Grimpoteuthis, ROBSON, 1932

Collins, Martin A., 2003, The genus Grimpoteuthis (Octopoda: Grimpoteuthidae) in the north-east Atlantic, with descriptions of three new species, Zoological Journal of the Linnean Society 139 (1), pp. 93-127 : 97-122

GENUS GRIMPOTEUTHIS ROBSON, 1932

New diagnosis

Small to large octopods with semigelatinous body of bell-shaped form. Thick primary web, intermediate (secondary) web absent. White (areolar) spots absent. Fins medium to large, lateral, with distinct lobe near the anterior fin insertion. Eyes lateral. Gills halforange in form. Optic lobe spherical; optic nerve passes through white body as a single bundle of fibres. Shell vestige U-shaped; lateral sides parallel, not tapered to fine points. Radula homodont or absent. Posterior salivary glands small or absent. Digestive gland entire (single lobe). Sucker sexual dimorphism present in some species, but with single enlarged field. Type species Cirroteuthis umbellata by subsequent designation.

Etymology

Named in honour of the eminent cephalopod researcher Georg von Grimpe.

GRIMPOTEUTHIS UMBELLATA ( FISCHER, 1883)

( FIGS 2, 17 ; TABLES 2, 11)

Cirroteuthis umbellata Fischer, 1883: 404 (in part) Cirroteuthis umbellata Fischer & Joubin, 1907: 318 Grimpoteuthis umbellata ( Robson, 1932: 137) Grimpoteuthis umbellata ( O’Shea, 1999: 45) Grimpoteuthis umbellata (Voss, 1988, in part) Grimpoteuthis umbellata ( Sweeney & Roper, 1998) Non Stauroteuthis umbellata ( Ebersbach, 1915: text fig. 1)

Non Cirroteuthis umbellata ( Massy, 1909: 4)

Non Cirroteuthis umbellata ( Joubin, 1920: 13)

Non Cirroteuthis umbellata ( Chun, 1913: 16)

Material examined

LECTOTYPE: MNHN 3.6.698: immature female, 25 mm ML; north-east Atlantic, Azores , between Fayal and San Miguel; 37∞55¢N, 20∞22¢W; 2235 m; Talisman 130; 16 August 1883.

Type locality

North-east Atlantic, Azores, between San Miguel and Fayal, 2235 m (37∞55¢N, 20∞22¢W)

Distribution

Known only from type locality.

Remarks

Known, with certainty, only from the type material, which is in poor condition, with the shell and reproductive system missing ( Fig. 2). The species was first described by Fischer (1883), from three specimens caught at different depths and locations in the northeast Atlantic. A slightly more detailed description was subsequently published by Fischer & Joubin (1907). Despite a thorough search, only two of the specimens were traced in this study. Measurements of them are given in Table 2, with additional data from unpublished notes of the late Gilbert Voss. Voss & Pearcy (1990) suggested that the smaller syntype may be a different species and, although in poor condition, it is clearly an Opisthoteuthis , on the basis of body shape and sucker form. The larger specimen is herein designated the lectotype and the smaller specimen removed from the type series. The lectotype was damaged upon capture and has subsequently been extensively dissected. Many specimens initially assigned to this species ( Table 1) are unlikely to be G. umbellata , given the limited bathymetric distributions of individual species (see below). Despite a thorough search, no well-preserved specimens have been found from the type location. The only specimen from near the location is in poor condition (see Indeterminate Material below). It is possible that G. wuelkeri (see below) is a junior synonym of G. umbellata , but until new material is obtained from the type locality this will not be possible to evaluate. G. wuelkeri has similar sucker counts, but differs from G. umbellata in the gill lamellae count.

GRIMPOTEUTHIS WUELKERI ( GRIMPE, 1920)

( FIGS 3-7 , 17 ; TABLES 3, 4, 11)

Synonymy

Stauroteuthis wülkeri Grimpe, 1920: 235

Cirroteuthis umbellata ( Chun, 1913: 16; in part) Stauroteuthis umbellata ( Ebersbach, 1915: text fig. 1) Grimpoteuthis wülkeri ( Robson, 1932: 152) Grimpoteuthis wuelkeri nomen dubium ( Voss, 1988a: 296)

Grimpoteuthis wülkeri ( Sweeney & Roper, 1998: 577) Enigmatiteuthis wülkeri ( O’Shea, 1999: 50) Grimpoteuthis sp. A . ( Collins et al., 2001)

Material examined

HOLOTYPE: ZUB 36372; immature female 46 mm ML; 35∞46¢N, 8∞16¢W; 2056 m; 8 May 1910; RV Michael Sars Station 25.

Other material examined

BMNH 20030368; 1 mature male; 49∞36¢N, 12∞27¢W; 1857–1910 m; OTSB; 30 May 1980; RRS Discovery 50802. BMNH 20030369; 1 mature female; 49∞36¢N, 12∞27¢W; 1857–1910 m; OTSB; 30 May 1980; RRS Discovery 50802. DC-SOC 50510 ; 1 mature female; 51∞05¢N, 13∞04¢W; 1924– 45 m; OTSB; 3 June 1979; RRS Discovery 50510. USNM 730896 ; 1 female; 36∞14¢N, 74∞30¢W; 1550–1640 m; Trawl; 11 November 1974; RV Gillis. USNM; 1 female; 57∞23¢N, 11∞29¢W; 1850–1900 m; Trawl; 16 November 1974; Walther Herwig 836/74. NMSZ 1999.4128 ; 1 female; 56∞41¢N, bars = 25 mm (A,B,F); 10 mm (C- E) .

09∞47¢W; 1775–1835 m; OTSB; 21 September 1993; RRS Challenger.

Diagnosis

Large species. Cirri short; 60–70 suckers deeply embedded in each of the arms, with suckers of MSD present over broad range of arms. Radula present, all teeth homodont; posterior salivary glands small. Shell robust, with thickened basal portion and fin attachment area strengthened; lateral wings terminate in two lobes, one of which extends to a fine point. Gills large, with 6–7 broad lamellae.

Description

Large species, body semigelatinous, bell-shaped ( Figs 3A , 4A,B ). Eyes and fins lateral. Mantle length 30% TL, and broadly rounded posteriorly. Head slightly wider than mantle, with no discernible constriction between head and mantle. Eyes of moderate size ( EDI 34). Pallial aperture small, closely surrounds funnel. Funnel moderately long (FuLI 35). Funnel organ V-shaped. Olfactory organs rounded, prominent; located just within mantle aperture, on either side of funnel. Fins large (mean FSI 69.6; mean FLI 96.6), positioned laterally with narrow muscular base, muscular area extending two-thirds of the fin length, tips rounded with slight point, anterior margin convex with distinct lobe.

Arms long (mean ALI 72.0), moderately stout, subequal in length, with arm formulae typically I.II.III.IV ( Fig. 4C ). Arms enveloped in single web, extending distally approximately 2/3 of arm length. Web extends distally further on the dorsal arms; formula A.B.C.D.E. Single web-supporting nodule present on the ventral side of each arm, usually located at level of sucker 28 on arm I, 26 on arm II, 24 on arm III and 22 on arm IV. Single row of suckers extends from mouth to arm tip ( Fig. 4D ). Suckers of moderate size (mean SDI 4.8), deeply set in semigelatinous tissue of all arms ( Fig. 4E ). Sucker count similar on all arms, range of 60–70. Suckers increase in size from mouth to 8th, then remain of similar size to approximately sucker 25, then decrease to arm tips. Cirri short (mean CLI 6.1 in females; 10.0 in single male), first appear between suckers 5 and 6 on most arms (occasionally between 4 and 5 or 6 and 7). Cirri extend to tips of arms; are longest on mid-portion of 1st arm.

Gills of half-orange (spherical) form ( Fig. 5A,B ), with 6-7 broad lamellae (6/ 6 in two specimens; 6/ 7 in two and 7/ 7 in three). Optic lobes slightly larger than semicircular brain, spherical, connected to the eye by single nerve bundle that passes through white body ( Fig. 5E ), which is a pink/grey colour in preserved specimens.

Entire outer surface of digestive tract deeply pigmented ( Fig. 5C ). Upper beak without special ridges, grooves or thickenings. Lower beak with rounded hood, raised above crest; broad wings with distinct ridge running diagonally across; lateral walls without ridge or fold ( Fig. 5D ). Radula ( Fig. 6A,B ) poorly developed with all teeth homodont. Anterior salivary glands small, attached to the base of the buccal mass, posterior salivary glands small, located posterior to brain and on the ventral surface of the oesophagus. Oesophagus twice length of intestine; crop absent. Caecum with single turn, connected to single-lobed digestive gland by two digestive gland ducts.

Fin-supporting cartilage (shell) robust, thickened U-shape, lateral wings with parallel outer edges ( Fig. 5F ). Basal portion with outer and inner surfaces convex and with distinct ridge on the outer surface (not visible in illustration). Lateral wings terminate in two lobes, one of which has a small point.

Male reproductive tract ( Fig. 7A ) composed of large, oval testis located in median dorsal portion of mantle; vas deferens short; seminal vesicle complex large, convoluted; accessory gland complex consisting of three glands, with the distal gland largest and with a shortterminal organ (penis) projecting from it.

Female reproductive tract ( Fig. 7B ) simple, unpaired. Oviducal gland in two parts; proximal part cream in colour; distal section green/brown. Large eggs (12–13 mm in length) were found in the ovary of three specimens. Egg from oviducal gland of one specimen ( BMNH 20030369 ) 14.0 mm in longest dimension .

Skin surface smooth, unpigmented on the oral and aboral surface in specimens preserved in Steadman’s solution, except for the posterior margin of fins, which are purple.

Type locality

Off the coast of Morocco (35∞46¢N; 8∞16¢W), 2056 m.

Distribution

Known from depths of 1550–2056 m in the North Atlantic. One specimen from the north-west Atlantic, all others from north-east Atlantic.

Etymology

Named in honour of Gerhard Wülker.

Remarks

The type specimen was originally identified as C. umbellata by Chun (1913) and was the undissected specimen illustrated by Ebersbach (1915). Grimpe (1920) examined the Michael Sars material and designated the specimen as the holotype of his new species, Stauroteuthis wülkeri . The specimen from the north-west Atlantic ( USNM 730896) was caught geographically and bathymetrically near to the type location of G. plena ( Verrill, 1885) . The type and only specimen of G. plena (see Tables 1, 12) is in poor condition and difficult to evaluate. Indices derived from Verrill’s data and illustration indicate close similarity with the material described here, but fewer suckers were present on the G. plena specimen (55).

GRIMPOTEUTHIS BOYLEI SP. NOV.

( FIGS 3 , 6 , 8-10 , 17 ; TABLES 5, 6, 11)

Synonomy

Grimpoteuthis sp. B ( Collins et al., 2001).

Material examined

HOLOTYPE: BMNH 20030370; 1 male 90 mm ML; 48∞47¢N, 16∞30¢W; OTSB; 4845–4847 m; 3 September 1989; RRS Discovery 11908#35.

PARATYPES: BMNH 20030371; 1 female 115 mm ML; 49∞35¢N, 14∞01¢W; OTSB; 4190–4255 m; 20 February 1982; RRS Discovery 51309. BMNH 20030372; 1 male 66 mm ML; 48∞27¢N, 15∞40¢W; OTSB; 4843– 4840 m; 2 May 1999; RRS Discovery 54902#1. BMNH 20030373; male 65 mm ML; 48∞53¢N, 16∞40¢W; OTSB; 4848– 4847 m; 10 July 1997; Discovery 13200#27. BMNH 20030374; male 63 mm ML; 48∞53¢N, 16∞40¢W; OTSB; 4848– 4847 m; 10 July 1997; Discovery 13200#27. USNM 1013613 ; male 35 mm ML; 48∞48¢N, 16∞16¢W; OTSB; 4836–4839 m; 12 September 1996; Discovery 12930#64. DC-SOC; immature female 43 mm ML; 48∞50¢N, 16∞29¢W; OTSB; 4840 m; 18 March 1998; Discovery 13368#51 .

Other material examined

Discovery 12600#23; 2 males, 1 female; 20∞53¢N, 31∞14¢W; OTSB; 4480–4565 m; 4 August 1993 (in BMNH Collections).

Diagnosis

Large species. Cirri long and suckers (52–58) large, cylindrical with MSD at web margin. Gills large with 7-8 broad lamellae. Radula present, teeth homodont; posterior salivary glands present. Fin-supporting cartilage (shell) U-shaped, smooth, with distinct depression in the posterior surface of the basal portion; outer edges of lateral walls parallel, with blunt termination.

Description

Large species; body semigelatinous, bell-shaped ( Figs 3B , 8A, B ). Eyes and fins lateral. Mantle 20% TL, broadly rounded posteriorly. Head slightly wider than mantle, with no discernible constriction between head and mantle. Eyes of moderate size (mean EDI 33.9). Pallial aperture small, closely surrounds funnel. Funnel moderately long (mean FuLI 46.7). Funnel organ V-shaped. Olfactory organs ( Fig. 9A ) rounded, prominent, located just within mantle aperture on either side of funnel. Fins long (mean FSI 55.4; mean FLI 94.6), straight, tips rounded with slight point; anterior margin convex with distinct lobe. Fins with narrow muscular base, muscular area extending two-thirds of the fin length.

Arms long (mean ALI 75.5), moderately stout, subequal in length, arm formula typically I.II.III.IV. Arms enveloped in single web, extends distally 2/3 of arm length ( Fig. 8C ). Intermediate web absent. Web extends further on the dorsal arms; web formula typically A.B.C.D.E. Single web-supporting nodule prominent on the ventral side of each arm, usually located at the level of sucker 31 on arm I, 28 on arm II; 25 on arm III and 24 on arm IV. Single row of suckers extends from mouth to arm tip ( Fig. 8D ). Suckers broad (mean SDI 6.1), cylindrical with diameter greater than sucker height ( Fig. 8E ). Sucker count (52–58), approximately equal on all arms. Suckers increase in size from mouth to the web margin ( MSD between suckers 25 and 35), then decline in size to arm tip. In some specimens there is a sharp decline in sucker diameter following sucker 35. Cirri long (mean CLI 12.2), first appear before the 5th, 6th or 7th sucker. Cirri extend to arm tips, longest on mid-portion of dorsal arms.

Gills large, of half-orange (spherical) form ( Fig. 9A ), with 7-8 broad lamellae (8/8 on six specimens; 8/7 on two and 7/7 on one). Optic lobes large, spherical, single bundle of optic nerves extend through the white body to each eye ( Fig. 9C ). White body, closely associated with eye, large, pale grey in preserved specimens.

Entire outer surface of digestive tract deeply pigmented ( Fig. 9B ). Upper beak without special ridges or folds. Lower beak with rounded hood, raised above crest; broad wings with distinct flexure running diagonally across; lateral walls without ridge or fold ( Fig. 9D ). Radula poorly developed, all teeth homodont ( Fig. 6C,D ). Anterior salivary glands attached to the base of the buccal mass, posterior salivary glands small, connect into single duct, which passes through the brain to the buccal mass. Oesophagus twice length of intestine; stomach small. Caecum with single turn, connected to single-lobed digestive gland by two digestive ducts. Intestine straight.

Fin supporting cartilage (shell) simple, broad, Ushape; smooth with distinct depression on the outer surface of the basal portion ( Fig. 9E ). Inner surface of basal portion convex. Lateral wings of shell with blunt termination.

Male reproductive tract ( Fig. 10A ) composed of large, oval testis located in the posterior median portion of mantle. Vas deferens short, seminal vesicle complex convoluted; accessory gland complex consisting of three glands. Terminal organ (penis) long, projecting from distal accessory gland.

Female reproductive tract ( Fig. 10B ) simple, unpaired. Oviducal gland large, in two parts, proximal third cream in colour, distal two-thirds green/ brown. Eggs large, capsules of mature ovarian eggs 18–20 mm in length ( BMNH 20030371).

Skin surface smooth, deep red in fresh specimens, becoming reddish-brown with preservation. Pigmentation greatest on the oral and aboral surfaces of the arms and less on the mantle. Posterior margin of fin pigmented (purple).

Type locality

Porcupine Abyssal Plain, 48∞47¢N, 16∞30¢W, 4845–4847 m.

Distribution

Abyssal species, currently known from the type locality (Porcupine Abyssal Plain), from the Porcupine Seabight and the Madeira Abyssal Plain ( Fig. 17 ). Probably distributed at abyssal depths from 20∞N to 50∞N in the north-east Atlantic.

Remarks

This species is similar to G. challengeri (see below), but can be distinguished by the shape of the shell, presence of posterior salivary glands, form of the gills, egg size and the number, form and arrangement of suckers. Stomach contents of single specimen ( USNM 1013613) examined included remains of polychaetes (2–3 individuals), calanoid copepods, amphipods ( Rhachotropis and Octorama) and isopods of the family Munnopsidae .

Etymology

Named in honour of Professor Peter Boyle in recognition of his contributions to cephalopod biology.

GRIMPOTEUTHIS CHALLENGERI SP. NOV.

( FIGS 3 , 6 , 11-13 , 17 ; TABLES 7, 8, 11)

Synonomy

Grimpoteuthis sp. C . ( Collins et al., 2001).

Material examined

HOLOTYPE: BMNH 20030375; male 58 mm ML; 48∞56¢N, 15∞45¢W; OTSB; 4828–4838 m; 22 March 1998; RRS Discovery 13369#1.

PARATYPES: USNM 1013614 ; male 75 mm ML; 48∞47¢N, 16∞25¢W; OTSB; 4853–4877 m; 12 September 1989; Discovery 11908#68. BMNH 20030376; male 55 mm ML; 48∞27¢N, 15∞40¢W; OTSB; 4843–4845 m; 2 May 1999; Discovery 54902#1. BMNH 20030377; female 70 mm ML; 48∞48¢N, 16∞30¢W; OTSB; 4843– 4849 m; 5 September 1989; Discovery 11908#44 . USNM 1013615 ; immature female 35 mm ML; 48∞53¢N, 16∞31¢W; OTSB; 4836–4840 m; 16 September 1996; Discovery 12930#77 .

Other material examined

Discovery 13369#2; male; 48∞55¢N, 15∞40¢W; OTSB; 4832–4836 m; 23 March 1998 ( DC-SOC). Discovery 54901#2; male; 48∞42¢N, 16∞52¢W; OTSB; 4811– 4837 m; 26 April 1999 ( DC-SOC). Discovery 12930#64; female; 48∞48¢N, 16∞16¢W; OTSB; 4836–4839 m; 12 September 1996 ( DC-SOC). UMML31.2890 ; female; RV Gillis St. 144; no record of locality .

Diagnosis

Large species. Cirri long and suckers (63–72) moderate with MSD at web margin. Gills with eight narrow lamellae. Fin-supporting cartilage (shell) robust U-shape; lateral wings well developed, each terminating in two lobes. Radula present; posterior salivary glands absent.

Description

Large species; body semigelatinous, bell-shaped ( Figs 3C , 11A,B ). Mantle 22% TL, and broadly rounded posteriorly. Head slightly wider than mantle, no discernible constriction between head and mantle. Eyes of moderate size (mean EDI 32.7). Pallial aperture small, closely surrounding funnel. Funnel, moderately long (mean FuLI 50.7). Funnel organ Vshaped. Olfactory organs rounded, prominent, located just within mantle aperture on either side of funnel. Fins very long (mean FSI 59.0; mean FLI 113.8), tips rounded with slight point, anterior margin convex with distinct lobe. Fins with narrow muscular base, muscular area extending two-thirds of the fin length.

Arms long (mean ALI 74.3), moderately stout, subequal in length, with arm formulae typically I.II.III.IV. Web single, occupying approximately 2/3 of arm length ( Fig. 11C ). Intermediate (secondary) web absent. Web extends further on dorsal arms, and generally of the formula A.B.C.D.E. Single web-supporting nodule on the ventral side of each arm, typically located at level of sucker 34 on arm I, 30 on arm II; 28 on arm III and 27 on arm IV ( Fig. 11C,D ). Suckers in single row, count similar on all arms (range 63–72), extend from mouth to arm tip. Suckers cylindrical ( Fig. 11E,F ) with sucker diameter similar to sucker height (mean SDI 4.6), increase in size from mouth to the web margin ( MSD between suckers 30 and 38), then decline in size to the arm tip. MSD similar in males and females; but suckers deeply embedded in arms in females ( Fig. 11E,F ). Cirri long (mean CLI 13.7), first appear before the 5th, 6th or 7th sucker. Cirri extend to the tips of the arms, longest on midportion of dorsal arms.

Gills moderate, of half-orange (spherical) form ( Fig. 12A ), normally with eight narrow lamellae (8/8 on eight specimens; 8/7 on one). Optic lobes large, spherical, with a single bundle of optic nerves running through white body to each eye ( Fig. 12C ). White body, closely associated with eye, large, pale grey in preserved specimens.

Outer surface of oesophagus and stomach deeply pigmented ( Fig. 12B ). Upper beak without well defined ridges or grooves. Lower beak with rounded hood, distinctly raised above crest; broad wings with distinct flexure running diagonally across; lateral walls without ridge or fold ( Fig. 12D ). Radula present ( Fig. 6E,F ), poorly developed, all teeth homodont. Anterior salivary glands small, attached to the base of the buccal mass; posterior salivary glands absent. Oesophagus twice the length of intestine. Stomach small, simple, appears as a swelling of the oesophagus. Caecum with single turn, connected to single-lobed digestive gland by two digestive ducts.

Fin supporting cartilage (shell) robust, thickened U-shape, with the outer walls of the lateral wings parallel. Inner surface of the basal portion convex; outer surface concave. Lateral wings each terminate in two lobes ( Fig. 12E ).

Male reproductive tract ( Fig. 13A ) composed of large, oval testis located in the posterior median portion of the mantle. Vas deferens short, seminal vesicle complex convoluted. Accessory organ complex consists of three glands, with proximal gland the largest. A long-terminal organ (penis) projects from the distal accessory gland.

Female reproductive tract ( Fig. 13B ) simple, unpaired. Oviducal gland in two parts, proximal third cream in colour, distal two-thirds green/brown. Eggs of moderate size, 13 mm in length in oviduct of specimen BMNH 20030377.

Skin surface smooth, reddish-brown following preservation. Pigmentation greatest on the oral and aboral surfaces of the arms and less on the mantle. Posterior margin of fins pigmented (purple).

Type locality

Porcupine Abyssal Plain, 48∞56¢N, 15∞45¢W; 4828–4838 m.

Distribution

Abyssal species, known from the type locality, caught at depths of 4800–4850 m ( Fig. 17 ). The capture location for the single specimen in the UMML collection is not known, but is from the north-west Atlantic .

Remarks

This species is similar to Grimpoteuthis tuftsi , which Voss & Pearcy (1990) described from abyssal depths in the North Pacific, but can easily be distinguished from the other north-east Atlantic species by the narrow lamellae of the gills.

Etymology

Named after RRS Challenger, which, together with RRS Discovery, was the vessel responsible for catching many of the specimens reported in the present study.

GRIMPOTEUTHIS DISCOVERYI SP. NOV.

( FIGS 3 , 14-17 ; TABLES 9 –11)

Synonomy

Grimpoteuthis spp. D & E ( Collins et al., 2001) Cirroteuthis umbellata ( Chun, 1913: 16, in part) Cirroteuthis umbellata ( Joubin, 1920: 13) Grimpoteuthis s.sp. D ( Piertney et al., 2003)

Material examined

HOLOTYPE: BMNH 20030378; male 50 mm ML; 49∞35¢N, 14∞01¢W; 4190–4255 m; OTSB; 20 February 1982; Discovery 51309.

PARATYPES: BMNH 20030379; male 45 mm ML; 49∞35¢N, 14∞01¢W; 4190–4255 m; OTSB; 20 February 1982; Discovery 51309. USNM 1012988 ; male 42 mm ML; 49∞35¢N, 14∞01¢W; 4190–4255 m; OTSB; 20 February 1982; Discovery 51309. USNM 1012989 ; female 35 mm ML; 49∞35¢N, 14∞01¢W; 4190–4255 m; OTSB; 20 February 1982; Discovery 51309. BMNH 20030380; female 55 mm ML; 50∞00¢N, 14∞07¢W; 4180–4310 m; OTSB; 30 March 82; Discovery 51414. BMNH 20030381; female 51 mm ML; 50∞00¢N, 14∞07¢W; OTSB; 4180–4310 m; 30 March 82; Discovery 51414. UMML 31.3166 ; 1 male 36 mm ML; 1 female 46 mm ML; 50∞03¢N, 13∞51¢W; 3749–3757 m; OTSB; 12 June 1977; Discovery 9640#1 .

Other material examined

ZUB 36379; 1 male, 1 female; 34∞59¢N; 33∞01¢W; 2615 m; Michael Sars 53; 8 June 1910 . MOM 295091; female; 39∞54¢N, 20∞27¢W; 4360 m; Trawl; 18 August 1896; Princesse Alice 753. MOM 295245; male; 44∞08¢N, 10∞44¢W; 5000 m; Trawl; 19 August 1910; Princesse Alice 2994. MOM 295344; male; 45∞07¢N, 7∞06¢W; 4870 m; Trawl; 17 August 1910; Princesse Alice 2986. NMSZ 1994128.45603 ; male; 54∞42¢N, 12∞16¢W; 2875–2900 m; OTSB; 21 October 1987 ; Challenger 20/87/3. NMSZ 1999158.158 ; female; 49∞50¢N, 14∞18¢W; 4143 m; OTSB; 9 August 1997 ; Challenger 134/6. NMSZ 1999158.160 ; female; 49∞50¢N, 14∞18¢W; 4143 m; OTSB; 9 August 1997 ; Challenger 134/6. NMSZ 1999158.159 ; male; 49∞58¢N, 14∞29¢W; 4139 m; OTSB; 17 August 1997 ; Challenger 134/24. NMSZ 1999158.161 ; male; 49∞58¢N, 14∞29¢W; 4139 m; OTSB; 17 August 1997 ; Challenger 134/24. Discovery 9756#3; 1 male, 1 female, 49∞48¢N, 14∞15¢W; 4080–4156 m; OTSB; 11 April 1978. Discovery 9756#5; female; 49∞49¢N, 14∞05¢W; 4012–4020 m; OTSB; 12 May 1978. Discovery 51012; 1 male, 2 females; 49∞51¢N, 13∞58¢W; 3880–3920 m; OTSB; 5 May 1981. Discovery 50512; female; 50∞14¢N, 13∞42¢W; 3022–3110 m; OTSB; 4 June 1979. Discovery 50811; 2 males; 49∞39¢N, 14∞32¢W; 4400– 4350 m; OTSB; 2 August 1980. Discovery 50910; male; 49∞50¢N, 14∞40¢W; 4265–4320 m; OTSB; 10 November 1980. Discovery 51216#4; 1 male, 1 female; 49∞49¢N, 14∞07¢W; 4000–3970; OTSB; 29 September 1981. Discovery 51803; female; 49∞37 N, 13∞49¢W; 3990– 3920 m; OTSB; 29 September 1983. Discovery 9638#2; 1 female, 1 male; 49∞50¢N, 14∞07¢W; 4043–4104; OTSB; 8 June 1977. Discovery 9640#1; male; 50∞03¢N, 13∞51¢W; 3749–3757 m; OTSB; 12 June 1977. Discovery 13369#1; female; 48∞56¢N, 15∞45¢W; 4828–4838 m; OTSB; 22 March 1998. Discovery 13369#2; female; 48∞54¢N, 15∞40¢W; 4828–4836 m; OTSB; 23 March 1998. Discovery 50514; male; 49∞44¢N, 14∞02¢W; 4017–4095 m; OTSB; 5 June 1979. Discovery 54301#6; male; 48∞47¢N, 16∞50¢W; 4837– 4846 m; OTSB; 20 October 1997. Discovery 52403#13; 2 males; 48∞52¢N, 15∞57¢W; 4805–4810 m; OTSB; 5 December 1986. Discovery 52403#14; female; 49∞02¢N; 16∞00¢W; 4850–4860 m; BN1.5; 5 December 1986. Discovery 11908#7; male; 49∞04¢N, 16∞04¢W; 4847– 4789 m; OTSB; 24 August 1989. Discovery 11908#51; female; 48∞46 N, 16∞32¢W; 4839–4856 m; OTSB; 7 September 1989. Discovery 11908#68; 1 female, 2 males; 48∞47¢N, 16∞25¢W; 4853–4877 m; OTSB; 12 September 1989. Discovery 12930#26; male; 48∞53¢N, 16∞43¢W; 4836–4843 m; OTSB; 4 September 1996. Discovery 12930#64; female; 48∞48¢N; 16∞16¢W; 4836–4839 m; OTSB; 12 September 1996. Discovery 51608#1; male; 49∞36¢N, 14∞30¢W; 4370– 4270 m; OTSB; 19 August 1982. Discovery 54901#2; female; 48∞42¢N, 16∞52¢W; 4811–4837 m; OTSB; 26 April 99. Discovery 54902#1; female; 48∞27¢N, 15∞40¢W; 4843–4845 m; OTSB; 2 May 99. Discovery 13906#1; male; 50∞12¢N, 14∞40¢W; 3972–4003 m; OTSB; 18 September 2000. Discovery 13925; 3 males; 48∞57¢N, 16∞55¢W; 4835–4845 m; OTSB; 7 October 2000.

Diagnosis

Small to medium sized species. Gills with seven (occasionally six or eight) broad lamellae. Radula and posterior salivary glands absent. Fin-supporting cartilage (shell) U-shaped, with outer edges of lateral wings parallel, terminating in two lobes. Eggs 10–11 mm. Oral suckers (1–4) small with distinct increase in size at sucker 6. Suckers of MSD present over broad range of arms; sexually dimorphic, larger in males. Cirri of moderate length.

Description

Small to medium species; body semigelatinous, bellshaped ( Figs 3D , 14A,B ). Mantle 25% TL, and broadly rounded posteriorly. Head slightly wider than mantle, with no discernible constriction between head and mantle. Pallial aperture small, closely surrounding funnel. Funnel, moderately long (mean FuLI 45.4). Funnel organ V-shaped. Olfactory organs rounded, prominent, located just within mantle aperture on either side of funnel. Fins lateral, moderately long (mean FSI 63.9; mean FLI 104.8); posterior margin straight; tips rounded with slight point, anterior margin convex with distinct lobe. Fins with narrow muscular base, muscular area extending two-thirds of the fin length.

Arms long (mean ALI 72.5), moderately stout, subequal in length, with arm formulae generally I.II.III.IV. Arms enveloped in single web, occupying approximately 2/3 of arm length ( Fig. 14C ). Intermediate (secondary) web absent. Web extends further on dorsal arms, generally of formula A.B.C.D.E. Single, prominent, web supporting nodule on the ventral side of each arm, located at approximately sucker 31 on arm I, 29 on arm II; 27 on arm III and 25 on arm IV ( Fig. 14C,D ). Suckers in single row from mouth to arm tip, approximately equal numbers on all arms (range 56–61). Suckers barrel-shaped and show distinct sexual dimorphism ( Fig. 14E,F ). In mature females sucker height approximately equal to SD (mean SDI 4.4). In mature males sucker height (distance from sucker base to tip) greatly exceeds SD. SD is significantly greater in males (mean SDI 6.4) than females. In both sexes the first 4 or 5 (oral) suckers are very small, and there is a distinct increase in SD at sucker 5 or 6; this increase is greatest in mature males. From sucker 6 to approximately sucker 40 the sucker size is constant. Terminal 15–20 suckers decrease rapidly in size. Cirri moderate (mean CLI 7.3), first appear before the 4th or 5th sucker. Cirri extend to the tips of the arms, longest on mid-portion of dorsal arms.

Gills large, of half-orange (spherical) form ( Fig. 15A ), normally with seven broad lamellae (7/7 on 29 specimens; 8/7 on 10; 8/8 on four, 7/6 on 2 and 6/6 on one). Eyes of moderate size (mean EDI 32.5). Optic lobes large and spherical, with a single bundle of optic nerves running through the white body to each eye ( Fig. 15D ). White body of characteristic shape, closely associated with eye, large, dark grey in preserved specimens.

Entire outer surface of digestive tract deeply pigmented ( Fig. 15B ). Upper beak lacking distinct ridges or grooves. Lower beak with rounded hood, slightly raised above crest; broad wings with distinct ridge running diagonally across; lateral walls without ridge or fold ( Fig. 15C ). Radula absent. Anterior salivary glands attached to the base of the buccal mass, posterior salivary glands absent. Oesophagus twice the length of the intestine; stomach small, a simple swelling of the distal oesophagus. Caecum with single turn, connected to single-lobed digestive gland by two digestive ducts.

Fin-supporting cartilage (shell) robust, thickened U-shape ( Fig. 15E ). Outer edge of lateral wings parallel, terminating in two lobes. Inner and outer surfaces of the basal portion convex.

Male reproductive tract ( Fig. 16A ) with large, oval testis located in the posterior median portion of the mantle. Vas deferens short; seminal vesicle complex convoluted; accessory gland complex comprising three glands; terminal organ (penis) of moderate length, projecting from distal accessory gland.

Female reproductive tract simple, unpaired ( Fig. 16B ). Oviducal gland in two parts, proximal third cream in colour, distal two-thirds either green or redbrown, depending on preservative. Oviducal eggs of maximum length 10–11 mm.

Skin surface smooth, oral and aboral surfaces of arms deeply pigmented (purple). Mantle with less pigmentation. Posterior margin of fins pigmented (purple).

Type locality

Porcupine Seabight; 49∞35¢N, 14∞01¢W; 4190–4255 m.

Distribution

Caught at depths of 2600–4870 m in the north-east Atlantic ( Fig. 17 ).

Remarks

Grimpoteuthis discoveryi is a variable form occupying a broad depth range and Collins et al. (2001) thought it may be two distinct species. Variability was noted in the cirrus length and shell form. Cirri length varied from 1.5 to 5.0 mm ( CLI 3.6–13.8), but this may be due to fixation. The form of the shell varied, but this may be attributable to sexual dimorphism and size (maturity). It is possible that the material examined represents more than one species but further detailed morphological, and perhaps molecular, work is required. In view of this the type series was selected from close proximity (50∞N, 40∞W) at a similar depth (~ 4000 m). The size of the suckers is highly dimorphic in mature animals, with males possessing large barrel-shaped suckers. The enlargement of suckers differs from that found in Opisthoteuthis in that there is only one, extended field of enlargement.

Etymology

Named after RRS Discovery, which was responsible for collecting most of the specimens.

Figures

Figure 3. Photographs of Grimpoteuthis wuelkeri (A), G. boylei sp. nov. (B), G. challengeri sp. nov. (C) and G. discoveryi sp. nov (D). Scale bars = 100 mm (A-C); 40 mm (D).

Figure 4. Grimpoteuthis wuelkeri: external anatomy. Ventral (A), dorsal (B) views of the holotype (ZUB 36372); oral view, orientated with dorsal arms up (C: BMNH???? (BMNH 20030369)); oral surface of first arm (D: holotype ZUB 36372), with detail of suckers and cirri (E: BMNH 20030369). Scale bars = 50 mm (A-C); 25 mm (D) and 5 mm (E).

Figure 5. Grimpoteuthis wuelkeri: internal anatomy. Ventral view of viscera of the holotype (A: ZUB 36372) and mature female (B: BMNH 20030369); digestive system (C); lower and upper beaks (D) optic nerve, white body and posterior salivary glands (E: all BMNH 20030369); dorsal ventral and lateral views of the shell of the holotype (F: ZUB 36372). Scales

Figure 6. Scanning electron micrographs of the radulae of Grimpoteuthis wuelkeri (A,B), G. boylei sp. nov. (C,D) and G. challengeri sp. nov. (E,F).

Figure 7. Grimpoteuthis wuelkeri: internal anatomy. Dorsal and ventral view of the male reproductive system (A: BMNH 20030368); ventral view of female reproductive system (B: DC-SOC: Discovery 50510). Scale bars = 15 mm (A); 25 mm (B).

Figure 8. Grimpoteuthis boylei sp. nov: external anatomy. Dorsal (A), ventral (B) and oral (C) views (all BMNH 20030371); oral view of first arm of male (D) with detail of suckers (E: both BMNH 20030370). Scale bars = 100 mm (A,B,C,D); 10 mm (E).

Figure 9. Grimpoteuthis boylei sp. nov: internal anatomy. Ventral view of the viscera in situ (A: BMNH 20030371); the digestive system (B); optic nerve and white body (C); lower and upper beaks (D); dorsal, ventral and lateral view of shell (E: all BMNH 20030374). Scale bars = 20 mm (A,B,E); 10 mm (C,D).

Figure 10. Grimpoteuthis boylei sp. nov: internal anatomy. Dorsal and ventral views of the male reproductive system (A: BMNH 20030373). Ventral view of the female reproductive system (B: BMNH 20030371). Scale bars = 15 mm (A); 30 mm (B).

Figure 11. Grimpoteuthis challengeri sp. nov: external anatomy. Dorsal (A), ventral (B) and oral (C) views (all BMNH 20030375); oral view of first arm (D: BMNH 20030375), with detail of female (E: BMNH 20030377) and male suckers (F: BMNH 20030375). Scale bars = 50 mm (A,B,D); 100 mm (C); 5 mm (E,F).

Figure 12. Grimpoteuthis challengeri sp. nov: internal anatomy. Ventral view of the viscera in situ (A); detail of the digestive system (B); optic nerve and white body (C) lower and upper beaks (D); dorsal, ventral and lateral view of shell (E: all BMNH 20030376). Scale bars = 15 mm (A); 20 mm (B,E); 10 mm (C,D).

Figure 13. Grimpoteuthis challengeri sp. nov: internal anatomy. Dorsal and ventral views of the male reproductive system (BMNH 20030376) (A). Ventral view of the female reproductive system (BMNH 20030377) (B). Scale bars = 10 mm.

Figure 14. Grimpoteuthis discoveryi sp. nov: external anatomy. Dorsal (A), ventral (B) and oral (C: all BMNH 20030380) views; oral view of first arm (D), with detail of female (E: BMNH 20030380) and male (F: BMNH 20030378) suckers and cirri. Scale bars = 50 mm (A-C); 25 mm (D); 5 mm (E,F).

Figure 15. Grimpoteuthis discoveryi sp. nov: internal anatomy. Ventral view of the viscera in situ (A: BMNH 20030380): note pallial adductor has been cut and is not illustrated; detail of the digestive system (B); lower and upper beaks (C); optic nerve and white body (D: all BMNH 20030381); dorsal, ventral and lateral view of shell (E: BMNH 20030380). Scale bars = 20 mm (A); 10 mm (B-D); 15 mm (E).

Figure 16. Grimpoteuthis discoveryi sp. nov: internal anatomy. Dorsal and ventral views of the male reproductive system (A: BMNH 20030378). Ventral view of the female reproductive system (B: BMNH 20030381). Scale bars = 20 mm.

Figure 17. Distribution of the genus Grimpoteuthis in the North Atlantic. Grimpoteuthis wuelkeri (A), G. boylei sp. nov. (B), G. challengeri sp. nov. (C), G. discoveryi sp. nov. (D), G. umbellata, G. plena and G. megaptera (E).