DISCOVERY OF THE FOURTH SPECIES OF THE ENIGMATIC
CHIROTEUTHID SQUID ASPEROTEUTHIS (CEPHALOPODA:
OEGOPSIDA) AND EXTENSION OF THE RANGE OF THE GENUS
TO THE SOUTH ATLANTIC
ALEXANDER I. ARKHIPKIN AND VLADIMIR V. LAPTIKHOVSKY
Fisheries Department, Falkland Islands Government, PO Box 598, Stanley, FIQQ 1ZZ, Falkland Islands
(Received 2 November 2007; accepted 5 February 2008)
ABSTRACT
A new species of deepwater squid Asperoteuthis nesisi is described based on a single specimen from the
Southwest Atlantic. The new species is characterized by rather thin but muscled mantle and arms,
and a wide ovoid funnel-locking apparatus with weak tragus and well-developed antitragus. Peculiar
integumental tubercles are present both on the mantle and head. The largest arm suckers have chitinous rings with 12 –14 separate sharp triangular teeth distally and a flat thin ridge proximally. Based
on the external morphology, A. nesisi n. sp. is more closely related to A. acanthoderma than to A. mangoldae. Two specimens of ‘Mastigoteuthis? A’ reported previously from waters of South Georgia probably belong to A. nesisi n. sp. Our finding extends the range of the genus Asperoteuthis from the
tropical and temperate Pacific to the sub-Antarctic Atlantic, supporting a cosmopolitan distribution
of this genus of deepwater squid.
INTRODUCTION
Until now, the genus Asperoteuthis (Chiroteuthidae) consisted of
three species of meso-bathypelagic squid. Two have been
found in the tropical northern Pacific (Young, Vecchione &
Roper, 2007) and one in temperate waters of New Zealand
(Salcedo-Vargas, 1999). The first species was described as
Chiroteuthis acanthoderma by Lu (1977) from the tropical western
Pacific. Nesis (1980) transferred this species to the new genus
Asperoteuthis. However, he considered C. acanthoderma Lu, 1977
to be a junior synonym of Asperoteuthis famelica (Berry, 1909),
described from a very damaged juvenile specimen caught near
the Hawaiian Islands. Later studies showed that A. famelica is
in fact Mastigoteuthis famelica (Mastigoteuthidae) (Young,
1978), so that the valid name for Lu’s specimens is Asperoteuthis
acanthoderma (see Young et al., 2007). Another species of the
genus, Asperoteuthis lui, was described from a very damaged
specimen taken from the stomach of ling (Pisces) using mainly
the tentacle morphology (Salcedo-Vargas, 1999). Recently, the
third species of the genus, A. mangoldae, was described from
waters of the Hawaiian Islands (Young et al., 2007).
We describe a fourth species of Asperoteuthis, caught in
sub-Antarctic waters of the Southern Atlantic. All previous
authors (Lu, 1977; Nesis, 1980; Salcedo-Vargas, 1999; Young
et al., 2007) note that squid of the genus Asperoteuthis have
many peculiar features (skin trabecules, funnel cartilage structure, armament of arm suckers, tentacle club structure with
wide protective membranes in proximal part) that are unique
among species in the family Chiroteuthidae.
SYSTEMATIC DESCRIPTION
Genus Asperoteuthis Nesis, 1980
Type species: Asperoteuthis acanthoderma (Lu, 1977)
Diagnosis: Tentacular club with suckers on distal half and wide
protective membrane on proximal half. Arms long, attenuated.
Correspondence: A. Arkhipkin; email: aarkhipkin@fisheries.gov.fk
Arms IV slightly longer in large subadults. Funnel valve
present. Funnel-locking apparatus varies from curved groove
without antitragus to inverted Y-shaped groove with weak and
long tragus and strong antitragus. Photophores absent from
viscera and arms IV. Long oval photophore on ventral surface
of each eye. Single, large aboral photophore on tip of each
tentacle.
Asperoteuthis nesisi new species
(Figs 1, 2)
Type material: Holotype Natural History Museum, London,
BMNH 20070615. Maturing female, 363 mm ML, with short
stumps of both tentacles; 53844’S 58846’W, collected by
Falkland Islands R/V Dorada pelagic trawl fishing near-bottom,
913 m depth off northern part of Burdwood Bank, Southwest
Atlantic, 20 July 2005 (Stn 2132, cruise ZDLH1-07-2005).
Etymology: Named after Dr Kir N. Nesis (1934 –2003) who
described the genus Asperoteuthis and whose outstanding scientific contribution benefited almost every field of cephalopod
biology.
Material examined: A single, damaged specimen collected from
the trawl wings. DNA was extracted from frozen tissue samples
and molecular data were collected from three mitochondrial
genes. Sequence data are available on Genbank via the following accession numbers for COI – EU421718, 16S rRNA –
EU421719 and 12S rRNA – EU421720.
Diagnosis: Rather thin but muscled mantle and arms; funnellocking apparatus wide, ovoid, weak tragus, well-developed
antitragus. Integumental tubercles on mantle and head.
Largest arm suckers with 12– 14 separate sharp triangular
teeth on distal inner ring.
Description: Mantle: bell-shaped, attenuated posteriorly in the
region of fins (Fig. 1), texture muscular before the cylindrical
Journal of Molluscan Studies (2008) 74: 203–207
# The Author 2008. Published by Oxford University Press on behalf of The Malacological Society of London, all rights reserved.
doi:10.1093/mollus/eyn007
A. I. ARKHIPKIN AND V. V. LAPTIKHOVSKY
Figure 1. Asperoteuthis nesisi n. sp. Holotype. A. Dorsal view. B. Ventral view (scale bar ¼ 10 cm). C. Sucker from the proximal part of arm III
(scale bar ¼ 1 mm). D. Funnel cartilage (scale bar ¼ 1 cm). E. Transversal row of radular teeth (scale bar ¼ 1 mm). F. Ventral view of the eye
with single photophore (scale bar ¼ 1 cm). Drawings by Z.N. Shcherbich.
Arms: long, attenuate; formula IV . II . I . III, difference between the first three pairs of arms small (Fig. 1). Arms
IV slightly thicker than others. Aboral keel low and thick, on
distal 3/4 of arms I, and along distal 4/5 of arms II –IV. All
arms with small protective membranes. Suckers globular,
stalked, arranged in two longitudinal series (Fig. 1C).
Proximal half of arms I –III with 41 –42 suckers, that of arms
IV with 48 suckers. Two series of suckers converge on the
distal 1/5 –1/6 of arms I– III into one series of much smaller
suckers that run almost to arm tip. Suckers absent on the distal
2/5 of arms IV. Largest suckers at a half of arm length. Arms
II with largest suckers, arms IV with smallest. Horny sucker
rings with 12–14 sharp, triangular, teeth distally. Proximal
margin of sucker ring smooth with flat thin ridge (Fig. 2C).
region between fins, wall thin, covered, especially anteriorly,
with shallow oval depressions with relatively darker pigment
(Fig. 2D). Muscle layer beneath depressions vacuolated. Skin
covered with tiny cartilaginous pointed tubercles (0.1 –0.2 mm
high), visible on skin dried by filter paper. Fins: muscular, distorted oval shape, widest at 1/3 of their anterior part, slightly
wider than long (Fig. 1A, B; Table 1). Free anterior lobes of
fins about 1/10 of the total fin length.
Locking-apparatus: funnel component low, asymmetrical
antitragus, long, weakly developed tragus (Figs 1D, 2A).
Mantle component with posterior, well-developed knob and
groove complimentary to funnel cartilage antitragus (Fig. 2B).
Funnel valve weak. Head small, cylindrical. Eyes large, with
single long longitudinal photophore on ventral surface (Fig. 1F).
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NEW SPECIES OF DEEPWATER SQUID
Figure 2. Asperoteuthis nesisi n. sp. Holotype. A. Frontal view of the funnel component of locking cartilage. B. Frontal view of the mantle
component of locking cartilage; top is anterior. C. Oral view of the largest sucker ring of arm III with 12 distal teeth. D. External view of the
ventral mantle surface. E. Lateral view of the lower mandible. F. Lateral-oblique view of the lower mandible. G. Lateral view of the upper
mandible with broad basal cusp on the shoulder. Abbreviations: at, antitragus; t, tragus; vac, vacuolated depressions.
Remnants of two tentacles short, slender stumps. Tentacles
may have been damaged in the trawl, or are absent in maturing specimens.
Gladius not extracted, posterior end broken. Buccal connectives attach to ventral side of arms IV.
Mandibles: yellowish-brown, rostra black (Fig. 2E –G).
Upper mandible with long lateral walls, shoulder with pointed
cusp in obtuse triangular shape. Lateral walls of lower mandible with well-developed longitudinal ridge. Radula with seven
teeth (Fig. 1E) in each row.
Standard measurements and sucker counts for the holotype
are given in Table 1.
the water mass suggests that the species may occur in mesoand bathypelagic layers of the Southern Ocean.
Remarks: Specific features of A. acanthoderma, A. mangoldae and
A. nesisi n. sp. are summarized in Table 2. Those of A. lui are
missing because of the extremely poor condition of the only
known specimen (Salcedo-Vargas, 1999). Asperoteuthis nesisi
n. sp. differs from other species of the genus by: (1) muscular
consistency of mantle, fin and arms; (2) fin shape; (3) presence
of oval skin depressions; (4) shape and structure of funnellocking apparatus; (5) higher number and triangular shape of
teeth on inner ring of arm sucker; (6) presence of broad-based
cusp on the shoulder of the upper mandible and (7) greater
number of suckers on the proximal half of the arms. Skin
depressions on the mantle and fin have not been reported in
any other species of Asperoteuthis, even in adult specimens in
Distribution: Single known specimen from the Antarctic
Intermediate Water Mass near the northern flank of
Burdwood Bank (Southwest Atlantic). The Antarctic origin of
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A. I. ARKHIPKIN AND V. V. LAPTIKHOVSKY
Table 1. Measurements and indices of Asperoteuthis nesisi n. sp.
Characters
Mantle length (with posterior end of the
Measurement
% of mantle
(mm)
length
Table 2. Specific features of the three species of Asperoteuthis
(Chiroteuthidae).
363
Character
A. acanthoderma
A. mangoldae
A. nesisi n. sp.
Consistency of
Muscular but
Almost
Muscular
arms and
gladius broken)
Head length
88
24.2
mantle
Head width
36
9.9
Fin shape
weak
Longitudinally
elongate oval
gelatinous
Transversally
Transversally
elongate
elongate oval
Distance between eyes
18
5.0
Maximum mantle width
100
27.5
oval with
with maximum
Fin length
220
60.6
maximum
width anterior
Fin width (only half of the fin)
123
33.9
width in the
middle
Arm length I
312
86.0
Arm length II
335
92.3
Skin tubercles
Present
Absent
Present
Arm length III
303
83.5
Skin vacuolated
Absent
Absent
Present
Arm length IV
377
103.9
depressions
Sucker diameter arm I
3
0.8
Funnel-locking
Narrow oval with
Narrow and
Wide ovoid with
Sucker diameter arm II
3.5
1.0
apparatus
Sucker diameter arm III
4
Sucker diameter arm IV
2.5
weakly
bent oval
weak tragus;
1.1
developed
with slightly
well-developed
0.7
tragus;
developed
antitragus
well-developed
tragus; no
Eye diameter
27
7.4
Eye photophore length
28
7.7
Number of suckers on proximal half of
41
antitragus
Number of
3 –4 broadly
8– 10 slender
12 – 14 triangular
truncated
sharp teeth
of arms
Arm sucker
41
chitinous ring
arm III
Number of suckers on proximal half of
41 – 48
proximal half
42
arm II
Number of suckers on proximal half of
24 –25
suckers on
arm I
Number of suckers on proximal half of
antitragus
Unknown
rounded teeth
teeth
bears
48
Club suckers
arm IV
9 blunt teeth on
25 truncated
the distal half of
teeth, large
the ring
on the distal
Unknown
half and
small on the
good condition (A. acanthoderma: Tsuchiya & Okutani, 1993;
A. mangoldae: Young et al., 2007). Asperoteuthis mangoldae Young
et al., 2007 differs from both A. acanthoderma (Lu, 1977) and
A. nesisi n. sp. by a set of important features including the
structure of the funnel cartilage (simple groove in A. mangoldae
vs elongated triangle with an antitragus in the other two),
length and structure of arms IV (long and specialized in
A. mangoldae vs similar in length to other arms and nonspecialized), consistency of the mantle (gelatinous in A. mangoldae vs
muscular) and type of skin (cartilaginous tubercles absent in
A. mangoldae vs present). The only feature of A. mangoldae
similar to A. acanthoderma is the structure of the tentacular club
with suckers present only on its distal half. In our opinion,
A. acanthoderma and A. nesisi n. sp. are more closely related to
each other than either is to A. mangoldae. Asperoteuthis lui was
described from a single rather digested specimen with its body
and funnel ‘mutilated’, and without horny sucker rings due to
digestion (Salcedo-Vargas, 1999). This species was based solely
on the features of the tentacle which was cited as ‘in good condition’. However, the tentacle suckers lack of horny rings,
casting some doubt on its condition. The lack of elongated
photophore on the tip of the tentacle club might also be due to
digestion. Salcedo-Vargas (1999) mentioned also that A. lui
had ‘enlarged suckers located in the middle to last portion of
arms II and III’, a condition that is unique among the species
of Asperoteuthis. Until a better specimen of A. lui is found, it is
impossible to identify a relationship between it and the other
three species of Asperoteuthis.
It should be noted that two partially digested specimens of
‘Mastigoteuthis? A’ taken from sperm whale stomachs in South
Georgia in 1964 (Clarke, 1980: 191, fig. 155) look very
proximal half
of the inner
ring
Terminal club
photophore
Large, oval,
Small, nearly
associated with
circular,
a short, blunt
associated
terminal papilla
with long,
Unknown
slender
terminal
papilla
Shoulder of
Slightly convex
Straight and
Straight with
upper
and irregular in
irregular in
prominent
mandible
outline
outline
wide-based
triangle cusp
Asperoteuthis lui is omitted because data for the holotype are extremely
limited. It is separated from all other species in the genus by structure of
tentacular club and position of largest arm suckers (see text).
similar to those of A. nesisi n. sp. Clarke (1980) did not
mention the characteristic skin tubercles and armature of
sucker rings, because these were absent because of digestion.
He assigned his specimens to the genus Mastigoteuthis ‘as a
temporary convenience since the beaks were grouped as
Mastigoteuthis before specimens became available’. Based on
both the illustrations and descriptions Clarke (1980) provided
of the fin and mantle, small funnel valve and characteristic
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NEW SPECIES OF DEEPWATER SQUID
funnel-locking cartilage, both Clarke’s specimens probably
belong to A. nesisi n. sp.
ACKNOWLEDGMENTS
We acknowledge the assistance of the crew of the R/V Dorada
in squid sampling. We are grateful to Dr R.E. Young
(Department of Oceanography, University of Hawaii,
Honolulu, USA) for his valuable comments on systematics of
the family Chiroteuthidae and editing of the English of the
manuscript, and to Annie Lindgren (Department of Evolution,
Ecology and Organismal Biology, Ohio State University,
Columbus, USA) for DNA processing. We thank Zhanna
Shcherbich (Fisheries Department, Falkland Islands) for drawings of the specimen.
DISCUSSION
In his description of Chiroteuthis acanthoderma, Lu (1977) noted
that the species ‘differs considerably from other species of the
genus [Chiroteuthis]’ and it is ‘unique in having cartilaginous
tubercles, . . .and suckers only on the distal half of the tentacular club’. However, he did not discuss the systematic position of
his newly described species, as the systematics of the genus
Chiroteuthis needed revision.
Nesis (1980) transferred C. acanthoderma to the genus
Asperoteuthis as A. famelica (Berry, 1909) and pointed out that
this species is distinguished from all other chiroteuthids by the
funnel cartilage structure, armament of arm suckers (without
large central tooth) and presence of skin tubercles. Moreover,
the only other chiroteuthid squid with the structure of the
arms IV similar to other pairs of arms, as in Asperoteuthis, is
Planctoteuthis levimana (Lönnberg, 1896).
The structure of the funnel-locking cartilage differs among
species within the Chiroteuthidae from a simple groove with
slightly developed longitudinal tragus (Planctoteuthis) to a very
specialized ear-like structure with well-developed tragus and
antitragus (Chiroteuthis) (Nesis, 1987; Young, 1978). Asperoteuthis
has an intermediate type of the cartilage either with welldeveloped antitragus (A. acanthoderma and A. nesisi n. sp.) or
without it (A. mangoldae). All species of Asperoteuthis probably
have the secondary fin supporting the posterior end of the
gladius as in A. acanthoderma (Tsuchiya & Okutani, 1993). This
secondary fin is almost always broken in a trawl catch, leaving
only the main fin with the broken gladius at its posterior tip.
Presence of the second fin in Asperoteuthis might indicate its
closer relation to another chiroteuthid Grimalditeuthis bonplandi,
however the structure of the funnel-locking cartilages is different in these two genera (Nesis, 1987). Further study of holotypes and molecular genetics will help to resolve the systematic
placement (Sweeney & Roper, 1998) of these strange looking
meso- and bathypelagic squid.
Similarity of environmental conditions of mesopelagic and
especially bathypelagic habitats in different parts of the world
ocean favours cosmopolitan distribution of deepwater animals.
This is especially characteristic for species that do not have
epipelagic planktonic larvae and therefore are not associated
with more latitudinally stratified epipelagic habitats. Squid
from two other genera of Chiroteuthidae (Chiroteuthis and
Planctoteuthis) are cosmopolitan in temperate and tropical
waters (Nesis, 1987). Our findings extend the range of
Asperoteuthis from the tropical Pacific to the sub-Antarctic
waters of the Southwest Atlantic, supporting a wider, or possibly cosmopolitan, distribution of squid from this deepwater
genus.
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