and, were thought to lack a sulcus (though a sulcus is present in both species). 
Proc. Linn. Sex:, n.s.w., 119. 1998 
A. REID 171 
Sepia rex differs greatly from S. mira in many other respects; the bone is much wider, 
has dorsal ribs and differs in shape, the club suckers are all small and arranged in 10-12 
transverse rows, and the animal is much larger, ranging up to approximately 10 cm ML. 
It is unlikely that the two species share close affinities. 
Until we develop some working hypotheses of the phylogenetic relationships 
among the sepiids, the position of S. mira and particularly its association with other nar- 
row-boned species remains equivocal. 
ACKNOWLEDGEMENTS 
I wish to thank Ken Graham from NSW Fisheries for his interest in collecting and retaining the Sepia 
mira specimens described here, and Ian Loch (AM) for the loan of some of this material. I thank Ken Graham 
also for providing the photograph of the freshly collected animal and C.C. Lu for the photographs of the cuttle- 
bone of the holotype. 
REFERENCES 
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Murray Expedition 1933-1934 11(1), 1-165, 46 plates. 
Cotton, B.C. (1932). Notes on Australian Mollusca with descriptions of new genera and species. Records of the 
South. Australian Museum 4(4), 537-547. 
Clarke, M.R. (1986). 'Handbook for the Identification of Cephalopod Beaks'. (Oxford Science Publications: 
New York). 
Dallwitz, M.J. (1980). A general system for coding taxonomic descriptions. Taxon 29, 4 1^46. 
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Kapala Cruise Report No. 116. (NSW Fisheries: Australia). 
Iredale, T. (1926). The cuttle-fish "bones" of the Sydney beaches. Australian Zoologist. 4: 186-196. 
Iredale, T. (1954). Cuttle-fish "bones" again. Australian Zoologist. 12(1): 63-82. 
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of the Australian Museum. 11, 1-109. 
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Contributions to Zoology. 
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Occasional Papers. Museum of Victoria 2: 21-36. 
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pp. (Levitov, B.S. trans, from Russian, Burgess, L.A. ed. 1987). 
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of Zoology (London) 236, 73-81. 
Nixon, M. and Dilly, P.N. (1977). Sucker surfaces and prey capture. In: 'The Biology of Cephalopods'. (Eds M. 
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(CSIRO Division of Entomology: Canberra). 
Rochebrune, A.T de (1884). Etude Monographique de la Famille des Sepiadae. Bulletin des Sciences, par la 
Societe philomathique. Paris 7(8), 7-44, plates 3-6. 
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ENDNOTE 
Sepia limata was misidentified as S. braggi Verco 1907 in Graham and Wood (1997), based on the work 
of Lu (in press) that includes S. limata as a synonym of S. braggi. Work in progress and in preparation for pub- 
lication by A. Reid has shown 5. limata to be valid and distinct from S. braggi. 
Proc. Linn. Soc. n.s.w., 119. 1998