Vampyroteuthidae

 

• Vampyromorpha (Robson, 1929)
• Vampyroteuthidae (Thiele, in Chun, 1915)
• Vampyroteuthis
• The Vampire Squid

A list of all nominal genera and species in the Vampyroteuthidae can be found below. The list includes the current status and type species of all genera, and the current status, type repository and type locality of all species and all pertinent references.

Nominal Genus-Level Taxa:

Danateuthis Joubin, 1929:375.

• CURRENT SYSTEMATIC STATUS. -- Vampyroteuthis Chun, 1903 [fide Pickford (1946:7)]
• TYPE SPECIES. -- Danateuthis schmidti Joubin, 1929 by monotypy

Hansenoteuthis Joubin, 1929:388.

• CURRENT SYSTEMATIC STATUS. -- Vampyroteuthis Chun, 1903 [fide Nesis (1987:276)]
• TYPE SPECIES. -- Hansenoteuthis lucens Joubin, 1929 by monotypy

Hymenoteuthis Thiele, 1916:4.

• CURRENT SYSTEMATIC STATUS. -- Vampyroteuthis Chun, 1903 [fide Pickford (1946:7)]
• TYPE SPECIES. -- Cirroteuthis macrope Berry, 1911 by monotypy

Melanoteuthis Joubin, 1912:1, figs 1-12.

• CURRENT SYSTEMATIC STATUS. -- Vampyroteuthis Chun, 1910 [fide Pickford (1939:1)
• TYPE SPECIES. -- Melanoteuthis lucens Joubin, 1912 by monotypy

Retroteuthis Joubin, 1929:383.

• CURRENT SYSTEMATIC STATUS. -- Vampyroteuthis Chun, 1903 [fide Pickford (1946:7)]
• TYPE SPECIES. -- Retroteuthis pacifica Joubin, 1929 by monotypy

Vampyroteuthis Chun, 1903:88, text-fig.

• CURRENT SYSTEMATIC STATUS. -- Valid genus [fide Robson (1932:98)]
• TYPE SPECIES. -- Vampyroteuthis infernalis Chun, 1903 by monotypy

Watasella Sasaki, 1920:168.

• CURRENT SYSTEMATIC STATUS. -- Vampyroteuthis Chun, 1903 [fide Nesis (1987:276)]
• TYPE SPECIES. -- Watasella nigra Sasaki, 1920 by monotypy

 

Nominal Species-Level Taxa (As Introduced Binomial)

 

Melanoteuthis anderseni Joubin, 1931:170, figs 1-5.

• CURRENT SYSTEMATIC STATUS. -- Vampyroteuthis infernalis Chun, 1903 [fide Pickford (1946:7)]
• TYPE REPOSITORY. -- ZMUC Holotype [fide Kristensen and Knudsen (1983:220)] [see also Pickford (1949:124)]
• TYPE LOCALITY. -- 7°30'N, 79°19'W (Pacific Ocean)

Melanoteuthis beebei Robson, 1929:470, figs 1-8.

• CURRENT SYSTEMATIC STATUS. -- Vampyroteuthis infernalis Chun, 1903 [fide Pickford (1946:9)]
• TYPE REPOSITORY. -- BMNH Holotype 1947.5.21.1 [not 1947.7.7.3-6 fide Lipinski et al. (2000:110); incorrect catalog number, same as Helicocranchia beebei syntypes]
• TYPE LOCALITY. -- South of Coccos Island, 4°45'N, 87°00'W (Pacific Ocean)

Vampyroteuthis infernalis Chun, 1903:88, text-fig.

• CURRENT SYSTEMATIC STATUS. -- Valid species [fide Pickford (1939:48)]
• TYPE REPOSITORY. -- ZMB Holotype Moll.-63850 [fide Glaubrecht and Salcedo-Vargas (2000:275)]
• TYPE LOCALITY. -- 1°56.7'S, 7°40.6'E (Atlantic Ocean) [fide Glaubrecht and Salcedo-Vargas (2000:275)]

Hansenoteuthis lucens Joubin, 1929:388, figs 17-23.

• CURRENT SYSTEMATIC STATUS. -- Vampyroteuthis infernalis Chun, 1903 [fide Pickford (1946:7)]
• TYPE REPOSITORY. -- ZMUC Holotype [fide Kristensen and Knudsen (1983:222)] [see also Pickford (1949:123)]
• TYPE LOCALITY. -- 14°38'N, 61°16'W (Caribbean Sea)

Melanoteuthis lucens Joubin, 1912:1, figs 1-12.

• CURRENT SYSTEMATIC STATUS. -- Vampyroteuthis infernalis Chun, 1910 [fide Pickford (1939:1)
• TYPE REPOSITORY. -- MOM Holotype [station 2118] [fide Belloc (1950:2)]
• TYPE LOCALITY. -- Sargasso Sea, 31°40[']30"N, 42°44'30"W

Cirroteuthis macrope Berry, 1911:589.

• CURRENT SYSTEMATIC STATUS. -- Vampyroteuthis infernalis Chun, 1903 [fide Pickford (1946:7)]
• TYPE REPOSITORY. -- NMNH Holotype 214317 [fide Sweeney et al. (1988:15)] [see also Pickford (1949:127)]
• TYPE LOCALITY. -- 32°54'20"N, 121°11'15"W (Pacific Ocean)

Watasella nigra Sasaki, 1920:168, pl 23 fig 1.

• CURRENT SYSTEMATIC STATUS. -- Vampyroteuthis infernalis Chun, 1903 [fide Pickford (1946:9)]
• TYPE REPOSITORY. -- NMNH Holotype 332892 [fide Roper and Sweeney (1978:10)] [see also Pickford (1949:128)]
• TYPE LOCALITY. -- Off Kii Province (33°23'N, 135°37'40"E), Japan

Retroteuthis pacifica Joubin, 1929:383, figs 13-16.

• CURRENT SYSTEMATIC STATUS. -- Vampyroteuthis infernalis Chun, 1903 [fide Pickford (1946:7)]
• TYPE REPOSITORY. -- ZMUC Holotype [fide Kristensen and Knudsen (1983:223)] [see also Pickford (1949:126)]
• TYPE LOCALITY. -- 7°30'N, 79°19'W (Pacific Ocean)

Danateuthis schmidti Joubin, 1929:375, figs 7-12.

• CURRENT SYSTEMATIC STATUS. -- Vampyroteuthis infernalis Chun, 1903 [fide Pickford (1946:7)]
• TYPE REPOSITORY. -- ZMUC Holotype [fide Kristensen and Knudsen (1983:224)] [see also Pickford (1949:122)]
• TYPE LOCALITY. -- 18°50'N, 79°07'W (Caribbean Sea)

Melanoteuthis schmidti Joubin, 1929:372, figs 3-6.

• CURRENT SYSTEMATIC STATUS. -- Vampyroteuthis infernalis Chun, 1906 [fide Pickford (1946)]
• TYPE REPOSITORY. -- ZMUC Holotype [fide Kristensen and Knudsen (1983:224)] [see also Pickford (1949:126)]
• TYPE LOCALITY. -- 17°56'N, 64°50'W (Caribbean Sea)

Type Repositories

• BMNH-The Natural History Museum, Cromwell Road, London SW7 5BD, England (formerly British Museum (Natural History)). [For type catalog see Lipinski et al. (2000)]
• MOM- Musée Océanographique, Avenue Saint-Martin, Monaco-Ville, MC 98000, Monaco. [For type catalog see Belloc (1950)]
• NMNH-National Museum of Natural History, 10th & Constitution Avenue, Smithsonian Institution, Washington, DC 20560, U.S.A. (formerly USNM). [For type catalogs see Roper and Sweeney (1978); Sweeney et al. (1988)]
• ZMB- Zoologisches Museum, Museum für Naturkunde der Humboldt-Universitat, Invalidenstrasse 43, D-1040 Berlin, Germany. [For type catalog see Glaubrecht and Salcedo-Vargas (2000)]
• ZMUC-Kobenhavns Universitet, Zoologisk Museum, Universitetsparken 15, DK 2100 Copenhagen, Denmark. [For type catalog see Kristensen and Knudsen (1983); See also Volsoe et al. (1962) for Steenstrup papers]

An octopodiform ...

• with long, slender filaments (probably modified arms II).
• with cirri but without suckers on the proximal halves of arms. 

Vertical Distribution

Off California trawling data show most vampires between depths of 600-1100 m with peaks at 700-800 m and 900-1000 m, and with small individuals of less than 20 mm being most abundant at the deeper peak (Roper and Young, 1975). ROV observations in Monterey Bay, California suggest that the vampire occupies the oxygen minimum layer at depths of600-900 m with oxygen levels centered around 0.4 ml/l (Hoving and Robison, 2012).

Off Hawaii, 10 of 11 captures came from depths of 800-1200 m but little towing was done in deeper water. Two captures were from opening-closing nets at depths of about 800-950 m.

In the Atlantic at 18° N, 25° W, the vampire shows a peak distribution between 700 and 1200 m but without a clear size/depth pattern (Clarke and Lu, 1975).

Figure. Vertical distribution chart modified from Clarke and Lu, 1975.

In the figure, all captures were made with opening/closing trawls. Bars represent a capture and the bar length indicates the depth range of the trawl while open. Yellow bars indicate a daytime capture and blue bars a nighttime capture. Fishing effort between 1000-1250 m was about twice that between 1250 and 1500 m, and effort between 1000-1500 m was about 5 times that between 1500 and 2000 m and about the same as that between 500 and 1000 m.

Numerous records exists for captures in excess of 1200 m (e. g., see Roper and Young, 1975) from open nets. Unfortunately, due to the rather high probability of contamination from shallower depths, these records are of questionable value.

 

 

Horizontal Distribution

The vampire squid is broadly distributed throughout the depths of the world's tropical and temperate oceans. Some geographical variation has been noted. Young (1972) found that the beaks of vampire squid from the Pacific Ocean off California were distinctly smaller than those from vampires of the Gulf of Guinea in the Atlantic Ocean. He also noted differences in sucker size and gill size in vampires from these areas. Vampires from off Monterey, California have a predominance of reddish rather than black chromatophores.

1.  Arms
   • Arms II (probably) modified to form retractile filaments (see details below).
   • Lateral cirri present over arm length; suckers present only on distal half of arms where they alternate with cirri.
   • Suckers without cuticular lining; sucker stalks constricted distally to narrow plate at base of sucker.
     
     Figure. Ventro-oral view of V. infernalis with arms folded aborally, showing cirri without associated suckers in proximal half of arms. Note the predominance of black pigment, compared to the previous two photographs, and the colorless, oral tips of the arms. Photographed in a shipboard aquarium in the North Atlantic by David Shale.
     
     
2. Retractile filaments
   • Origin of the filaments:
     • The primitive (plesiomorphic) arrangement of arms in coleoid cephalopods is thought to be 10 equal arms. This arrangement is known in some fossil coleoids (belemnoids) and the presence of ten unequal arms in modern decapods is easily derived from such a condition. Therefore, octopods, with eight arms, have apparently lost one pair. If the vampire filaments represent modified arms II and if this is the pair that is lacking in octopods, then strong support would exist for a vampire-octopod affinity. Embryological evidence suggests that the missing arm pair in octopods is either arms II or III (Boletzky, 1978-79).
     • Pickford (e.g., 1940) concluded that the filaments were modified arms II based on their position in series with the arms, venous connections and innervation. Young (1967) clarified the innervation and argued that the filaments were not arms but homologues of the preocular tentacles of Nautilus. The axial nerves of arms send most of their fibers to the anterior subesophageal mass (brachial lobe) of the brain. Young found that the filament nerves bypass this lobe with efferent fibers arising from the middle subesophageal mass and the afferent fibers reaching an enlarged ventral magnocellular lobe. Dilly, et al. (1977), however, demonstated that large nerve bundles from the tentacles of the deep-sea squid Mastigoteuthis also pass to an enlarged ventral magnocellular lobe. Presumably the combined sensory function of thousands of suckers carried by the tentacles of Mastigoteuthis requires the large nerve bundle and the enlarged ventral magnocellular lobe. Similarly the still poorly understood functions of the filaments of Vampyroteuthis probably emphasized the connection to and size of the ventral magnocellular lobe while a loss of coordination with the principal arms resulted in a great reduction (at least) in the connection to the anterior subesophageal mass. As a result, the innervation of the filaments became a weak argument against filament-arm homology.
     • The homology of the filament appeared to be resolved when J. Z. Young (1977) reported a connection between the axial nerve of the vampire filament and ganglia of the circumoral commissure that connects the axial nerves of all arms. This evidence, however, could not be confirmed by Young and Vecchione (1996).
     • Recent confirmation of Pickford's earlier evidence (Pickford, 1946) that the vampire hatchling has relatively thick (i.e., more arm-like) filaments supports the homology between arms and filaments (Young and Vecchione, 1999)
       
       
       
       Figure. Vampyroteuthis infernalis hatchling. Figures modified from Young and Vecchione, 1999. Top - Dorsal and slightly lateral photograph of the lightly stained (methylene blue) hatchling. Bottom - Lateral, dorsal and anterior drawings of the same hatchling. Morphologically the specimen is an advanced embryo with a large internal yolk sac.
       
       
     • The weight of evidence indicates that the filaments represent arms II. Proof may depend on examining the embryonic development of the vampire squid which is not feasible at present.
       
       
   • The filament:
     • Each long filament has muscles that run the along one side. Contraction of these muscles causes coiling of the filament and its retraction into the small filament pocket that lies at the base of the web. The following images show a filament that has been removed from a recently dead vampire. Most of the filament has an orange pigmentation; the cut base (arrow), however, is abruptly broader and has a pink color. Much of the tight coiling has been disrupted by handling.
       Figure. Filament of Vampyroteuthis infernalis.
       Left - The entire filament from the cut base (arrow) to the filament tip.
       Right - Enlargement of the upper part of the left image showing how tight some of the undisturbed coils can be. Photographed in a small dish aboard ship by Henk-Jan Hoving.
     • Under the dissecting microscope numerous, straight "hairs" are seen extending from the filament. These "hairs" are thought to aid in capturing food. 
       Figure. V. infernalis filament. Photomicrograph of a portion of the intact, fresh filament. Photograph by H. Hoving.
     • The Histology of the filaments has been examined by Hoving and Robison, 2012. The "hairs" could not be detected in the paraffin-embedded sections but could be seen in SEM photographs collapsed onto the surface of the filament; there, they appear to be tree-like branching structures rather than simple hairs. The surface of the filament consists of a layer of "vacuolated" cells with little internal structure. In SEM photographs, these cells are also collapsed and the "hairs" seem to arise at the junctures between these cells. Beneath the vacuolated cell layer is a layer of simple cuboidal cells with larger cells broadly interspersed. These larger cells are apparently nerve cells as they have extensions that pass into a large, axial nerve. A large muscle exists along one side of the filament and its contraction results in the coiling of the filament.
     • The "hairs" apparently secure food particles (Hoving and Robison, 2012). The retraction of the filaments is a slow process and the vampire squid may swim to the portion of the tentacle holding the food (Hunt, 1996) and enclose it within the web. Next, the filament is drawn between the arms, wrapped in mucous produced by cells on the suckers, sucker stalks and cirri, and passed to the mouth (Hoving and Robison, 2012).
3. Head
   
   • Eyelid circular, without anterior optic sinus.
   • Beaks:
     
     • Lower beak:
       

       	V. infernalis, sex ?, 13 mm GL, 1.0 mm LRL, Hawaiian waters. Photographs by R. Young	V. infernalis, sex ?, ~40 mm ML, 2.2 mm LRL, Hawaiian waters. Photographs by R. Young.	V. infernalis, sex ?, ML ?, 5.3 mm LRL, Hawaiian waters. Photographs by R. Young.
       Side View
       Oblique View
       Oblique View
       “Top” view
       Oral view
       Front View
       Posterior Oblique View

     • About anaglyph 3D: Photographs here are in 3D, using anaglyph techniques which combines two photographs one in red and the other in cyan (blue + green) color. To get the 3D effect, you must use glasses with red and cyan lenses (filters) over your regular glasses. The form of the beak is far easier to interpret in 3D and we strongly recommend to the viewer that the glasses be obtained (this is especially helpful when viewing enlarged images - i.e. click on the image). These beak photographs were made using Red and Cyan Anachrome Aviator Glasses (see: http://www.anachrome.com/glassbuy.htm or http://www.amazon.com) which cost under $10.00 (USD).
     • Upper beak:
       

       	V. infernalis, sex ?, 13 mm GL, 1.0 mm LRL, Hawaiian waters. Photographs by R. Young	V. infernalis, sex ?, ~40 mm ML, 2.2 mm LRL, Hawaiian waters. Photographs by R. Young.	V. infernalis, sex ?, ML ?, 5.3 mm LRL, Hawaiian waters. Photographs by R. Young.
       Side View
       Oblique View
       Oblique View
       “Top” view
       Oral view
       Front View
       Posterior Oblique View

   • Nuchal cartilage present even though head and mantle broadly fused.
   • Central nervous system with incipient inferior frontal lobe system; superior buccal lobes adjacent (fused at edges) to posterior buccal lobes; suprabrachial commissure lies within brain.
4. Funnel
   
   • Funnel valve absent.
5. Mantle
   
   • Dorsal mantle cavity absent.
   • Mantle apex a highly-flexible muscular cone filled with gelatinous tissue located posterior to the cone of the gladius.
6. Fins
   
   • Two pairs of fins present during early ontogeny.
7. Spermatangia
   
   • Receptacle (deep sac) for spermatangia located anterior to each eye in females.
8. Shell
   
   • Shell a gladius with broad median field and broad conus with or without a short, narrow cyclindrical rostrum.
9. Photophores
   
   • Large circular, lidded organs (= "fin-base" organs or "large posterior photophores") present posterior to each adult fin .
   • Numerous small organs distributed over ventral surfaces of mantle, funnel, head and aboral surface of arms and web (="skin-nodule" organs or "epidermal photophores"). Two patches on dorsal surface of head look like aggregrated small photophores (see images in Introduction) but are photoreceptors (Herring, et al., 1994) and may be "Nuchal Organs" (see Parry, 2000).
   • Arm-tip organ(s) produce luminescent clouds consisting of microscopic glowing particles (Robison, et al., 2003).
   • Arm-tip organs that flash or glow (Hunt, 1996).
10. Pigmentation
    
    • Photographs, of the vampire squid, taken in situ at two different locations show the geographic differences in coloration:
      
      
      Figure. Vampyroteuthis infernalis.
      Left - Side view, central North Pacific.
      Right - Oblique-anterior view, Monterey Bay. Note the difference in color between the two. © MBARI
      
      
    • These photographs of Vampyroteuthis were taken in situ by an ROV of the Monterey Bay Aquarium Research Institute. The two photographs were taken from widely different areas in the North Pacific showing geographic differences in coloration. The left image is a side view of the vampire in an unusual pose. Note the peculiar nature of the web in this posture which seems to suggest how the web can contract and expand. Also peculiar are the small epidermal photophores on the arms that seem to align in series. In the right image, the arms are aligned and the presence of a web is hardly noticed.
11. Viscera
    
    • Oviducal glands at terminal end of oviducts.
    • Visceropericardial coelom extends posteriorly as a slender duct, to a small organ at the apex of the conus of the gladius. This duct is possibly a remnant siphuncle.
    • Photosensitive vesicles located immediately dorsal to funnel and, possibly, on dorsal integument in nuchal region.

Comments: Mantle Cavity

Ventral view of an immature, female V. infernalis with the mantle cavity of cut open. Posteriorly the lining of the mantle cavity has been cut away to better view the visceral nucleus. Note that there is no abdominal septum, no ink sac, no anal flaps. The visceropericardial coelom (dark region surrounding the ventricle, stomach and caecum) is large. Posteriorly, in the ventral midline, the mantle muscle terminates on the conus of the gladius. The photograph on the right shows a mature female. The arrangement of the viscera looks very different. The gladius, with a small rostrum and a dark "apical organ", is visible because surrounding tissue has been removed.

Figure. Vampyroteuthis infernalis.

Left - Ventral view, immature female, with the mantle cavity opened. Drawing by R. Young.
Right - The mantle cavity of a mature female. Photograph by R. Young.

Figure. View of a portion of the mantle cavity of V. infernalis. The stellate ganglion (arrow), a characteristic structure of cephalopods, appears as only a slight enlargement of a large mantle nerve. The nerve running toward the oviducal gland is the fin nerve. Photograph by R. Young.