Wunderpus photogenicus n. gen. and sp., a new octopus from the shallow waters of the Indo-Malayan Archipelago (Cephalopoda: Octopodidae)

Abstract

Wunderpus photogenicus n. gen. and n. sp. is a spectacular long-armed species that occurs on soft sediment habitats in shallow waters (typically less than 20 m deep) in Indo-Malayan waters. It is characterized by small eyes on elongate stalks, a long, conical papilla over each eye and a dramatic and fixed color pattern of white bars and spots over a brown-red background. The distribution of the species is centered in the Indo-Malayan Archipelago and extends from Vanuatu to Papua New Guinea, Indonesia and Malaysia, north to the Philippines. Animals typically emerge at dusk and dawn to forage in the twilight, primarily catching small crustaceans and fishes by flaring the arms and webs over patches of sand or coral rubble to trap enclosed (and typically buried) prey. The species also extends its arms into holes to probe for potential prey. The distinctive color pattern of this species is most pronounced when the octopus is disturbed or threatened by real or perceived attackers. It appears to be a warning display and may represent one of two scenarios: either 1) it warns that the octopus is directly toxic or venomous by nature or; 2) it represents impersonations of toxic or venomous creatures with similar color patterns which co-occur in the same habitat. The new genus and species is compared with, and distinguished from, other long-armed octopuses.

Full text

Molluscan Research 26(3): 128–140
Magnolia Press
http://www.mapress.com/mr/ ISSN 1323-5818
128 COPYRIGHT © 2006 MALACOLOGICAL SOCIETY OF AUSTRALASIA
Wunderpus photogenicus n. gen. and sp., a new octopus from the shallow waters of the
Indo-Malayan Archipelago (Cephalopoda: Octopodidae)
F. G. HOCHBERG1, MARK D. NORMAN2 AND JULIAN FINN2, 3
1Department of Invertebrate Zoology, Santa Barbara Museum of Natural History, 2559 Puesta del Sol Road, Santa Barbara, California
93105-2936 USA
Corresponding author fghochberg@sbnature2.org
2Sciences, Museum Victoria, GPO Box 666E, Melbourne, VIC 3001, AUSTRALIA
EMAIL mnorman@museum.vic.gov.au
3School of Biological Sciences, La Trobe University, Bundoora, VIC 3086, AUSTRALIA
EMAIL jfinn@museum.vic.gov.au
Abstract
Wunderpus photogenicus n. gen. and n. sp. is a spectacular long-armed species that occurs on soft sediment habitats in shallow
waters (typically less than 20 m deep) in Indo-Malayan waters. It is characterized by small eyes on elongate stalks, a long,
conical papilla over each eye and a dramatic and fixed color pattern of white bars and spots over a brown-red background. The
distribution of the species is centered in the Indo-Malayan Archipelago and extends from Vanuatu to Papua New Guinea,
Indonesia and Malaysia, north to the Philippines. Animals typically emerge at dusk and dawn to forage in the twilight,
primarily catching small crustaceans and fishes by flaring the arms and webs over patches of sand or coral rubble to trap
enclosed (and typically buried) prey. The species also extends its arms into holes to probe for potential prey. The distinctive
color pattern of this species is most pronounced when the octopus is disturbed or threatened by real or perceived attackers. It
appears to be a warning display and may represent one of two scenarios: either 1) it warns that the octopus is directly toxic or
venomous by nature or; 2) it represents impersonations of toxic or venomous creatures with similar color patterns which co-
occur in the same habitat. The new genus and species is compared with, and distinguished from, other long-armed octopuses.
Key words: Wunderpus, octopus, Octopodidae, Cephalopoda, Indo-Malayan Archipelago, taxonomy
Introduction
In recent years, increased interest in the marine life of the
tropical Indo-Malayan Archipelago (particularly Indonesia)
has lead to the discovery of a diverse and largely unstudied
fauna. The spread of dive resorts and dive boat charters
throughout this region, combined with significant
improvements in underwater photographic and video
technology, has resulted in a flood of images to the outside
world of the spectacular life in these waters.
Amongst the images and reports emerging from
Indonesia in the mid 1980’s was a remarkable long-armed
octopus, which quickly received the nickname “Wunderpus”
(also referred to as “Wonderpus”). At the same time,
collectors in the Philippines and Indonesia started supplying
the home and public aquarium trade with occasional live
specimens of this octopus.
Despite all the reports and images, specimens required
for formal description of this species proved difficult to
obtain. Over the past two decades relatively few specimens
of this octopus have been preserved and archived in museum
collections. This distinctive animal is described here and
distinguished from another recently described species of
long-armed octopus from the same region with which it
frequently has been confused, the “Mimic Octopus”
(Norman and Hochberg 2005).
Material and Methods
A total of ten specimens were amassed from diverse sources.
Alex Kerstitch kindly donated two specimens; one he
collected in Vanuatu, the other was obtained from an
unknown locality through commercial aquarium suppliers.
Two specimens, collected by B. Bruguier, from unspecified
localities in Bali, Indonesia were supplied by Bruno Condé,
Director of the Aquarium Tropical, Musée de Zoologie,
Université de Nancy, France. An additional mature female,
obtained through the commercial aquarium trade, was
donated by John Forsythe of the National Research Center
for Cephalopods, Galveston, Texas, USA. Most recently
three specimens, probably collected in the Philippines, were
purchased at an aquarium store in Berkeley, California.
They were donated by Dr. Roy Caldwell following study at
the University of California, Berkeley. The above eight
specimens have been lodged in the Invertebrate Zoology
collection of the Santa Barbara Museum of Natural History
(SBMNH). Two more specimens were found in the
collections of the California Academy of Sciences, San
Francisco, California, USA (CASIZ) and the Australian
Museum, Sydney, Australia (AMS).
Additional color photographic images, videos,
distributional data and behavioral information were kindly
supplied by a number of keen amateur and professional
underwater photographers and filmmakers. The second

WUNDERPUS PHOTOGENICUS N. GEN. AND SP. FROM INDO-MALAYAN ARCHIPELAG 129
author observed four live individuals in situ while diving in
northern Sulawesi in 1998 and 1999.
The taxonomic methodology and diagnostic characters
used below follow Norman and Sweeney (1997) and
Norman et al. (1997).
Abbreviations: coll.—collected by; ML—mantle
length. Institutions: AMS—Australian Museum, Sydney,
Australia; CASIZ—California Academy of Sciences,
Invertebrate Zoology, San Francisco, USA; SBMNH—Santa
Barbara Museum of Natural History, California, USA; MV–
Museum Victoria, Melbourne, Australia.
SYSTEMATICS
Family Octopodidae d’Orbigny, 1840
Genus Wunderpus n. gen.
Type Species
Wunderpus photogenicus n. sp.
Diagnosis
Body small; mantle thin-walled, weakly muscular,
elongate ovoid to amphora-shaped; mantle aperture wide;
funnel organ small, W-shaped, lateral limbs very short;
interbrachial water pore system absent; stylets present, short,
chitinous (not mineralized); eyes small on elongate stalks;
gills with 6–7 (rarely 5) lamellae per demibranch plus
terminal lamella; arms long, thin and muscular, >5 times
mantle length, dorsal arms shortest, arm formula typically
4=3=2>1; arm autotomy occurs at the level of the 8th to 12th
proximal sucker; webs thin and elastic, extend entire length
of arms as well-developed ventral margin; suckers biserial,
small and widely spaced; suckers normal, functional to tips
of all arms other than hectocotylized arm of males; enlarged
suckers absent in both sexes; right arm III hectocotylized,
very short, less than 1/3 length of opposite arm; copulatory
organ with ligula and calamus; ligula size small to moderate
(<6% of arm length) with distinct groove; crop with distinct
diverticulum; posterior salivary glands larger than buccal
mass; intestine relatively short and robust without distinct U-
shaped loop; terminal organ (penis) short with simple short
diverticulum; spermatophores unarmed; egg size small
(chorion capsule of mature ovarian eggs, 2.9–3.6 mm long).
Skin largely smooth between erectile primary and secondary
papillae; single elongate, conical papilla present medially
over each eye (supraocular), tip rounded; body uniformly
brown to red color, patch and groove lattice system absent;
diagnostic pattern of distinct white bands along arms and
white spots and short transverse bars on mantle, head and eye
stalks; ocelli absent.
Etymology
Wunderpus: from the German ‘wünder’ meaning
“marvel or wonder”.
Remarks
The taxonomy of the family Octopodidae at the generic
level is in need of major revision. Over 90% of the 200+
species coined within this family have been placed in the
catchall genus Octopus Cuvier, 1797 (Hochberg et al. 2005;
Norman and Hochberg 2005b). A number of taxonomic
studies based on morphology have recognized distinct
species-complexes within the genus Octopus as it currently
stands (e.g., Robson 1929; Norman 1992a; Norman and
Hochberg 1994, 2005b; Norman and Sweeney 1997).
Recent molecular studies into the phylogeny of the octopuses
of the family Octopodidae (Hudelot 2000; Takumiza et al.
2005; Guzik et al. 2005; Strugnell et al. 2005) have
demonstrated that the genus Octopus is polyphyletic. Many
of the clades proposed in earlier morphological studies are
supported by molecular studies. The first step in the review
of the generic level taxonomy within the family Octopodidae
is to delineate taxa clearly distinct from the genus Octopus
sensu stricto as represented by the type species of this genus,
O. vulgaris Cuvier, 1797, and its relatives (a group treated in
the literature as the "O. vulgaris species-complex", Norman
and Sweeney 1997).
Wunderpus n. gen. is distinguished from Octopus s.s. by
the following characters: eyes small, on elongate stalks
(versus moderate to large-sized unstalked eyes); single,
elongate, blunt-tipped papilla over each eye (versus two or
more smaller supra-ocular papillae); characteristic two-tone
color pattern with fixed, relatively limited repertoire (versus
highly variable color patterns); skin smooth (versus a "patch
and groove" lattice-like sculpture); arm autotomy occurs at a
set level at the arm bases (versus no autotomy); male
hectocotylised arm short, ~30% of opposite arm length
(versus >70%); enlarged suckers absent (versus present on
arms of mature males); and funnel organ W-shaped with very
short outer limbs, length <40% of medial limbs (versus
>80%).
At present we recognize only one species in the new
genus. See Discussion below for delineation of Wunderpus
n. gen. from other long-armed octopuses in the family
Octopodidae.
Wunderpus photogenicus n. sp.
Figures 1–6; Table 1
Octopus horridus – Hanlon and Messenger 1996: 72; photo fig. 5.6I
[photo: J. Forsythe—collection locality not stated].
Octopus sp. 7.—Gosliner et al. 1996: 190; photo 679 [photo: M.
Severns—Luzon, Philippines].
Octopus sp. 20.—Norman 2000: 13, 304–305, 315 [photos: A.
Kerstitch—Malekula Island, Vanuatu Islands; M.D. Norman
and D. Nielsen-Tackett— both North Sulawesi, Indonesia].
“Banded Octopus”.—Kerstitch 1989: 375.—Kerstitch 1992: 34–35
[photo: A. Kerstitch—Malekula Island, Vanuatu Islands].
“Poulpe Zébré”.—Louisy 1996: 27 [photo: P. Louisy—Bali,
Indonesia]
“Ornate Octopus”.—Steene 1998: 250 [photo: R. Steene—Lembeh
Strait, North Sulawesi, Indonesia].
“Wunderpus”.—Norman 2000: 304, 315.—Norman and Reid
2000: 78 [photos: A. Kerstitch—Malekula Island, Vanuatu
Islands].—Halstead 2000: 272 [photos: B. Halstead—Madang,
Papua New Guinea; Vanuatu Islands].
“Wonderpus”.—Norman and Hochberg 2005a: 60 [Table 1].
“Mimic Octopus” (in part).—Butvill 2003: 38–40 [photos: D.
Nielsen-Tackett—collection locality not indicated].

HOCHBERG (2006) MOLLUSCAN RESEARCH, VOL. 26
130
TABLE 1. Wunderpus photogenicus n. sp. Counts and
measurements (mm) of males. Abbreviations: A–E: web sectors
starting from dorsal sector; D = damaged; Hc = hectocotylized arm
of male (right arm 3); max = maximum depths, lengths or counts
per arm pair; + = arm tip missing; ** = all or most arms missing.
Type Material
Holotype: male (mature), 24 mm ML; INDONESIA,
Bali, specific locality and depth not indicated, coll.
Neumann, date unknown, AMS C304110.
Paratypes: 1 female (mature, gravid), 27 mm ML;
VANUATU ISLANDS, Malekula Island, 150 m off shore,
~16º15’S, ~167º30’E, on sand bottom under dead coral slabs
and rocks, 20 m, coll. A. Kerstitch, M/V Kalinda, field no.
AK 881124, 24 November 1988, by hand with quinaldine;
SBMNH 369471. 1 female (mature, gravid), 36 mm ML;
PHILIPPINES, specific locality not indicated, coll.
commercial aquarium collector, spring 1986, SBMNH
345682 [ex National Resource Center for Cephalopods,
Galveston, Texas; maintained in aquarium, died 28 April
1986; photos by John Forsythe of live animals in aquarium].
1 male (mature), 20 mm ML; INDONESIA, Bali, specific
locality and depth not indicated, coll. B. Bruguier, 22 March
–18 April 1996; SBMNH 345680 [ex Musée de Zoologie,
Nancy, France; photos by Patrick Louisy of live animal;
DNA voucher]. 1 female (mature, gravid), 35 mm ML;
INDONESIA, Bali, additional locality data not available,
coll. B. Bruguier, 14 March – 12 April 1996; SBMNH
345681 [ex Musée de Zoologie, Nancy, France; photos by
Patrick Louisy of live animal; DNA voucher].
Material Examined
Holotype and paratypes (see above). 1 female
(submature), 23 mm ML, locality presumed to be the
PHILIPPINES (collected by commercial aquarium
collectors; locality and depth details not available),
purchased in 1986 by Steinhart Aquarium from tropical fish
dealer in Brisbane, Australia, CASIZ 081006 [photos by
Dustin Chivers of live animal in aquarium]. 1 female
(submature), 35 mm ML, purchased in 1985 by Alex
Kerstitch from aquarium store in San Francisco, California,
SBMNH 345679 [source originally provided as ‘Indian
Ocean’; videos and color photos by Alex Kerstitch of live
animal in aquarium]. 1 male (mature), 20 mm ML,
purchased in 2005 by Roy Caldwell from aquarium store
(The Octopus’s Garden) in Berkeley, California; SBMNH
369187 [maintained in aquarium, died April 2005; DNA
voucher]. 1 female (submature) 32 mm ML; purchased in
2005 by Roy Caldwell from aquarium store (The Octopus’s
Garden) in Berkeley, California; SBMNH 369188
[maintained in aquarium, died April 2005; DNA voucher]. 1
male (mature), 20 mm ML, purchased in 2005 by Roy
Caldwell from aquarium store (The Octopus’s Garden) in
Berkeley, California; SBMNH 369470 [maintained in
aquarium, died April 2005; DNA voucher].
Photographic and/or Video Records Examined
(specimens not captured)
INDONESIA: Lembeh Strait, North Sulawesi,
01º26’N, 125º13’E, 1 mi north of Bitung, 10–15 ft [3–5 m],
photos by Bill MacDonald, 03 October 1996; north of
Bitung, near Police Pier, on black sand, 9 m, photos by Mark
D. Norman, 14 October 1998. Sarena Kecil Island, off south
side on sandy slope just above rubble area, 50 ft [15 m],
Repository AMS SBMNH
Catalog number C304110 345680
Status Holotype Paratype
Sex male male
Maturity mature mature
Total Length 197 149
Total Wet Weight (g) 11.2 7.0
Dorsal Mantle Length 24 20
Ventral Mantle Length 16 14
Mantle Width 12.2 10.8
Head Width 12.8 13.2
Funnel Length 10.0 10.1
Free Funnel Length 5.5 3.0
Funnel Organ Length -lateral limb 1.0 1.3
-medial limb 2.4 3.4
Web Depths (max per pair) A 13 12
B 13 12
C 13 13
D 9 D
E 10 11
Web Depth Formula C=B=A>E>D C>A=B>E
Arm Lengths (max per pair) 1 123 110
2 173 122
3 111+ 123
Hc 47 38
4 131+ 120+
Arm Length Formula 2>1** 3=2>1
Arm Width 4.0 4.0
Sucker Diameter 2.0 1.8
Sucker Counts (max per pair) 1 186 160
2 - 162
3 D 152+
Hc 64 64
4 D D
Gill Count 5–6 6–7
Ligula Length 1.6 2.0
Calamus Length 0.5 1.0
Terminal Organ Length 4.3 4.1
Diverticulum Length 1.6 1.6
Spermatophore Count 31 24
Spermatophore Length 9.9 10.4
Spermatophore Width 0.4 0.3
Spermatophore Reservoir Length 4.6 5.0
Spermatophore Cord Whorls 32–36 ~30
Ejaculatory Apparatus Whorls ~17 -

WUNDERPUS PHOTOGENICUS N. GEN. AND SP. FROM INDO-MALAYAN ARCHIPELAG 131
photos by Denise Nielsen-Tackett, between August and
October 1996. Aer Perang (site of fresh water spring), sandy
area with scattered rubble, 9 m, photos by Mark D. Norman,
13 October 1998; 35–40 ft [to 12 m], photos by Annie
Crawley, 2002. Alor Island, Timor; photos by Roberto
Sozzani, date unknown.
NEW GUINEA: Milne Bay, 10º22’S, 150º30’E, 40 ft
[~12 m], photos by Fred Bavendam, M/V Telita, date
unknown; depth unknown, photos by Ron and Valerie
Taylor, date unknown. Madang, ~5º00’S, ~145º30’E, depth
unknown, photos by Bruce Halstead, date unknown. Irian
Jaya, locality details and depth unknown, photos by Robert
Sozzani, 2002.
PHILIPPINES: Luzon, Batangas, ~14º00’N,
~121º00’E, depth unknown, photos by Mike Severns, date
unknown [= Octopus sp. 7 in Gosliner et al. 1996]. Anilao,
photos by Jeffrey Rosenfield, 2002; photos by Paul Osmond,
date unknown.
SOLOMON ISLANDS: Florida Island Group, Gavutu,
~9º00’S, ~160º00’E, depth unknown, photos by Ted DeFeo,
1998.
MALAYSIA: Kapalai, photos by Nils Aukan, date
unknown.
VANUATU ISLANDS: Location and depth, photos by
Bruce Halstead, date unknown.
Diagnosis
Small-bodied animals (mantle lengths to 36 mm; total
lengths to over 230 mm; total weight weight to at least 26 g);
elongate, delicate species; eyes small on long stalks, head
appears Y-shaped; funnel organ W-shaped with short outer
limbs; stylets short, chitinized; gills with 6–7 (rarely 5)
lamellae per demibranch plus terminal lamella; arms long
(typically 5–7 times mantle length), almost equal in length,
dorsal arms slightly shorter (arm formula 4=3=2>1);
evidence of arm autotomy present between 8th and 12th
proximal sucker; right third arm of males hectocotylized,
very short (30% length of opposite arm); copulatory organ
small (length 3.4–5.3% of hectocotylized arm length)
consists of distinct ligula and calamus; ligula squat,
copulatory groove well developed, with single medial row of
small raised bumps; calamus distinct; enlarged suckers
absent in both sexes. Sucker counts on normal arms range
between 160–230; hectocotylized arm with 64 suckers in two
available males. Webs thin and retractile, extend along
ventral margin to tips of all arms; webs shallow in preserved
material (web depths 7.5–11% of longest arm) but capable of
considerable expansion in live animal; relative lengths of
web sectors variable in preserved material with sector A
typically most shallow. Terminal organ in males short and
straight with simple diverticulum; spermatophores short
(around 50% mantle length), produced in moderate numbers
(24 and 31 in storage sac of examined males);
spermatophores unarmed; mature ovarian eggs small
(chorion capsule length 2.9–3.6 mm). Skin color uniform
brown to red; distinct pattern of defined white bands on arms
and white spots and stripes on mantle; skin generally smooth,
primary papillae present and often conspicuous in white
areas on dorsal mantle; patch and groove system absent;
single elongate, conical papilla present medially over each
eye (supra-ocular), tip bluntly rounded.
Description
The following description is based on detailed
examination of two mature males and five females (three
submature and two mature). Counts and measurements for
this material are presented in Table 1–2. Indices are included
in the text: ranges in (parentheses); means in italics. One
submature female (34.5 mm ML, SBMNH 345679) is poorly
preserved, the body and arms are very soft and stretched.
Due to this distortion, data on arm and mantle lengths, and
web depths for this specimen are excluded from calculations
in the following description.
Small-bodied species (Fig. 1A), mantle lengths to at
least 24 mm in mature males, 36 mm in mature females; total
lengths to over 230 mm; weights (wet) to at least 26 g.
Mantle elongate-ovoid to amphora-shaped (Figs. 1A–C);
walls of mantle thinly muscular; mantle width approximately
half mantle length (males and females 50.7–54.7–63.3%;
gravid female with distended mantle 69.8%). Head of
moderate width, around half mantle length (males 53.6–
59.8–66.0%; females 37.4–44.8–49.1%; gravid female
27.9%). Head typically wider than mantle in males,
narrower than mantle in females due to influence of ovary
(head width of males 104.9–113.6–122.2% of mantle width;
of females 73.7–79.7–88.0%; of gravid female 40.0%). Eyes
small and stalked, often raised in live animals so head
appears Y-shaped. Neck region distinct. Stylets present,
short (4.9 mm, 13.9% of mantle length), non-mineralized
(Fig. 1D). Pallial aperture of moderate width, approximately
1/2 mantle circumference. Funnel thin walled, moderately
long (39.3–43.4–50.5% of mantle length); free about half its
length (29.7–46.1–56.2%). Funnel organ W-shaped (Fig.
1E), located in middle of dorsal funnel; large, medial limbs
about 1/3 length of funnel (24.0–30.7–37.0%); limbs wide
with lateral limbs much shorter than medial limbs (20.0–
36.3–41.7%).
Webs thin, shallow relative to arm length (deepest 7.5–
9.5–10.6% of longest arm length); dorsal web typically
shortest but formulae variable (see Table 1–2). Web margins
well developed on ventro-lateral edge of arms, extend along
entire length of all arms; webs frequently flared by live
animals during displays (Fig. 4A) or when foraging (Fig.
4D). Arms long, longest 5–7 times mantle length (5.4–6.3–
7.2% of mantle length); width moderate (12.3–16.3–20.0%
of mantle length); arms roughly triangular in cross-section
(aboral surface as apex), arm width increases from mouth
through proximal quarter of arm length, width uniform in
middle quarter, tapers evenly in distal half. Dorsal arms
appear shortest in few intact specimens; ventral or lateral
arms typically longest (see Table 1–2). Numerous arms
missing in several specimens; typically severed between
proximal suckers 8–12 suggesting arm autotomy likely
occurs (see life history section below for discussion).
Suckers biserial; small and widely spaced (Fig. 1F);

HOCHBERG (2006) MOLLUSCAN RESEARCH, VOL. 26
132
infundibulum fleshy with distinct radial cushions; rim
scalloped; diameter of suckers small, slightly larger in males
(males 8.4–8.7–9.0% of mantle length; females 5.3–6.7–
8.3%). Enlarged suckers absent in both males and females.
Sucker counts range between 160–230 on normal arms (male
162–186 in two specimens; female 233 in single specimen
where counts were possible). Hectocotylized arm with 64
suckers in both males.
Third right arm hectocotylized in males; very short (1.9,
2.0 times mantle length), less than 1/3 length of opposite arm
(30.9% in male specimen with intact opposite arm).
Copulatory organ (Figs. 1G–H) small (3.4–4.4–5.3% of
hectocotylized arm length). Ligula squat with well-
developed copulatory groove; single medial row of small
FIGURE 1. Wunderpus photogenicus n. sp. A. Stylized dorsal view of mature male (paratype, 20 mm ML; SBMNH 345680); B–C, F.
Details of mature female (paratype, 27 mm ML; SBMNH 369471); B. Mantle, dorsal view; C. Mantle, left lateral view; D. Left stylet
(female, 36 mm ML; SBMNH 345682); E, G–H. Details from mature male (holotype, 24 mm ML; AMS C304110); E. Funnel organ on
dorsal surface of funnel; F. Arm section showing suckers and color pattern; G. Copulatory organ, oral view; H. Copulatory organ, left lateral
view. Abbreviations: C = calamus; CG = copulatory groove; DMP = dorsal mantle papilla; Hc = hectocotylized arm; L = ligula; PMWS =
posterior mantle white spot; SOP = supra-ocular papilla; SpG = spermatophore groove.

WUNDERPUS PHOTOGENICUS N. GEN. AND SP. FROM INDO-MALAYAN ARCHIPELAG 133
raised bumps present in groove; calamus low, small (31.3–
40.7–50.0% of ligula length). Spermatophore groove well
developed and fleshy.
Gills with 6–7 lamellae (rarely 5) per outer demibranch
plus terminal lamella.
Digestive tract (Fig. 2A). Buccal mass large, about 2/3s
length of digestive gland. Anterior salivary glands of moderate size, approximately 1/3 length of buccal mass;
posterior salivary glands large, as long or longer than buccal
FIGURE 2. Wunderpus photogenicus n. sp. A–E. Digestive system (paratype, female 27 mm ML; SBMNH 369471); A. Entire digestive
tract; B. Upper beak, left lateral view; C. Lower beak, left lateral view; D. Lower beak, ventral view; E. Radula. Abbreviations: AF = anal
flaps; ASG = anterior salivary gland; BM = buccal mass; Ca = caecum; CD = crop diverticulum; Cr = crop; I = intestine; IS = ink sac; L1 =
first lateral tooth; L2 = second lateral tooth; M1 = first marginal tooth; MP = marginal plate; O = oesophagus; P = pancreas; PSG = posterior
salivary gland; R = rachidian tooth; S = stomach.
O

HOCHBERG (2006) MOLLUSCAN RESEARCH, VOL. 26
134
mass. Crop with distinct diverticulum. Stomach bipartite;
caecum with single whorl, ducts to digestive gland short and
wide. Intestine relatively short, wide; rectum short,
muscular; distinct U-shaped loop in intestine absent.
Digestive gland approximately ovoid with slight posterior
lobes. Pancreatic tissue evident in posterior 20% of digestive
gland. Ink sac small, narrow and shallow, only slightly
embedded in surface of digestive gland. Anal flaps present,
elongate.
Beaks (Figs 2B–D). Upper beak hood relatively small,
around 30% of beak length (30.3% in SBMNH 64198);
rostrum bluntly hooked, cutting edge rounded; crest slightly
rounded. Lower beak, hood and crest rounded; rostrum
pointed, cutting edge sharp; lateral walls flared with concave
posterior margin (Fig. 2D). Radula (Fig. 2E) with 7 teeth
plus 2 marginal plates present per transverse row. Rachidian
tooth with very tall mesocone and single lateral cusp on each
side in symmetrical arrangement. Lateral cusps migrate
from medial to lateral position over 3 rows. First lateral teeth
almost square with slight cusp on lateral edge. Second
lateral teeth with single distinct cusp at medial end. First
marginal teeth tusk-shaped. Marginal plates roughly square.
FIGURE 3. Wunderpus photogenicus n. sp. A–B. Male reproductive system (holotype, 24 mm ML; AMS C304110); A. Reproductive tract
of mature male, ventral view; B. Spermatophore; C–D. Female reproductive system (paratype, 36 mm ML; SBMNH 345682); C.
Reproductive tract of mature female, ventral view; D. Mature ovarian egg showing capsule and stalk. Abbreviations: Ap = appendix; AG =
accessory gland; D = diverticulum; DO = distal oviduct; EA = ejaculatory apparatus; O = ovary; OC = oral cap thread; OE = ovarian egg; OG
= oviducal gland; PO = proximal oviduct; SG = spermatophoric gland; SR = sperm reservoir; SS = spermatophore storage sac; T = testis; TO
= terminal organ.

WUNDERPUS PHOTOGENICUS N. GEN. AND SP. FROM INDO-MALAYAN ARCHIPELAG 135
FIGURE 4. Wunderpus photogenicus n. sp. Photographs of live animals: A. Alarm display of female from Vanuatu; aquarium photograph
(27 mm ML, SBMNH 369471; Photo: A. Kerstitch); B. Female from Philippines stretched out on substrate; photographed in Steinhart
Aquarium (23 mm ML, CASIZ 081006; Photo: D. Chivers); C. Foraging animal showing suppressed color pattern; in situ photograph,
Lembeh Strait, North Sulawesi, Indonesia (Photo: M.D. Norman); D. Foraging animal showing flared web of speculative pounce posture; in
situ photograph, Lembeh Strait, North Sulawesi, Indonesia (Photo: D. Nielsen-Tackett).

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Male reproductive tract (Fig. 3A). Terminal organ short
and robust (18.0–19.3–20.5% of mantle length);
diverticulum simple single loop (37.2–38.1–39.0% of organ
length); genital aperture subterminal. Vas deferens duct thin,
long. Spermatophoric gland robust with distinct recurved
coil. Accessory gland robust, distal end reflexed. Appendix
present at junction of spermatophore storage sac and
spermatophoric ducts. Spermatophore storage sac broad and
short, spermatophores visible through thin sac wall;
spermatophores present in moderate numbers (24 and 31 in
storage sacs of examined males).
Spermatophores (Fig. 3B) short (9.9, 10.4 mm), around
50% of mantle length (41.4–46.7–52.0%); narrow without
swellings (2.9–3.5–4.0% of spermatophore length);
FIGURE 5. Wunderpus photogenicus n. sp. Photographs of live animals: A. Alarm display; in situ photograph, Lembeh Strait, North
Sulawesi, Indonesia (Photo: M.D. Norman); B. Detail of male from Bali showing small stalked eyes; photographed in Nancy Aquarium (20
mm ML, SBMNH 345680; Photo: P. Louisy); C. Detail of female possibly from Philippines showing raised papillae on dorsal mantle and
head; aquarium photograph (35 mm ML; SBMNH 345681; Photo: A. Kerstitch); D. Foraging animal showing regenerating arms (left arms 2
& 3, right arm 2); photographed in situ in Lembeh Strait, North Sulawesi, Indonesia (Photo: M.D. Norman); E. Male (on right) clasping
female in mating posture; photographed in situ in Lembeh Strait, North Sulawesi, Indonesia (Photo: D. Nielsen-Tackett).

WUNDERPUS PHOTOGENICUS N. GEN. AND SP. FROM INDO-MALAYAN ARCHIPELAG 137
unarmed. Sperm reservoir about 50% spermatophore length
(46.5–47.3–48.1%); sperm cord coiled in approximately 30–
35 coils.
Female reproductive tract (Fig. 3C). Ovary large;
proximal oviducts short, about one-half length of distal
oviducts; oviducal gland dark in distal half, without evident
radiating chambers; distal oviducts slightly swollen; genital
aperture subterminal. Ovarian eggs (Fig. 3D) relatively
small with short stalks (capsule length 2.9–3.6 mm; 8.2–
10.1% of mantle length); 4 follicular folds present.
Approximately 2000 mature ovarian eggs present in single
gravid female examined. Spawned eggs carried in arms of
female (Miske and Kirchhauser 2006).
Colour of live animals (Figs. 4, 5). Body uniformly red
dorsally and ventrally; with large, conspicuous white spots
and stripes on dorsal, lateral and ventral mantle, head and
eyes; large, round white spot present on posterior dorsal tip
of mantle; thin white strip extends from posterior ventral eye
down stalk onto lateral anterior mantle; conspicuous white
bands alternate with red and extend down dorsal and lateral
arms from base to tips of arms; each white band complex
consists of wide medial band plus narrow band on either side
of medial band (Fig. 4C); small round white spot often
visible dorsally in center of wide white arm band; elongate
papilla over each eye, red with white tip. Dark ocellus on
arm base below eyes absent. Ventral mantle and arms light
in color; sucker bases yellowish cream. Dorsal mantle and
frontal white spot complexes (sensu Packard and Sanders
1971) not evident.
Color and patterns of well-preserved specimens similar
to those seen in live animals and photographs; body
uniformly light red; larger white spots and stripes easily
visible on mantle, head and arms; smaller spots and stripes
often not observed.
Skin smooth; patch and groove system absent.
Conspicuous primary papillae present on dorsal and lateral
mantle only in areas where white spots and stripes are
present; slight variations in patterns apparent with each
individual (Figs. 1B–C, 5C). Single elongate, conical papilla
present medially over each eye (supra-ocular), tip bluntly
rounded.
Etymology
Specific name in recognition of the considerable
photographic interest in this spectacular species in the media
in recent years.
Common Name
“Wunderpus”.
Distribution
Wunderpus photogenicus n. sp. is known from shallow
waters of the tropical Indo-Malayan Archipelago from
Indonesia and the Philippines, east to Papua New Guinea,
and the Solomon Islands and Vanuatu (Fig. 6). Reports from
underwater photographers and data from collected
specimens indicate a depth range from 0.5 to at least 20 m.
FIGURE 6. Wunderpus photogenicus n. sp. Map of distribution of
specimens and photographs examined. Symbols: = holotype;
= paratypes and additional specimens; = photographic records
and observations.
Taxonomic remarks
There are a number of other species of long-armed
octopuses within the family Octopodidae (see Table 1,
Norman and Hochberg 2005a). Wunderpus photogenicus n.
sp. previously has been confused with a similar Indo-
Malayan species that occurs in the same localities and habitat
types. The “Mimic Octopus” (Thaumoctopus mimicus
Norman and Hochberg, 2005) has been described in detail.
W. photogenicus n. sp. can be clearly distinguished from T.
mimicus on the basis of the following characters: a) single
long supra-ocular papilla over each eye, inflated and blunt-
tipped (versus very elongate and finely pointed); b) anterior-
ocular papilla on margin of each eye absent (versus present);
c) lower number of suckers on hectocotylized arm (64 versus
130–146); e) higher number of suckers on normal arms (to
233 versus to 283); f) shorter arm lengths (5.4–7.2 times
mantle length versus 7.4–10.2); g) fewer gill lamellae per
demibranch (5–7 versus 9); and h) well-defined color pattern
with a large, round white spot on the posterior tip of the
mantle (versus less clearly delineated color pattern with a U-
shaped white patch) (Figs. 4A, D; Norman and Hochberg
2005a).
Another genus of long-armed octopus from Australia
shows some similarities with Wunderpus photogenicus n. sp.
“The Banded Drop-arm Octopus” (Ameloctopus litoralis
Norman, 1992) is an intertidal species that also has long thin
arms (5–10 times mantle length) with alternating light and
dark purple-brown bands along their length. W. photogenicus
n. sp. can be easily distinguished from A. litoralis based on
the following characters: a) ink sac and anal flaps present
(versus absent); b) webs along the ventral edges of all arms
present (versus absent); c) spermatophores narrow (versus
inflated or swollen); d) funnel organ well-developed, W-
shaped (versus vestigial, consisting only of four small pads);
e) terminal organ short, squat with well-developed
diverticulum (versus linear in shape, without a diverticulum);
and f) distinct white markings on the mantle present (versus
absent) (Norman 1992).
Wunderpus photogenicus n. sp. has been reported in the

HOCHBERG (2006) MOLLUSCAN RESEARCH, VOL. 26
138
past under the name Octopus horridus d’Orbigny, 1826
(Hanlon and Messenger 1996). W. photogenicus n. sp. is
easily distinguished from O. horridus based on the following
characters: a) longer arms (5.4–7.2 times mantle length
versus 4.2–4.7); b) shallower webs (7.5–10.6% of arm length
versus 15%); c) lower number of suckers on hectocotylized
arm (64 versus about 90); d) higher number of suckers on
the normal arms (to 233 versus 125–135); e) enlarged
suckers in mature males absent (versus present); and f)
longer ligula length (3.4–5.3% of hectocotylized arm length
versus 1.6–2.0%).
Life history and behavior
Little is known of the life history and behavior of this
distinctive octopus. It has primarily been encountered on
soft sediment substrates where it occupies the burrows of
other animals or may dig its own burrow. Individuals were
found to occupy the same burrow for periods of at least three
weeks. This species appears to have a crepuscular activity
pattern, emerging to forage during half-light periods at dusk
(about 1600 hrs) and dawn. Reports of day activity from
photographers and film crews were determined to be
observations of another long-armed species, the “Mimic
Octopus” (see Norman and Hochberg 2005a).
TABLE 2. Wunderpus photogenicus n. sp. Counts and measurements (mm) of females. Abbreviations: A–E: web sectors starting from
dorsal sector; D = damaged; ind = indistinct; Hc = hectocotylized arm of male; + = arm tip missing; ** = all or most arms missing.
Repository SBMNH SBMNH SBMNH SBMNH CASIZ
Catalog number 345682 345681 369471 345679 81006
Status Paratype Paratype Paratype - -
Sex female female female female female
Maturity gravid mature submature submature submature
Total Length 234 128 D distorted D
Total Wet Weight (g) 26.5 20.9 7.6* 21.6 6.8*
Dorsal Mantle Length 36 35 27 35 23
Ventral Mantle Length ~29 D 20 23 11
Mantle Width 25.0 17.9 17 18 12
Head Width 10.013.213.114.611.0
Funnel Length 14.3 14.2 10.5 15.5 10.8
Free Funnel Length 5.2 6.9 5.9 8.3 5.5
Funnel Organ Length -lateral limb 1.6 1.5 1.4 ind 0.8
-medial limb 4.4 3.7 3.4 ind 4.0
Web Depths (maximum per pair) A D 12 10 9 10
B D 15 12 16 11
C D 18 16 20 10
D 16 20 12 18 12
E 12 19 12 15 11
Web Depth Formula D>E D>E=C>B>A C>B>E=D>A C>D>B=E>A D>B=E>C=A
Arm Lengths (max per pair) 1 D 155 D 187+ D
2 175+ 155+ D 249 D
3 190+ 191 D 195+ D
4 D 176 D 205 140+
Arm Length Formula D 3>4>1 D 2>4** D
Arm Width 4.4 4.9 4.4 distorted 4.2
Sucker Diameter 1.9 2.1 1.9 1.6 1.9
Sucker Counts (max per pair) 1 D D D D D
2 176+ D D D D
3 182+ D D D D
4 D 233 D D 120+
Gill Count 77677
Egg Length (chorion capsule) 3.6 2.9 1.6 submat. - -
Egg Width 1.2 0.9 0.7 submat. - -
Egg Number ~2000 - - - -

WUNDERPUS PHOTOGENICUS N. GEN. AND SP. FROM INDO-MALAYAN ARCHIPELAG 139
Two foraging methods have been observed for this
species:
Speculative “probing” of individual arms down
burrows in the mud. When prey are encountered they
are seized and withdrawn using suckers on the distal
tips of the arms.
Speculative “web-casting”, where the octopus arches its
narrow arms over an area of sand or mud forming an
umbrella-like empty frame. The thin, semi-transparent
webs are then rapidly extended to fill in the umbrella
and enclose any small fishes or crustaceans contained
within the area of the “cast” (Fig. 4D). Arm tips are
used to flush buried prey into the waiting web and
suckers.
The diet appears to consist of small fishes and crabs
(e.g. Calappa spp.) that co-occur in sand, mud and rubble
substrates. The second author observed one octopus
consuming a crab with a carapace as large as the mantle of
the octopus, which prevented the octopus from retracting
back into its lair while feeding. Captive animals have been
observed to be day active and readily fed on shrimps and
pieces of fish meat (A. Kerstitch and R. Caldwell, pers.
comm.).
As mentioned above, several of the specimens
examined were missing numerous arms, typically severed
around the level of the 8th to 12th proximal sucker.
Regeneration had commenced from some stumps. One wild
animal encountered by the second author possessed three
arms regenerating from the base (Fig. 5D). This species thus
appears to be capable of arm autotomy (severing an arm at a
set cleavage plane near the base), as has been documented in
other long-armed octopuses (Norman 1992b). Such behavior
provides a wriggling decoy or food offering to attackers,
while the animal escapes to regenerate the lost limb.
Predators are unknown, although they are likely preyed
upon by flounders and scorpion fishes. One individual
octopus rapidly fled from threats by an aggressive mantis
shrimp (stomatopod), which was resident within the
octopus’s home range. This reaction suggests that the shrimp
may be a direct competitor, a potential predator, or both.
The third author observed mating in Wunderpus
photogenicus n. sp. in the wild. The male mounts the female
and inserts the short hectocotylized arm (~30% of length of
opposite arm) into the mantle of the female (Fig. 5E). This
contrasts with the more typical distance or remote mating
and insemination behavior observed in other octopus species
which possess a long hectocotylized arm (e.g. Young 1962;
Norman and Finn 2001).
Mature ovarian eggs reach a capsule length of at least
3.6 mm long (approximately 10% of mantle length). Egg
size relative to mantle length has been used in other octopus
species to successfully predict hatchling behavior. Egg size
predicts planktonic hatchlings (see Miske and Kirchhauser
2006).
Certain postures and behavioural displays observed in
Wunderpus photogenicus n. sp. have been proposed to
represent mimicry of other animals. The display shown in
Figure 4A has been suggested to be impersonation of
scorpaenid lionfishes (Pterois spp.), which possess long
poisonous spines advertised by similar banded markings
(Kerstitch 1989). W. photogenicus n. sp. also has been
observed to sit with six arms down a burrow while two
opposite arms are undulated to produce the appearance of
another poisonous animal, a banded sea snake (Laticauda
sp.; Nielsen-Tackett and Tackett 2000).
Wunderpus photogenicus n. sp. is primarily active at
dusk and dawn on soft sediment substrates, which offer little
cover or refuge from fishes and other predators. The
dramatic color pattern of W. photogenicus n. sp. may have
evolved as a defensive strategy against these predators
through one of two scenarios. Color patterns and potential
mimicry behavior may have evolved to impersonate other
species, specifically banded poisonous animals such as lion
fishes or sea snakes. An alternative explanation is that this
species is itself poisonous and is advertising its toxicity in a
similar fashion to species of the deadly blue-ringed or blue-
lined octopuses (Hapalochlaena spp.) of Australia and Asia.
Brilliant iridescent blue markings (rings or lines) on these
animals warn potential attackers of the high toxicity of their
saliva. Toxicology of the salivary glands and other tissues of
W. photogenicus n. sp. may provide further insights into the
significance of the dramatic color pattern of this species.
Discussion
Wunderpus photogenicus n. sp. joins a growing list of
species of small octopuses with long arms that live on soft
sediment and/or in intertidal habitats around the world. The
majority of these taxa exhibit arm autotomy at a set plane
near the arm base. These taxa include Abdopus species of
the Indo-West Pacific (Norman and Finn, 2001); Amelocto-
pus litoralis Norman, 1992 of northern Australia; Macro-
tritopus defillipi (Verany, 1851) of the Atlantic Ocean and
Mediterranean Sea (and the potentially related ‘Octopus’ sp.
17 in Norman, 2000); Thaumoctopus mimicus Norman and
Hochberg, 2005 of the Indo-West Pacific (and the potentially
related ‘Octopus’ sp. 18 in Norman, 2000) and a number of
unplaced species including ‘Octopus’ mutilans Taki, 1942 of
Japan (Tsuchiya et al. 2002), ‘Octopus’ harpedon Norman,
2001 from northern Australia and ‘Octopus’ sp. B Voss and
Williamson, 1972 found in Hong Kong.
Generic placement of most of the latter species awaits
their thorough morphological description. The single largest
impediment to this process is access to well-preserved
material. Many of these species are best known from live
animal photographs, however little or no voucher material
has been collected. Because of the rarity of these animals
and the remote locations in which many live, problems occur
with collecting permits and access to appropriate expertise
and preservation techniques. The limited material that is
available, including that of the new species described herein,
comes primarily from the marine aquarium trade.
The octopod fauna of Indonesia and elsewhere in the
Indo-Malayan Archipelago still remains largely unstudied.

HOCHBERG (2006) MOLLUSCAN RESEARCH, VOL. 26
140
At least 50 undescribed species have been recognized by the
authors throughout the region. The group is in need of
extensive revision and sources of well-preserved voucher
specimens need to be developed.
Acknowledgements
The authors are very grateful for the assistance of the following
people. Donation of specimens: Roy Caldwell and Christine
Huffard (University of California, Berkeley); Bruno Condé
(Aquarium in Nancy, France); John W. Forsythe (National Resource
Center for Cephalopods, Galveston, Texas); Alex Kerstitch
(Tucson, Arizona); John McCosker (Steinhart Aquarium, San
Francisco, California). Assistance with museum collections: staff
(AMS); Dustin Chivers (CASIZ); Fred Naggs (The Museum of
Natural History, London, England). Sigurd von Boletzky
(Laboratoire Arago, Banyuls-sur-Mer, France) facilitated contact
with Patrick Louisy and Bruno Condé at the Nancy Aquarium.
Access to photographic images, videos and/or general information
on live animals: Fred Bavendam; Annie Crawley, Patrick Louisy,
Bill MacDonald; Susan Ritman-MacDonald; Ron and Valerie
Taylor; Denise Nielsen-Tackett and Larry Tackett. Information
from field observations: Christine Huffard (University of
California, Berkeley). Illustrations were prepared by Peter Gaede
(SBMNH). A preliminary draft was reviewed by Christine Huffard
and Michael Sweeney. Logistical and financial support for visits to
Indonesia by the second and third authors: Joe Kennedy, Zebra
Films, BBC Natural History Unit, and the Discovery Channel.
Financial assistance was provided to both senior authors by the
Museum National d’Histoire Naturelle, Paris through Visiting
Professor grants. This manuscript was completed with the
assistance and generosity of Renata Boucher-Rodoni, Guy Boucher,
and Bernard Metivier.
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