A remarkable new jellyfish (Cnidaria: Scyphozoa) from coastal Australia, representing a new suborder within the Rhizostomeae
ABSTRACT A new genus and species of rhizostome jellyfish, which cannot be placed in any known
family or suborder, is described from central eastern Australia. The Ptychophorae
suborder nov., can be separated from the two recognised suborders of the Rhizostomeae
in having hooded rhopalia rather than open pits, unbranched evenly-spaced radial
canals, and a large circular stomach. It also possesses a unique combination of some of
the primary diagnostic characters of rhizostome families in both current suborders.
Moreover, Bazinga rieki gen. nov., sp. nov. is unique among all rhizostomes in its very
small mature size. It is probable that this species has been previously mistaken for
juveniles of other species.
- SourceAvailable from: Allen G. Collins[Show abstract] [Hide abstract]
ABSTRACT: A stable phylogenetic hypothesis for families within jellyfish class Scyphozoa has been elusive. Reasons for the lack of resolution of scyphozoan familial relationships include a dearth of morphological characters that reliably distinguish taxa and incomplete taxonomic sampling in molecular studies. Here, we address the latter issue by using maximum likelihood and Bayesian methods to reconstruct the phylogenetic relationships among all 19 currently valid scyphozoan families, using sequence data from two nuclear genes: 18S and 28S rDNA. Consistent with prior morphological hypotheses, we find strong evidence for monophyly of subclass Discomedusae, order Coronatae, rhizostome suborder Kolpophorae and superfamilies Actinomyariae, Kampylomyariae, Krikomyariae, and Scapulatae. Eleven of the 19 currently recognized scyphozoan families are robustly monophyletic, and we suggest recognition of two new families pending further analyses. In contrast to long-standing morphological hypotheses, the phylogeny shows coronate family Nausithoidae, semaeostome family Cyaneidae, and rhizostome suborder Daktyliophorae to be nonmonophyletic. Our analyses neither strongly support nor strongly refute monophyly of order Rhizostomeae, superfamily Inscapulatae, and families Ulmaridae, Catostylidae, Lychnorhizidae, and Rhizostomatidae. These taxa, as well as familial relationships within Coronatae and within rhizostome superfamily Inscapulatae, remain unclear and may be resolved by additional genomic and taxonomic sampling. In addition to clarifying some historically difficult taxonomic questions and highlighting nodes in particular need of further attention, the molecular phylogeny presented here will facilitate more robust study of phenotypic evolution in the Scyphozoa, including the evolution characters associated with mass occurrences of jellyfish.Integrative and Comparative Biology 09/2010; 50(3):436-55. · 3.02 Impact Factor
Article: Synopsis of the Medusae of the World[Show abstract] [Hide abstract]
ABSTRACT: This work was started many years ago in the form of a card-index for my own use. At the beginning the building up of the index involved a great deal of work, but later on it saved me time for my studies, and it was not difficult to keep it up to date. One day my friends, Dr F. S. Russell, director of the Marine Biological Laboratory, Plymouth, and Dr W. J. Rees, British Museum (Nat. Hist.), London, saw this index, and they repeatedly urged me to put it into a form suitable for publication for the benefit of other students of medusae. I agreed, though hesitatingly, because my time was much occupied. A grant from the Browne Research Fund of the Royal Society of London, however, enabled me to employ an assistant, Mr J. Liitzen, who under my supervision could carry out the necessary revision and the typing, as well as some translation from Danish or other languages into English; he has done it remarkably well, and I wish to thank him for his industrious and painstaking collaboration.Journal of the Marine Biological Association of the UK 10/1961; 40:7 - 382. · 1.13 Impact Factor
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ABSTRACT: Inaug.-Diss.--Königsberg. Vita. "Abdruck aus 'Bibliotheca zoologica, hrsg. von dr. Rud. Leuckart und dr. Carl Chun. Cassel 1888. Heft III.'" "Benutzte literatur": 1 p. at end.
A remarkable new jellyfish (Cnidaria: Scyphozoa)
from coastal Australia, representing a new suborder
within the Rhizostomeae
CSIRO Marine and Atmospheric Research, GPO Box 1538, Hobart, Tasmania 7000. Email:
Peter J.F. DAVIE
Queensland Museum, PO Box 3300, South Brisbane, Queensland 4101.
A new genus and species of rhizostome jellyfish, which cannot be placed in any known
family or suborder, is described from central eastern Australia. The Ptychophorae suborder
nov., can be separated from the two recognised suborders of the Rhizostomeae in having
hooded rhopalia rather than open pits, unbranched evenly-spaced radial canals, and a
large circular stomach. It also possesses a unique combination of some of the primary
diagnostic characters of rhizostome families in both current suborders. Moreover, Bazinga
rieki gen. nov., sp. nov. is unique among all rhizostomes in its very small mature size. It is
probable that this species has been previously mistaken for juveniles of other species. ?
Scyphozoa, Rhizostomeae, Bazingidae, blubber jelly, new species, taxonomy.
Rhizostome jellyfish are typically the most
obvious of all gelatinous life-forms, often being
large and brightly coloured, and often having
bizarre exumbrellar morphology or conspic -
uous trailing clubs and filaments. Most rhizo -
stomes are relatively large, and certain edible
species support a significant commercial fishery.
The biggest in the group is the infamous 2 m
wide, 200 kilogram Nemopilema nomurai Kishi -
nouye, 1922, which has been plaguing Japanese
fishing grounds by the billions, particularly
since 2000 (Kawahara et al. 2006). While there
are a few rhizostomes in the 35–50 mm range,
such a small size is exceptional.
The present work documents an intriguing
little jellyfish that is particularly striking by
reaching maturity at below 20 mm. It is also
significantly different from other rhizostomes
by its unusual combination of morphological
characters, and by having a suite of unique
features that have not otherwise been observed
in any rhizostome family or suborder.
While Bazinga rieki gen. nov., sp. nov. is likely
to be relatively common, it has probably been
overlooked because its small size makes it
easily mistaken for a juvenile of other species.
In particular it is superficially similar to the
common Catostylus mosaicus (Quoy & Gaimard,
1824), and similar sized juveniles of that species
were also collected at the same time at one
MATERIALS AND METHODS
Specimens were fixed in a solution of about
5% concentrated formalin in seawater. Measure -
ments were made on preserved specimens with
digital calipers. Character evaluations were
made on preserved specimens examined under
a dissecting scope and from photographs and
video of live specimens.
Peripheral portions of the radial canals were
studied by dissecting away the subumbrellar
circular muscles and exumbrellar warts.
All type material has been deposited in the
Queensland Museum. Abbreviations: QM,
Queensland Museum; BD, bell diameter; RC,
Memoirs of the Queensland Museum — Nature ? 2013 ? 56(2) ? www.qm.qld.gov.au 625
Citation: Gershwin, L. & Davie, P.J.F. 2013 06 30. A remarkable new jellyfish (Cnidaria: Scyphozoa) from
coastal Australia, representing a new suborder within the Rhizostomeae. Memoirs of the Queensland Museum
— Nature 56(2): 625–630. Brisbane. ISSN 0079–8835.
Order RHIZOSTOMEAE Cuvier, 1799
Ptychophorae suborder nov.
Diagnosis. Body globular. Oral arms coalesced
into a single short, ridged column; without
scapulets. Rhopalia hooded, lacking typical
pits. Velar lappets 4 per octant; 2 asymmetrical
ocular lappets per octant. Annular muscles
conspicuous. Subgenital ostia very small,
round. Stomach circular, large. Radial canals 4
per octant, proximally unbranched, fluted;
peripherally coalesced into vast open sinus
with patchwork of jelly matrix.
Etymology. From the Greek ptychos (fold, leaf,
layer) and phoras (bearing), in reference to the
Bazingidae fam. nov.
Diagnosis. As for the suborder.
Bazinga gen. nov.
Type species. Bazinga rieki sp. nov. by present
Diagnosis. As for the suborder.
Etymology. The name Bazinga has been confer -
red for two reasons. Firstly, bazinga is a slang
term in present popular culture, meaning
'fooled you!' or 'stung you!' (e.g., the fictional
Dr Sheldon Cooper in the television series The
Big Bang Theory), and this is appropriate as the
type species, B. rieki, is so small that it has
probably been overlooked in the past as a
juvenile of a larger species. Secondly, a seven-
string harp is also called a bazinga (Cuppy
1950), and the straight radial canals of this new
species are reminiscent of such strings.
Bazinga rieki sp. nov.
(Figs 1, 2)
Material examined. HOLOTYPE: QM-G331996, male
(18.7 mm BD), Brunswick River, NSW, high tide in
shallow water, Denis Riek, 04.12.2011.
PARATYPES: QM-G331997, male (15.5 mm BD),
gravid female (14.0 mm BD), Cudgen Creek, Hastings
Point, NSW, high tide, under road bridge, Denis
Riek, 16.11.2011. QM-G331998, gravid female (16.4
mm BD), male (13.7 mm BD), Seagull Rocks,
Brunswick Heads, NSW, rock pools, low tide, Denis
Description. Body thick, globular, rounded,
tending to flattened above; small, reaching
maturity at less than 20 mm BD (Fig. 1A).
Exumbrella without any form of central hump
or papillae clusters. Entire aboral surface evenly
and densely covered in minute warts; each
minute wart resembles small spherical sessile
tag with yellowish core, possibly filled with
zooxanthellae. Peripheral region of bell turned
vertically downward parallel to body axis.
Oral arms (Fig. 1B) coalesced into single short
column, ridged along column wall; arms
formed into narrow folded sheets with
mouthlets arranged along distal margin. Sheets
contain scattered zooxanthellae throughout.
Column wall and arm sheets finely granulated
with many minute warts. Filaments, clubs,
knobs, and other appendages lacking (Fig. 1F).
Subumbrellar surface (Fig. 1C) evenly covered
with fine warts from oral arm column to just
before edge of stomach; warts mostly longer
proximally and rounder and smaller distally.
Gastrogonadal cavity open to outside only at 4
small, round, perradial holes (Fig. 1D) located
on subumbrellar portion of stomach wall very
close to margin of stomach, about midway
between oral arm column and proximal muscle
bands. Circular muscle bands numbering about
12; in form of broad flat lamellae; richly impreg -
nated with zooxanthellae.
Stomach circular in outline, large, occupying
more than half body diameter; margin plainly
visible through subumbrellar wall. Stomach
extends via broad, evenly spaced, irregular
flute-shaped radial canals, 4 per octant;
peripherally coalesced into a broadly open
circular sinus, punctuated with an irregular
patchwork of jelly matrix (Fig. 2A). All radial
canals arise at same level and are of same
length. No radial canals reach bell margin, and
no specifically differentiated rhopalial canal;
radial canals dissipate in circular sinus. Proximal
straight portions of canals clearly visible in live
animal (Fig. 1E), but precise nature of periph -
eral gastrovascular system otherwise obscured
by surface warting and subumbrellar muscle
bands. Defined ring canal lacking.
Gonads comprised of heavily folded tissue
sheets, plainly visible through subumbrellar
wall as dark masses rich with zooxanthellae.
626 Memoirs of the Queensland Museum — Nature ? 2013 ? 56(2)
Gershwin & Davie
Memoirs of the Queensland Museum — Nature ? 2013 ? 56(2) 627
A remarkable new jellyfish
FIG. 1. Bazinga rieki gen. nov., sp. nov., in life. A, habitus; B, oral arms; C, subumbrellar view; D, gastro-
gonadal pores; E, bell on power stroke (note straight radial canals visible through bell); F, bell relaxed.
Velar lappets 4 per octant, short, squared off,
not as long as ocular lappets, defined from one
another proximally by narrow region without
warts. Ocular lappets two per octant, long and
pointed, asymmetrical in preserved specimens,
with right-hand lappets in ventral view folding
to point obliquely along bell margin, left-hand
lappets folding axially (Fig. 2B); bases of right
and left ocular lappets folded axially to create a
straight-sided gap between, slightly rounded
marginally (Fig. 2C). All lappets heavily
granulated with fine warts.
Rhopalia 8; 4 perradial and 4 interradial;
deeply embedded under exumbrellar hood, in
cavity formed by large fold on axial side of each
ocular lappet (Figs 2B, C). Typical rhizostome
exumbrellar smooth or radially ridged,
rhopaliar pits lacking.
Colour in life. Bell jelly translucent and
colourless; subumbrellar muscle folds golden
brown due to zooxanthellae; gonads darker
brown; after 2–3 months preserved in formalin
solution, whole animal pale yellowish with
Variation. Smaller paratypes have the radial
canals in the shape of a simple broad straight
line, without the flared sides of the holotype;
mid-size specimens have the sides of the canals
somewhat more divergent like a funnel.
Observations on live animals. Video taken by
Denis Riek in both natural and laboratory
settings accompanies the type series. The
swimming pattern consists of continuous rapid
pulsations at a rate of more than 200 per
minute, and even though the bell is being
continually flexed, there appears to be little
effective propulsion. In an aquarium these
pulsations were insufficient to lift it off the
substrate. Thus it would seem that Bazinga rieki
is largely at the mercy of currents.
Etymology. The species name, rieki, is to
honour Mr Denis Riek, whose keen interest in
photographing and identifying the sea-life of
northern New South Wales led to the discovery
of this most intriguing species.
Distribution. Bazinga rieki is only known from
coastal waters off northern New South Wales.
However, having been found at three different
collecting sites on three different dates it seems
likely to be relatively common, and could be
expected to be found more widely along the
central east Australian coast, particularly south
of the Tropic of Capricorn.
The order Rhizostomeae was divided into two
suborders by Stiasny (1921), based on a suite of
structural features. Members of the Dactylio -
628 Memoirs of the Queensland Museum — Nature ? 2013 ? 56(2)
Gershwin & Davie
FIG. 2. Bazinga rieki gen. nov., sp. nov. A, peripheral
canal sinus with subumbrellar muscles dissected
away; B, subum brellar view of rhopaliar niche (note
asymmetrical lappets); C, exumbrellar view of rhopaliar
niche (note protective hood in place of open pit).
phorae are characterised by having radial folds
on the surface of the rhopaliar pits; subgenital
pits narrowed by conspicuous papillae;
annular subumbrellar muscles; oral arms that
are three-winged and may have scapulets; and
by the anastomosing network of canals not
being in communication with the gastric cavity.
In contrast, the Kolpophorae has smooth-
surfaced rhopaliar pits; subgenital ostia without
papillae; muscles variable in form from radial
and straight to annular to feather- like arcs;
anastomosing canals in communication with the
gastric cavity; and oral arms that lack scapulets
and are dichotomous, triangular, or three- winged.
The primary diagnostic features for both sub -
orders are summarised and compared in Table 1.
Bayha et al. (2010) constructed a phylogeny
for the scyphozoan families using sequence
data from 18S and 28S rDNA nuclear genes.
Their results were largely consistent with prior
morphological hypotheses, but importantly they
concluded that the dactyliophorids were para -
phyletic with respect to the kolpophorids. Since
Bazinga falls well outside both of these groups
morphologically, it seems likely that it is either
the sister group to, or basal to, all other rhizo -
stomes. Moreover, its hooded rhopalia are more
reminiscent of some semaeostomes than of the
rhizostomes, again suggesting a basal affinity.
Obtaining DNA and testing these hypotheses
should be considered a high research priority.
Rhizostomes are famously large, some reach -
ing diameters of 350–500 mm or even more, and
while there are a few small species in the size
range 35–50 mm, e.g., Cassiopea ndrosia Agassiz
& Mayer, 1899 (50 mm, Fiji), Mastigias gracilis
(Vanhöffen, 1888) (35 mm, Red Sea), and
Acromitus tankahkeei Light, 1924 (45 mm, China),
this is unusual. However, Bazinga rieki is mature
at less than 20 mm, making it by far the smallest
rhizostome yet discovered. While this is note -
worthy in itself, Bazinga rieki is also unique in its
morphology, differing in major ways from all
The ocular lappets of Bazinga rieki are remark -
able in being asymmetrically folded, and this is
consistent across all specimens. Such an occur -
rence has apparently not been described in any
The unusual canal system is also utterly
unlike any other rhizostome, and can only be
awkwardly described using standard termin -
ology. In most rhizostomes, the radial canals
arise at different levels along the cross-shaped
stomach, with those arising from the interradii
(i.e., the arms of the cross) being the shortest,
those arising from the perradii (i.e., between
the arms of the cross) being the longest, and
those in between being midway in length. In
Bazinga the arms are all the same length because
they arise from the same level off the circular
stomach. Moreover, rhizostomes are taxonomi -
cally separated based on a) how many radial
canals extend past the ring canal to reach the
margin, and b) whether the anastomosing net -
work communicates with the stomach. However,
Bazinga has no ring canal, and no radial canals
extend to the margin, but rather, the coelenteric
region covered by the circular muscles consists
of a vast open sinus with scattered small
patches of jelly matrix, giving the impression of
a reversed meshwork, where the swollen and
coalesced area of the anastomosed canals is far
greater than the area between.
If one looks at individual families within the
Rhizostomeae, irrespective of suborder, then
Bazinga rieki does share some important char -
acters with particular families. Like Cassio -
peidae, the stomach of Bazinga is circular;
however, in the only described cassio peid
genus, Cassiopea, it is very small compared to
the whole diameter, whereas in Bazinga it is
very broad, spanning more than half the bell
diameter. Bazinga further differs from Cassiopea
in all other primary diagnostic characters such
as the number of rhopalia, the form of the
subumbrellar muscles, the form of the oral
arms, the form and number of the radial canals,
and the general habits of the live animal.
Bazinga is reminiscent of Cepheidae in having
small subgenital ostia; however, the Cepheidae
is characterised by a large central dome on the
exumbrella which often bears conspicuous
papillae or sometimes a large knob. Even in
Cotylorhiza, which lacks papillae or a knob, the
central dome is well defined and dominates the
bell. Bazinga has no such central dome; instead,
the bell is globular and often somewhat flat -
tened in life. Species in the Cepheidae are also
characterised by having appendages on the oral
arms such as filaments, stalked suckers, or
spindles, whereas Bazinga completely lacks any
Memoirs of the Queensland Museum — Nature ? 2013 ? 56(2) 629
A remarkable new jellyfish
such appendages. Furthermore, the oral arms
of Bazinga are coalesced whereas those of
cepheids are not. Finally, the conspicuous
annular muscles of Bazinga compared to the
fine radial muscles of the cepheids, the circular
stomach of the former compared with the octag -
onal stomach of the latter, and the completely
different radial canal pattern between the two
groups convincingly separates them.
Like some Kolpophorae families such as
Mastigiidae, Versurigidae, and Thysanostomidae,
Bazinga has well developed annular muscles on
the subumbrella. However, along with many
other structural features, the continuous sub -
genital porticus and separate oral arms of these
other families would serve to immediately
distinguish them from Bazinga.
Similarly, the Dactyliophorae families have
annular muscles, but they have conspicuous
papillae in the subgenital ostia, whereas Bazinga
Even the families with coalesced oral arms
would not be easily mistaken for Bazinga, with
both the Rhizostomatidae and Stomolophidae
also having scapulets, whereas Bazinga does not.
One of the key features that separates the two
orders, Kolpophorae and Dactyliophorae, is the
surface texture of the horseshoe-shaped rhopaliar
pits, being smooth in the former and decorated
by radial folds in the latter. However, the
rhopalia of Bazinga are completely different,
i.e., completely covered by an exumbrellar
hood and embedded in a subumbrellar cavity
formed by the folded lappets.
Therefore, while Bazinga does share some
features with other Rhizostomeae suborders
and families, its large circular stomach,
unbranched evenly spaced radial canals, and
hooded rhopalia are unique, and warrant new
subordinal status. This decision is further
reinforced by its unusual combination of other
primary diagnostic characters such as coalesced
oral arms and lack of scapulets and subgenital
papillae, that also make it wholly unlike any
other known family or suborder.
We are indebted to Denis Riek for bringing
this splendid little medusa to our attention. We
sincerely thank Michelle Baker for her out -
standing artwork, and Wolfgang Zeidler and
Dale Calder for helping improve the manu -
script in review. Travel costs for the senior
author to visit the Queensland Museum were
partially supported by travel grants from the
Asperger’s Services Australia and the Common -
wealth Scientific and Industrial Research
Agassiz, A. & Mayer, A.G. 1899. Acalephs from the
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M.S., & Haddock, S.H.D. 2010. Evolutionary relation -
ships among scyphozoan jellyfish families based
on complete taxon sampling and phylogenetic
analyses of 18S and 28S ribosomal DNA.
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stomeae) in East Asian waters. Marine Ecology
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630 Memoirs of the Queensland Museum — Nature ? 2013 ? 56(2)
Gershwin & Davie
Gershwin, L. & Davie, P.J.F. 2013 06 30. A remarkable new jellyfish (Cnidaria: Scyphozoa) from coastal Australia, representing a
new suborder within the Rhizostomeae. Memoirs of the Queensland Museum — Nature 56(2): 625–630.
The following table was omitted in error from the original printed paper
Memoirs of the Queensland Museum — Nature ? 2013 ? 56(2)
A remarkable new jellyfish
SUBUMBRELLAORAL ARMSRADIAL CANALSSTOMACH
annular, or in
laterally into a
Table 1. Comparison of diagnostic structural features separating the suborders of Rhizostomeae, synthesised
primarily from: Mayer (1910), Stiasny (1921, 1923), and Kramp (1961).