BULLETIN OF MARINE SCIENCE, 72(3): 941–953, 2003 NEW TAXA PAPER
A NEW SPECIES OF NEMATOLAMPAS (CEPHALOPODA: OEGOPSIDA) FROM THE WESTERN CENTRAL ATLANTIC WITH AN OVERVIEW OF THE FAMILY LYCOTEUTHIDAE
Freddy Arocha
ABSTRACT A new squid species of the family Lycoteuthidae, Nematolampas venezuelensis is rec- ognized from the central western Atlantic Ocean. The new taxon is characterized by the threadlike, greatly elongated ventro-lateral arms and the presence of a single, large pho- tophore embedded in the musculature of the fin/mantle complex. An overview of the Family Lycoteuthidae is presented based on the examination of the holotypes of Nematolampas regalis and Oregoniateuthis springeri, reported specimens of Selenoteuthis scintillans and Lampadioteuthis megaleia, and a review of the literature on the family.
Among the cephalopods collected during the four cruises of the R/V DR. FRIDTJOF NANSEN off the coasts of Venezuela in 1988, were four lycoteuthid squids with very elongated, threadlike ventro-lateral arms. Gross anatomical features initially suggested that the speci- mens were attributable to Nematolampas regalis Berry, 1913. However, closer examina- tion revealed distinctions from Berry’s definition of N. regalis, and showed them to rep- resent a new species from the central western Atlantic Ocean (Arocha, 1991). The Family Lycoteuthidae is poorly known due to the lack of sufficient comparative material that would enable a full review. However, in the past decade several advances have been made. There have been important collections of lycoteuthid species made off New Zealand in the southwest Pacific (Förch and Uozumi, 1990) and Namibia in the southeast Atlantic (Villanueva and Sanchez, 1993), that have led to the clarification of the Lycoteuthis diadema problem (Toll, 1983; see ‘Discussion’). In addition, the type specimens for N. regalis and Oregoniateuthis springeri Voss, 1956, and previously re- ported specimens of Selenoteuthis scintillans Voss, 1958 and Lampadioteuthis megaleia Berry, 1916 were available for examination in the present study. These circumstances, together with a review of the literature, enabled the comparison of the morphological characters of the recognized species and allowed a clarification of the classification of the family members. Measurements and indices are as defined by Roper and Voss (1983). The following abbreviations are used: USNM, U.S. National Museum of Natural History; UMML, Marine Invertebrate Museum, Rosenstiel School of Marine and Atmospheric Science, University of Miami; the R/V NANSEN, is a fisheries research vessel from Norway.
Family LYCOTEUTHIDAE Berry, 1914 Subfamily Lycoteuthinae Pfeffer, 1908 Genus Nematolampas Berry, 1913
Diagnosis.—Lycoteuthid with ventro-lateral (III) arms longer than mantle, devoid of suckers in distal part of arms in males (females not known). Photophores present near tips
941 942 BULLETIN OF MARINE SCIENCE, VOL. 72, NO. 3, 2003
the four dorsal arms and along the entire length of the ventro-leteral (III) arms. Ten pho- tophores within mantle cavity. Males with single genitalia. Type species.—Nematolampas regalis Berry, 1913.
Nematolampas venezuelensis new species (Figures 1–4, Tables 1,2)
Material Examined.—Holotype: One male, mantle length 87.5 mm, R/V NANSEN sta 1143, northeast of Golfo Triste, Venezuela, 10°55’N, 67°57’W, 371–383 m, bottom trawl, December 2, 1988, USNM 817596. Paratypes: One male, mantle length 67.7 mm, R/V NANSEN sta 1129, off Punta Chuspa, Venezuela, 11°02’N, 62°21’W, 355–365 m, bottom trawl, November 28, 1988, UMML 31.3143. Two males, mantle lengths 65.0 and 58.2 mm, R/V NANSEN sta 146, south of Los Hermanos Islands, Venezuela, 11°20’N, 64°26’W, 298 m, bottom trawl, February 19, 1988, UMML 31.3144. Comparative Material.—Nematolampas regalis. Holotype, One male, mantle length 32 mm, R.S. BELL, on beach Sunday Island, Kermadec Island Group-New Zealand, 1910, USNM 815721. Paratype, one male, mantle length 26 mm, R.S. BELL, on beach Sunday Island, Kermadec Island Group-New Zealand, 1910, USNM 816592. Lycoteuthis springeri. Holotype, One male, mantle length 80.0 mm, R/V OREGON sta 382, 29°11.5'N, 88°7.5'W, 367 m, June 21, 1951, USNM 575090. One male, mantle length 97.0 mm, R/V OREGON sta 3296, 28°36'N, 89°48'W, 439–936 m, August 21, 1961, UMML 31.376. One female, mantle length 124 mm, R/V COLUMBUS ISELIN sta 256, 23°37'N, 77°6'W, 1360 m in stom- ach of Synaphobranchus brevidorsalis, November 1, 1974, UMML 31.1735. One fe- male, mantle length 62 mm, R/V OREGON sta 481, 28°57'N, 88°40.5'W, 378 m, Septem- ber 7, 1951, UMML 31.174. Two females, mantle length 48–56 mm, R/V SILVER BAY sta 1198, 24°11'N, 83°31'W, 360 m, June 9, 1959, UMML 31.228. Selenoteuthis scintillians. One male, mantle length 17 mm, R/V COLUMBUS ISELIN sta 206, 23°44.2'–23°46.0'N, 81°21.5'–81°24.5'W, 10'IKMT, 80 m, February 19–20, 1974, UMML 31.2271. One fe- male, mantle length 26 mm, R/V GILLISS sta 25, 20°26.5'N, 83°57'W, 4383 m, July 23, 1972, UMML 31.1503. two females, mantle length 29–30 mm, R/V OREGON sta 2191, 25°11'N,89°50'W, 3240 m, May 23, 1958, UMML 31.227. Lampadioteuthis megaleia. One male, mantle length 31 mm, DANA sta 4173, 40°46'N, 18°35'W, 400 m, June 3, 1931, UMML 31.2968. Diagnosis.—Lycoteuthid with greatly elongated, ventro-lateral arms in males (females not known), with threadlike distal portions, which are devoid of suckers. Ten photo- phores on viscera; numerous photophores on dorso- and ventro-lateral arms, head and mantle; five photophores arranged in one row on the ventral periphery of each eye. A single, large photophore embedded in musculature of fin/mantle complex near the base of tail, and on dorsal side, five beadlike photophores serially arranged on elongated pos- terior end of mantle. Description.—Known only from immature males, largest 87.5 mm ML. Mantle mod- erately stout, cylindro-conical, tapering posteriorly to a narrow pointed tail; mantle width about one third of length (MWI 26.0–34.0–37.3); anterior dorsal margin slightly pro- duced; ventral margin somewhat indented beneath funnel (Fig. 1). Fins large, muscular and sharply angled laterally, anterior margin convex, concave posterior margin termi- nally narrows and extends along elongated end of mantle. Combined width about equal AROCHA: NEW SPECIES OF LYCOTEUTHID SQUID 943
Figure 1. Nematolampas venezuelensis n. sp. Dorsal and ventral view (87.5 mm–ML, holotype). to mantle length (FWI 74.7–84.6–96.6) and length about one third of mantle length (FLI 33.9–36.4–38.8). Anterior fin margins thin with conspicuous notch where they fuse to mantle; posterior margins thick (Fig. 1). Gladius narrow and darkly pigmented (Fig. 2). Rachis narrow anteriorly (RWI 4.6– 4.6–6.2), with straight convergent margins and forming dorsally a strong keel. The vanes are hourglass shaped being slightly widened in midsection, then constricted and posteri- orly slightly expanded. The posterior tip of gladius is missing, although it appears to terminate in a cup-shape conus. Funnel large and stout with two large dorsal supports, anterior quarter projects free of head. Mantle locking apparatus composed of simple, straight funnel member and corre- sponding slightly raised ridge on inner surface of mantle. Funnel organ indistinguishable in available specimens. Head moderate in size, rounded, flat dorsally and slightly excavated ventrally; eyes large (EDI 18.3–19.7–21.5), eyelids with well-defined anterior sinus. Ventral surface of each eyeball has five photophores arranged in a single row (Fig. 3A), round central one clearly the largest, remaining four round and small, all with distinct reddish hue. Arms unequal in length, in order 3 > 2 > 4 > 1 except in smallest specimen (Table 1). Dorsal pair of arms with slight fleshy median ridge on distal halves. Dorso-lateral pair almost as long as mantle (ALI II 80.3–92.5–119.8), keeled on proximal half, and attenu- ated distally into filament. Proximal third of each dorso-lateral arm bears nine equally 944 BULLETIN OF MARINE SCIENCE, VOL. 72, NO. 3, 2003
Figure 2. Nematolampas venezuelensis n. sp. Gladius, dorsal view with cross sections and side view (87.5 mm–ML, holotype).
spaced round light organs arranged aborally in single series embedded in dermal layer of skin. Ventro-lateral arms more than 1 1/2 times length of mantle (ALI III 171.4–178.3– 191.9), keeled on proximal third and distally attenuated to threadlike filament. Proximal two thirds of each arm bears 19 equally spaced, round light organs arranged aborally in single series as on dorso-lateral arms. Protective trabeculate membranes on suckered portions of all arms. Arms II and III devoid of suckers on filamentous distal portions. Arm suckers stalked, hooded and biserial. Chitinous rings of suckers on midportions of arms with 9–12 sharp teeth on distal border, proximal edge with bluntly pointed teeth (Fig. 3B). Ventral pair of arms strongly keeled. Hectocotylus absent in available males, all immature. Tentacles, severed in all specimens except the holotype in which the left tentacle re- mained, long (TtLI 125.7), stout, thick and fleshy at base, attenuate for distal two-thirds. Single available club in poor condition; aboral surface with well-developed swimming keel extending from dactylus to distal part of manus; oral surface bordered by trabeculate AROCHA: NEW SPECIES OF LYCOTEUTHID SQUID 945
Figure 3. Nematolampas venezuelensis n.sp. A) photophore pattern on eye (87.5 mm–ml, holotype); B) large sucker from basal portion of arm IV (87.5 mm–ml, holotype); C) Upper mandibule and lower mandibule (69.7 mm–ML); D) radular teeth of one side (69.7 mm–ml); E) Internal photophore pattern and connective tissue (87.5 mm–ml, holotype). protective membranes. Suckers on carpus in four series. Manus and dactylus with small suckers in four series. Two large photophores embedded on oral surface of tentacular stalk, one at about one quarter the length of tentacle and second at base of club. Buccal membrane reddish purple in preserved specimen, with eight lappets and eight connectives, attached dorsally to arms I, II and IV, ventrally to III; oral surface thickly covered with fleshy papillae. Beaks are as illustrated (Fig. 3C). Lower beak with thick ridge bisecting lateral wall; dark spot near posterior base of each lateral wall evident in 946 BULLETIN OF MARINE SCIENCE, VOL. 72, NO. 3, 2003
Table 1. Mantle length and indices of type specimens of Nematolampas venezuelensis new species.
U3MML31.3144 U6MML31.314 USNM 81759 Pearatypes Holotyp MmL 5m8.2 m 6m5.0 m 6m9.7 m 87.5 m Index M4WI 325. 337. 307. 26. H2LI 293. 128. 188. 16. H9WI 202. 252. 178. 23. E9DI 108. 250. 231. 18. F4LI 386. 398. 333. 36. F6WI 976. 759. 877. 74. ALI I3559. 461. 431. 30. I8I 1319. 880. 8+1. 88.0 I9II 1+91. 1911.3 1472. 171. I7V 499. 465. 492. 38. A4L formula 31:2:1: 31:2:4: 31:2:4: 3:2:4: T---7tLI 125. C---4LI 22. G4LI 926. 876. 834. 74. G5WI 171. 79. 79. 4. R2WI 61. 56. 49. 3.
lower beak of 69.7 mm–ML. Radula as shown in figure 3D, with homodont teeth; mar- ginal plates absent. Outer integument almost intact in type specimen at capture. Head bears eight photo- phores: two at bases of third arms, one on either side on dorsal surface near posterior margin of head, one small photophore at anterior edge and another at posterior edge of each eyelid. On dorsal surface of mantle, anterior to fins, integument bears five pairs of photophores. On dorsal surface of fins, integument bears three pairs of photophores, two pairs on dorsal posterior margins of fins, and one pair on anterior end of fin insertion on mantle. On the ventral surface of mantle, six pairs of photophores are present near or along lateral margins. On the fin/mantle complex, one large photophore lies in pocket in musculature at base of tail, postero-dorsal to conus of gladius. Opening in musculature dorsal to photophore covered by translucent connective tissue that forms a dorsally directed window; three small photophores are found around translucent tissue. Five round photophores deeply buried in dorsal midline of tail terminating in teardrop-shaped organ at distal extremity. One pair of closely set photophores present laterally near midpoint of tail and one photo- phore ventral to embedded large photophore. Photophores within the mantle cavity consist of two round anal photophores, one on each side of the anal opening, a set of three abdominal photophores lying transversely on the visceral mass, and one elongate photophore at the base of each gill. A complex of three photophores are located transversely in the posterior ventral mantle fastened to the inner wall of the mantle. All photophores within the mantle cavity are connected by a AROCHA: NEW SPECIES OF LYCOTEUTHID SQUID 947
Figure 4. Geographical distribution of Nematolampas venezuelensis n.sp. Holotype: star; Paratypes: filled circles.
strip of tissue, with the exception of the anal organ (Fig. 3E). Total of 127 photophores present on entire specimen. Male with single genitalia, which is on the right side of visceral organs. All specimens were immature, but in the largest, the holotype, a short inconspicuous penis is present. Holotype.—Male ML 87.5 mm, U.S. National Museum of Natural History, USNM 817596. Type locality.—Northeast of Golfo Triste, Venezuela. 10°55'N, 67°57'W, December 2, 1988, 371–383 m, bottom trawl. Distribution.—Nematolampas venezuelensis is known only from the waters off the coasts of Venezuela between 10–12° N and 64–68° W (Fig. 4). All specimens were caught near the bottom during the night at 298–385 m, where bottom temperatures were 12–15° C. Etymology.—The specific name venezuelensis is given after the country from whose waters the specimens were found.
DISCUSSION
NEMATOLAMPAS VENEZUELENSIS.— The four male N. venezuelensis specimens examined in this study were all immature and collected in three different trawl stations off Venezuela. Because the purpose of the R/V NANSEN research cruises, during which the specimens were taken, was to survey demersal fishery resources (Anon., 1989) and not to collect specimens for scientific study, the majority of the cephalopods taken were badly dam- aged. In addition, most of the specimens were held frozen prior to permanent preserva- tion in 70% ethanol at the laboratory. Only the holotype, was fixed in 10% formalin immediately after capture and subsequently transferred to 70% ethanol. Nonetheless, the four specimens of N. venezuelensis were in sufficiently good condition to determine the distinctive characters needed to verify a new species. Nematolampas regalis, known only from Berry’s original two specimens (Berry, 1913, 1914) is closely related to the new species. An examination of Berry’s specimens (the 948 BULLETIN OF MARINE SCIENCE, VOL. 72, NO. 3, 2003
holotype body in good condition, portion of arms missing) showed them to be males, and a comparison of the two species showed them to share greatly elongated ventro-lateral arms with threadlike distal portions devoid of suckers (now left arm only present in holo- type of N. regalis); ten similarly placed photophores within the mantle cavity; five photo- phores arranged on one row on the ventral periphery of each eye; single male genitalia; and no modification on the ventral arms that would indicate hectocotylization. Nematolampas regalis, differs in the presence of one photophore near the tip of each dorsal and dorso-lateral arm, in contrast to N. venezuelensis, in which only the dorso- lateral arm pair bears photophores, which are embedded in the dermal layer in the proxi- mal third, and the dorso-lateral arms attenuate distally into a filament, a condition absent in N. regalis for this arm pair. Additionally, a pair of photophores are found on the ventral surface of the mantle near the posterior end of body at the attachment points of the poste- rior edges of the fins to the mantle in N. regalis, an arrangement absent in N. venezuelensis. Also, the elongated posterior end of the mantle in N. venezuelensis displays a distinctive large photophore embedded in the musculature of the fin/mantle complex and five bead- like, serially arranged photophores, features absent in N. regalis. The number of photo- phores on each ventro-lateral (III) arm in N. regalis exceed 31, in contrast to N. venezuelensis which has 19 photophores on each arm III. The 32 mm ML holotype of N. regalis was a mature male with the Needham sac full of spermatophores. No signs of hectocotylization were observed on either the ventral or dorsal arms. The largest male of the new species, the 87.5 mm–ml holotype, showed no evidence of advanced maturity, or of hectocotylization. These findings indicate that N. regalis males mature at a much smaller size than N. venezuelensis. OVERVIEW OF THE FAMILY.—Examination of representatives of five of the six species known to occur in the Lycoteuthidae (see comparative material) yields a better under- standing of the relationships within the family. The six species can be differentiated by the characters presented in Table 2. Lampadioteuthis megaleia is separated from the rest of the species by its unique form of the gladius, which has diamond-shaped, broad vanes with anterolateral borders slightly concave and a conus with a terminal acute rostrum as also observed in earlier studies (Young, 1964; Toll, 1983). Hess (1987) also found that spermatophore morphology separated L. megaleia from the other species. Other distinc- tive features include the presence of a hectocotylus on the right ventral arm and the dis- tinct pattern of the photophores in the mantle cavity, and on the eyes and tentacles. Sev- eral authors have suggested (Voss, 1962; Young, 1964; Nesis, 1987) that the single male genitalia present in L. megaleia was an additional character that helped separate this species into a separate subfamily within the Lycoteuthidae. The findings of the present study disprove this proposed difference as both species of Nematolampas also have single male genitalia. Regardless, the remaining distinctive characters of this species, including differences in photophore arrangement, continue to support its separation into the sub- family Lampadioteuthinae. With the addition of N. venezuelensis, the subfamily Lycoteuthinae now comprises three genera and five species (Table 2). Lycoteuthis diadema was first described by Chun (1900) as Enoploteuthis diadema, but in 1903 he placed this species in Pfeffer´s genus Lycoteuthis. Later, in 1912, Chun’s specimens were shown to be identical to Pfeffer´s type Lycoteuthis jattai Pfeffer, 1900, which Pfeffer (1912) considered “jattai” to be a “museum name” and abandoned it (Vecchione and Young, 1999). Voss (1958, 1962) dem- onstrated that Pfeffer´s paralarval Astenoteuthion planctonicum and Robson´s AROCHA: NEW SPECIES OF LYCOTEUTHID SQUID 949
x
n
y
e
i
l
d
p
l
I
t
n
a
a
I
i
a
a
I
r
e
d
I
a
l
n
s
e
e
e
e
a
a
n
s
e
r
m
m
n
e
r
o
t
r
r
V
m
n
o
n
a
,
h
e
d
r
.
i
t
o
e
e
r
e
d
a
e
l
c
n
t
i
,
v
e
i
p
l
c
s
f
t
c
a
i
d
a
a
n
o
n
s
l
i
r
t
e
*
h
t
a
e
o
a
h
n
s
s
s
p
r
s
r
c
e
t
p
e
a
l
d
o
n
t
s
i
n
l
n
e
h
e
a
t
s
h
o
r
f
a
a
e
s
a
g
g
c
p
r
h
l
o
n
n
v
e
e
n
t
o
i
i
b
p
s
g
r
t
z
a
e
,
n
r
n
u
i
u
r
e
l
o
m
i
s
o
l
i
s
a
u
r
c
r
s
a
l
a
h
o
,
y
d
a
a
o
w
o
u
a
t
n
s
b
p
d
t
n
a
p
h
h
e
i
l
a
p
o
e
y
b
t
,
l
r
l
p
c
g
+
l
m
m
Z
a
a
s
n
l
a
a
d
o
a
o
2
a
e
e
y
a
f
a
t
r
c
t
l
n
2
l
n
t
e
i
r
n
6
n
m
t
w
o
c
o
o
i
r
o
u
3
a
o
f
n
e
i
o
e
t
n
l
e
e
m
h
d
o
s
n
r
n
n
o
o
h
:
s
m
r
a
s
a
i
r
i
l
L
h
p
e
N
e
n
e
e
p
y
,
e
o
o
t
l
e
g
i
s
l
t
t
a
n
r
m
e
m
f
m
f
o
o
o
s
e
r
s
d
a
n
a
b
a
e
r
o
o
f
e
,
v
i
e
o 2
a
f
n
b
i
I
w
w
w
o e
N
A
V
V
f
t
t
t
t t
p
p 9
M
I m
b
N
N
a
o
O
o
S
f
s
y
,
o
l
I
r
e
l
I
t
o
a
e
e
d
a
t
I
i
a
s
n
h
n
l
i
n
a
i
s
p
y
e
i
i
l
b
m
l
y
r
m
m
m
t
a
e
r
o
e
o
6
r
i
a
m
a
d
p
f
d
e
1
d
t
b
f
l
o
e
b
m
e
a
d
a
o
a
n
e
r
r
5
m
n
r
o
t
n
e
r
e
a
2
e
o
a
n
,
n
o
l
c
d
i
1
i
r
e
I
e
e
a
r
a
e
r
V
l
I
a
t
o
f
:
v
e
e
c
r
.
t
c
c
m
e
n
t
s
i
l
i
a
e
u
n
e
n
s
r
t
a
e
f
s
h
t
p
a
m
o
o
t
t
n
r
s
i
a
l
p
m
e
n
u
e
a
s
d
n
a
t
h
i
r
e
e
e
e
o
s
s
e
o
t
e
r
f
n
e
F
r
e
e
g
p
c
i
o
g
i
s
h
o
r
n
o
n
t
e
n
p
t
c
s
n
s
s
h
o
d
m
o
z
a
i
n
o
e
a
,
y
t
i
u
e
c
r
l
l
r
s
a
,
l
i
r
r
,
n
L
m
l
a
n
g
p
a
s
l
i
t
y
d
o
a
d
r
s
t
n
e
o
g
b
a
a
8
e
h
l
s
i
u
A
l
a
o
n
y
s
t
9m
p
a
p
e
o
l
h
o
i
c
g
t
o
a
r
l
a
d
−
o
c
l
h
s
h
o
m
t
w
f
a
.
o
(
h
t
t
n
n
n
t
e
e
i
s
o
e
o
s
h
c
r
n
u
t
u a
8
o
e
n
n
,
a
h
w
l
r
l
e
i
p
e
a
e
s
n
o
s
e
o
:
p
t
d
s
a
p
h
r b
h
o
e
t
u
e
i
e e
y
t
y
u
,
t
o
s
l
r
r
t
h
p
n
r
m
e
e
h
n
m
c
c
o o
o
i
o
+
5
e
a
r
r
t
a
e
o
o
g
v
o
y
r a
h
f
0
i
a
0
i
o
u
w w
w
n
n
V
Li Vg
f
f
t t
t
Pt Pe M Nd
p i
i
Lg Nd
o
e
c
3
1+
e
t
o
s
,
l
c
I
r
e
l
I
t
a
e
y
a
t
I
a
s
n
h
n
L
i
n
r
a
i
s
p
y
i
i
l
e
b
m
y
m
l
r
m
m
l
h
m
t
i
a
r
o
e
o
m
r
i
p
a
a
d
p
d
e
s
m
d
t
3
i
b
f
l
e
b
a
e
1
a
1
a
o
a
n
e
m
F
r
1
m
d
r
o
t
n
e
r
e
e
:
n
o
a
n
,
o
c
d
i
H
e
a
i
h
e
e
r
a
e
l
t
r
V
l
t
f
v
I
e
a
n
e
c
r
.
f
t
c
I
r
t
i
a
e
u
n
o
n
s
m
e
r
t
s
s
t
p
a
o
e
o
h
n
s
s
t
a
l
m
p
F
e
e
e
d
n
r
u
a
t
h
i
e
a
e
s
s
e
a
o
o
c
r
f
e
r
r
m
e
g
c
e
S
s
h
o
e
r
n
o
n
e
n
t
p
s
i
m
s
g
h
o
d
e
z
a
s
e
a
,
i
t
i
u
c
r
s
l
l
s
h
r
l
i
7
r
,
n
t
e
e
a
g
a
l
i
o
y
d
8
a
d
r
r
h
l
t
n
b
a
a
t
10
e
o
s
i
u
l
a
o
m
y
s
t
p
−
e
h
l
h
o
i
c
g
o
g
a
r
l
a
d
c
p
r
h
h
o
n
s
f
a
o
f
h
t
t
n
5
i
n
n
o
e
i
s
u
t
o
s
t
h
c
r
n
u
u a
1
o
t
e
n
s
a
p
l
r
l
o
e
p
e
a
e
o
s
n
n
r
o
:
n
p
r
t
s
a
r b
h
h
o
e
e
e
a
e e
a
a
y
u
,
t
o
l
p
r
s
s
t
r
n
m
e
e
c
p
n
m
c
c
o o
o
i
o
a
r
b
b
e
a
o
c
e
v
o
y
a
h
2
f
i
o
w w
w
s
n
n
A)
As
Li Vg
V
f
f
t t
t
9+
Pd M
p i
5
i
Lg Nd
Oc
o
n
i
d
n
e
e
l
r
o
s
e
c
i
l
o
t
u
s
t
a
t
u
n
e
s
s
m
x
l
n
b
r
a
e
r
s
i
e
s
a
e
r
e
l
r
t
a
t
e
e
e
t
m
t
o
l
r
l
p
c
s
y
a
n
n
n
h
a
o
i
t
a
n
m
i
i
o
o
o
p
e
r
h
m
r
l
m
D
o
a
o
t
p
s
s
s
s
u
t
c
/
h
t
n
n
c
o
n
e
e
e
e
o
i
i
t
i
y
r
r
r
r
a
a
e
C
l
r
h
h
o
l
o
o
o
o
i
I
a
t
e
.
m
p
m
i
p
h
I I V
h
h
h
h
t
s
l
n
2
I I I
m
a
t
p
r
c
p
p
p
p
a
u
n
a
r
a
a
y
t a
a
i
i
s s s
l
e
o
o
o
o
t
i l g
r
l
t
t
t
t
F
d
a
m
i
l
e
a
n u o
t
/
m m m
i
b
o
o
o
o
b
a
v
z
r r r
e c e
l
h
o a
n
a
h
h
h
h
i
u
a
i
Ag Ag Ad
T4 Tt
f
T
P
P
P8
Pe
c
Gd Oy Ge
C
Go
S5
S 950 BULLETIN OF MARINE SCIENCE, VOL. 72, NO. 3, 2003
s
l
i
u
d
d
p
n
e
n
u
o
l
c
f
u
p
c
a
s
l
a
r
o
o
,
a
a
,
f
p
t
r
s
m
e
s
e
d
o
e
n
e
i
h
v
e
s
e
t
0
n
y
o
e
a
a
n
l
r
v
s
a
3
g
l
c
b
i
o
o
p
v
a
>
d
n
n
e
i
e
l
l
o
e
u
N
d
:
o
o
d
l
t
z
a
q
n
c
i
a
e
d
d
e
l
d
g
e
a
e
l
o
s
l
n
n
l
e
e
y
o
r
,
i
a
r
t
c
y
a
a
l
g
b
l
e
h
m
o
a
w
a
n
a
m
m
n
t
e
t
e
d
d
i
c
t
n
s
e
a
n
d
u
r
n
b
i
,
n
i
i
o
r
r
a
e
l
m
F
o
t
e
t
r
a
h
h
t
a
p
e
l
m
c
s
t
o
b
t
a
m
a
a
i
n
n
e
o
f
o
u
d
u
m
l
r
e
a
h
y
x
o
h
e
d
o
e
b
a
l
t
s
t
m
r
o
l
Z
a
b
,
e
-
d
e
o
r
o
e
t
a
a
y
i
m
i
r
d
t
s
i
n
1
p
r
r
u
w
r
c
d
d
r
a
a
o
n
u
3
t
t
n n
p
e
i
a
c
l
i
e
e
a
a
o
b
t
o
s
o
a
r
n
n
n
a
o o
:
h
i
o
s
t
p
r
p
l
l l
n
u
N
e
r
r
e
e
e
e
,
p
h
m
t
r
l
a
g
i
h
s
s
s
t t
e
e
p
m
e
m
m
f
l
o
s
a t
r
g
t
t
a
r
u
n
t
b
b
b
i i
o o
f
o
i
v
a
a
i
o
e
n
a
n
o
r
i
w
A
A
A
A
L
f
f
f
t
M
p 2
p
D w
L
N N R
O
a
c
a
S
r
y
n
l
I
o
l
i
i
I
s
r
a
i
u
e
e
s
e
t
d
p
n
n
s
e
r
m
e
a
a
o
r
e
r
a
o
e
r
t
,
r
a
m
V
c
p
m
n
o
e
n
h
.
i
f
l
f
t
s
o
d
f
e
r
0
e
c
o
e
c
o
e
,
o
v
e
t
3
p
a
l
t
x
s
t
c
e
a
d
a
:
p
e
n
n
i
s
i
p
n
s
i
e
h
s
r
e
i
a
o
t
n
e
h
a
t
l
s
t
p
t
s
a
c
t
a
b
h
e
a
t
d
l
s
t
n
f
a
n
l
l
e
t
h
a
e
s
m
i
o
r
o
o
a
a
s
a
r
t
a
g
e
r
c
b
i
h
e
l
n
a
n
t
e
n
t
e
r
i
b
F
l
e
p
i
g
s
s
z
a
n
n
,
o
r
s
u
c
n
u
i
r
i
l
a
u
m
o
a
l
s
e
h
u
r
c
i
b
o
l
m
a
l
b
o
t
y
e
a
p
o
d
w
o
,
s
t
a
n
l
d
t
l
n
t
m
i
a
o
t
h
e
i
A
a
o
l
t
y
a
b
g
l
,
h
m
n
l
c
l
t
t
5
g
a
o
c
s
n
l
a
d
.
e
e
s
a
o
u
a
e
y
a
h
a
n
f
r
t
n
2
t
t
n
e
i
i
g
r
n
e
m
p
o
c
o
i
r
r
t
u
3
a
n
n n
o
e
i
I
w
e
e
m
f
a
o
o
e
n
r
s
n
e
o o
I
:
h
,
l
m
r
s
t
i
h
r
l
l l h
o
I
e
e
n
e
e
e
e
t
p
e
o
o
t
l
g
r
i
s
r
e t t t
e
n
e
e
f
a
m
d
o
s
i
r
l
d
d
d
a
n
r
i
b
a
r
o
o o o
n
v
n
a
i
o
e
e
b
n
n
i
i
w
h
o e
At
V
f
t
t
t t
Nt
Mm
p 23
Pe
p
m
Sa Ng Ng Nd
a
e
O
a
S
h
t
s
u
l
e
l
e
t
r
a
t
a
t
t
a
e
s
r
n
o
n
i
n
a
h
i
o
y
c
i
l
p
b
h
m
y
l
i
m
e
s
m
t
r
a
o
l
Le
o
r
i
t
e
a
d
n
d
e
d
n
p
s
t
b
o
e
b
e
a
3
e
l
a
a
n
l
e
*
s
a
m
m
r
a
o
d
r
n
r
e
/
s
t
o
a
n
,
i
l
o
n
d
n
i
m
n
i
a
s
e
i
e
r
e
o
r
V
e
l
f
t
n
e
u
n
I
e
c
v
r
.
t
c
i
e
I
t
z
i
n
a
l
e
e
n
s
e
i
r
t
I
l
n
s
e
t
p
a
l
I
o
n
n
o
s
I
u
e
a
a
p
e
e
m
e
z
n
t
g
h
r
d
s
e
e
m
r
s
e
V
o
a
e
r
f
e
s
a
n
e
c
m
g
f
l
s
h
o
r
n
e
r
n
e
t
f
e
s
n
s
i
h
o
v
a
z
e
e
o
a
,
v
t
a
i
u
c
l
l
i
s
s
r
n
l
i
s
,
n
f
r
a
a
g
a
e
t
l
m
i
y
d
a
d
O
r
r
a
r
e
t
n
b
a
a
d
e
p
s
i
u
o
o
l
,
5
a
o
S
t
y
.
n
p
e
h
e
o
h
h
c
g
l
m
a
r
l
a
a
7
l
d
n
c
s
h
p
o
a
s
f
a
o
(
a
h
8
t
a
l
n
n
n
e
o
i
x
u
o
s
h
c
e
t
n
t
o
u a
r
>
o
n
m
e
t
p
l
r
e
p
a
b
l
o
e
n
o
s
o
:
p
r
a
s
r b
l
h
o
e
n
b
p
h
e
e
a
y
u
,
t
i
i
s
l
g
t
p
r
n
m
m
c
n
r
e
c
o o
l
o
o
m
e
a
r
n
e
a
i
o
e
a
x
v
o
r
i
h
f
o
2
8
i
o
i
w w
a
w
n
N
Vg
V
fl
f
t t
t
t
Pt
Mm
p
Lg
Nd
i
Cc
c
o
Sd
s
19
1
e
l
t
n
a
y
t
m
i
/
v
n
a
i
.
f
c
n
e
n
l
r
o
e
c
i
w
l
o
t
s
t
a
o
t
.
u
n
e
s
n
m
l
n
d
b
a
e
k
r
s
i
s
a
e
e
r
r
t
t
a
t
e
e
e
m
u
t
o
l
r
l
o
s
y
n
a
n
n
n
h
a
n
o
i
t
i
n
i
t
i
o
o
o
p
e
h
s
m
r
l
m
D
n
o
t
e
p
s
s
s
s
u
t
l
/
o
t
n
n
c
o
n
e
e
e
e
a
o
i
i
t
i
y
r
r
r
r
a
a
C
l
r
h
h
o
m
o
o
o
o
i
I
a
e
.
m
p
m
e
i
x
p
h
I I V
h
h
h
h
t
s
f
l
2
I I I
m
e
a
t
p
r
c
p
p
p
p
a
l
u
n
t
r
a
a
t
a
a
i
i
l
s s s
l
e
o
o
o
o
p
g
i
l
r
l
t
t
t
t
F
u
d
a
l
e
o
n
u
a
t
m m m
i
b
o
o
o
o
d
b
a
m
z
r r r
e
e
c
l
h
o a
a
h
h
h
h
i
u
o
A
Ag Ag Ad
T7 T)
T
P,
P
P8
Pe
Ga
Ge
Oy
c
C
Go
Sm
S
* AROCHA: NEW SPECIES OF LYCOTEUTHID SQUID 951
Leptodontoteuthis inermis belongs in the synonymy of L. diadema. Oregoniateuthis springeri Voss, 1956 characterized by having elongate dorso-lateral arms and an elongate tail, was found to be the sexually dimorphic male of L. diadema (Toll, 1983). Toll (1983) determined that the species described by Steenstrup in 1875 as Onychoteuthis loligera was a male with the basic proportions of a male L. diadema. The type specimen (180 mm–ML), which was in poor condition (Voss, 1962), had as type locality the ‘South Sea’. Forch and Uozumi (1990) redescribed L. lorigera from a single male and found that it lacked the long tail characteristic of L. diadema, and the females collected were indis- tinguishable from L. diadema. Later, in a cephalopods study of the cephalopods of the Benguela Current off Namibia, Villanueva and Sanchez (1993) examined a number of L. lorigera of both sexes, all of which were mature specimens, and argued that L. lorigera males are con-specific with L. diadema females, based on indirect biological evidence in which L. lorigera males are found with L. diadema females in different areas of the southern hemisphere: off New Zealand (Förch and Uozumi, 1990) and in the Benguela Current off Namibia. Based on geography, L. diadema is assumed to be a synonym of L. lorigera, and the North Atlantic ‘L. diadema’ is L. springeri. Although, no specimens of L. lorigera were examined in the present study, the author is inclined to believe like Nesis (1987), Villanueva and Sanchez (1993) and Vecchione and Young (1999) that L. lorigera is the species described by Chun (1910) as Thaumatolampas diadema (= L. diadema), and that L. springeri is the species with the mantle drawn out into a short tail in females and a long tail in males, bearing a single large photophore in the musculature at the apex of the fins in females, and eight slender, rodlike photophores in the tail of males (Voss, 1956, 1962; Toll, 1983). Both species of Lycoteuthis can be differentiated from other members of the family by the long dorso-lateral arms in males. Females cannot be easily differentiated because of the presence of a light organ at the end of mantle in L. springeri. Because females of Nematolampas are not known, the separation of Lycoteuthis females from other members of the family remains unclear. Selenoteuthis scintillans Voss, 1958 is well separated from the other species by the presence of a large globular photophore at the posterior end of mantle in both sexes and three photophores on the tentacles (Herring et al., 1985). Male specimens also can be separated by the absence of long lateral arms and the presence of a single photophore at tips of arms II and III. The high level of sexual dimorphism in the family has been noted by several authors (Voss, 1962; Toll, 1983; Nesis, 1987; Villanueva and Sanchez, 1993). Examination of male specimens in five of the six species in the family and study of the literature showed that a hectocotylus is present in the single species of the subfamily Lampadioteuthidae, and absent in the five species of the subfamily Lycoteuthinae. Within the Lycoteuthinae, the males of Lycoteuthis have very long dorso-lateral arms and the males of Nematolampas have very long ventro-lateral arms, both genera with modified distal portions, which may be called ‘secondary sexual modification’ [perhaps of use in caressing or holding the females, as proposed by Voss (1980) in the Cranchiidae]. In contrast, the only known species of Selenoteuthis does not have elongated lateral arms. The Lycoteuthinae genera also differ in the photophore pattern on the lateral arms. In Selenoteuthis, a single photo- phore is present on the tip of each arm II and III, while in Lycoteuthis and Nematolampas several photophores are present in dorso- and ventro-lateral arms. In addition, males of 952 BULLETIN OF MARINE SCIENCE, VOL. 72, NO. 3, 2003
Lycoteuthis and Selenoteuthis have paired genitalia, in contrast to Nematolampas, which have single male genitalia. Other features that heighten the level of sexual dimorphism in the family are the loca- tions, arrangements and sizes of the photophores on the arms and body. As these squids are mesopelagic to benthic-bathyal (Nesis, 1987), one of the functions of these photo- phores may be to differentiate sexes (and probably species) at time of courtship. Herring et al. (1985) suggested the tail photophores may be used as specific sexual signaling organs. The present study enhances our knowledge of the Lycoteuthidae with the addition of the new species to the family and reviews the characters that distinguish the species within the family. A full review of the family is needed to understand the position of the Lycoteuthidae in the teuthoid classification, but must await additional material of both sexes, in particular mature specimens.
ACKNOWLEDGEMENTS
The author expresses his gratitude to N. Voss for the encouragement, advise and support during the development of this study. Her reviews provided important suggestions that enhanced the manu- script. I also wish to thank C. F. E. Roper, M. Vecchione, and R. Toll for their comments and suggestions in earlier versions of the manuscript. Special thanks are given to L. Marcano and R. Cipriani for having a good eye for picking up odd-looking squids, and the drawing skills of R. Cipriani in the full specimen. Special thanks are also given to M. Sweeney for lending the type material deposited at the National Museum of Natural History and to N. Voss for providing space and the cephalopod collections during my visit to the Marine Invertebrate Museum-Rosenstiel School of Marine and Atmospheric Science. Special thanks are given to R. E. Young, M. Vecchione and N. Voss for their constructive criticisms of the final manuscript.
LITERATURE CITED
Anonymous. 1989. Surveys of the fish resources in the shelf areas between Suriname and Colom- bia. Reports on surveys with the R/V DR. F. NANSEN. NORAD/UNDP/FAO Programme. Inst. Marine Research, Bergen. 135 p. Arocha, F. 1991. A new species of Nematolampas (Cephalopoda: Lycoteuthidae) from the central western Atlantic. [Abstract] Bull. Mar. Sci. 49: 660 p. Berry, S. S. 1913. Nematolampas, a remarkable new cephalopod from the South Pacific. Biol. Bull. 25: 208–212. ______. 1914. Notes on a collection of cephalopods from the Kermadec Islands. Trans. N.Z. Inst. 46: 134–149. ______. 1916. Cephalopoda from the Kermadec Islands. Proc. Acad. Natl. Sci. Phil. 1916: 45– 66. Chun, C. 1900. Aus den Tiefen des Weltmeers. Gustav Fischer, Jena, pp i-viii+1–549, 390 figs., 46 pls. ______. 1910. Die Cephalopoden. 1. Teil:Oegopsida. Wiss. Ergebn. “Valdivia,” 18: 1–552, atlas of 61 pls. Förch, E. C. and Y. Uozumi. 1990. Discovery of a specimen of Lycoteuthis lorigera (Steenstrup, 1875) (Cephalopoda: Teuthoidea) from New Zealand and additional notes on its morphology. New Zealend J. of Mar. and Freshwater Res., 24: 51–258. Herring, P. J., P. N. Dilly and C. Cope. 1985. The photophore morphology of Selenoteuthis scintillans Voss and other lycoteuthids (Cephalopoda: Lycoteuthidae). J. Zool., Lond., (A) 206: 567–589. AROCHA: NEW SPECIES OF LYCOTEUTHID SQUID 953
Hess, S. C. 1987. Comparative morphology, variability, and systematic applications of cephalopod spermatophores (Teuthoidea and Vamyromorpha). Ph.D. Diss., University of Miami, Miami. 597 p. Nesis, K. 1987. Cephalopods of the world. Squids, cuttlefishes, octopuses, and allies. T.F.H. Publi- cations, Inc., Neptune City, New Jersey, 351 p. Pfeffer, G. 1912. Die Cephalopoden des Plankton Expedition. Ergebnisse Plankton Expedition der Humboldt-Stiftung 2:1–815, atlas of 48 pls. Roper, C. F. E. and G. L. Voss. 1983. Guidelines for taxonomic descriptions of cephalopod species. Mem. Nat. Mus. Victoria 44: 49–63. Toll, R. 1983. The lycoteuthid genus Oregoniateuthis Voss, 1956, a synonym of Lycoteuthis Pfeffer, 1900 (Cephalopoda: Teuthoidea). Proc. Biol. Soc. Wash. 96: 365–369. Vecchione, M. and R. E. Young. 1999. Lycoteuthidae. http://phylogeny.arizona.edu/tree/eukary- otes/animals/mollusca/cephalopoda/coleoidea/decapodiformes/enoploteuthid-families/ lycoteuthidae/lycoteuthis/lycoteuthis.html. Villanueva, R. and P. Sanchez. 1993. Cephalopods of the Benguela Current off Namibia: new addi- tions and considerations on the genus Lycoteuthis. J. Nat. History 27: 15–46. Voss, G. L. 1956. A review of the cephalopods of the Gulf of Mexico. Bull. Mar. Sci. Gulf & Carib. 6: 85–178. ______. 1958. The cephalopods collected by the R/V ATLANTIS during the West Indian Cruise of 1954. Bull. Mar. Sci. Gulf and Carib. 8: 369–389. ______. 1962. A monograph of the Cephalopoda of the North Atlantic I. The family Lycoteuthidae. Bull. Mar. Sci. Gulf and Carib. 12: 264–305. ______. 1980. A generic revision of the Cranchiidae (Cephalopoda:Oegopsida). Bull. Mar. Sci. 30: 365–412. Young, R. E. 1964. A note on three specimens of the squid Lampadioteuthis megaleia Berry, 1916 (Cephalopoda:Oegopsida) from the Atlantic Ocean, with a description of the male. Bull. Mar. Sci. 14: 444–452.
DATE SUBMITTED: June 8, 2001. DATE ACCEPTED: August 26, 2002.
ADDRESS: Instituto Oceanográfico de Venezuela, Universidad de Oriente, Apartado de Correos No. 204, Cumaná 6101, Venezuela. E-mail: