Description of a new midwater medusa, Tiaropsidium shinkaii n. sp. (Leptomedusae, Tiaropsidae)

2005·1.4 MB·

PlanktonBiol. Ecol. 52(2): 100-106,2005
plankton
biology & ecology
<: The Plankton Society ofJapan 2005
Description of a new midwater medusa, Tiaropsidium
shinkaii n. sp. (Leptomedusae, Tiaropsidae)
MlNORU KlTAMURA, DHUGAL J. LINDSAY & HlROSHI MlYAKE
MarineEcosystem Research Group,JapanAgencyforMarine-Earth Scienceand Technology(JAMSTEC), 2-15Natsushima-cho,
Yokosuka, Kanagawa237-0061, Japan
Received20 February2004; Accepted 15 June2005
Abstract: A new species, Tiaropsidium shinkaii (Leptomedusae, Tiaropsidae) is described based on
a single specimen collected by submersible from the midwater zone of Sagami Bay, Pacific coast of
Japan. The presence of compound sense organs, four radial canals, and two kinds of tentacles indi
cate thatthe presentspecimen is a member ofthe genus Tiaropsidiumwithin the family Tiaropsidae.
Of the currently described Tiaropsidium species, the present specimen resembled T. roseum most
closely as the number of long tentacles was four. However, it was distinguishable from T. roseum by
(1) deep umbrella, (2) thick jelly, (3) sinusoidal gonads located on distal portion of radial canals, (4)
presence of baso-abaxial projection on each tentacle bulb, and (5) black pigmentation in interradial
portion of innerwall ofstomach. This is the seventh species in the genus Tiaropsidium.
Key words: new species, Tiaropsidium shinkaii, Leptomedusae, Tiaropsidae, midwater
servational recordoffeeding byAtolla sp. (Hunt& Lindsay
Introduction
1998). Taxonomic reports on deep sea jellyfishes around
Since the introduction ofcrewed submersibles, remotely Japan are mainly old papers based on plankton net samples
operated vehicles (ROV) and other sampling devices, it has (e.g. Bigelow 1913, Uchida 1928, 1947), but recently Mat-
become apparent that the species diversity ofmid and deep sumoto et al. (2003) described a new midwater scyphome-
water jellyfishes is high. Pugh described many new dusa from studies using submersibles.
siphonophore species based on specimens collected using In the present study, we describe a new species ofmid
submersibles (e.g. Pugh & Harbison 1987, Pugh & Young- water leptomedusa collected using the submersible Shinkai
bluth 1988). In the Atlantic, Larson et al. (1991) reported 2000, from southeast of Hatsushima Island, Sagami Bay,
the presence of many undescribed species from observa Pacific coast ofJapan. Crewed submersibles and ROVs are
tions using a submersible. Recently, Gili et al. (1998, 1999) effective platforms not only fortaxonomic but also foreco
described several new species collected by moored sedi logical studies. For ecological studies, identification of
ment traps in the Mediterranean Sea. The manuscripts of species through observation rather than collection is often
Pugh et al. illustrate the effectiveness ofsubmersibles as de necessary. Hence, we also describe the macro-morphology
vices enabling taxonomic studies on midwater cnidarians. and swimming behavior of the present species in situ as a
In order to examine a suite ofmorphological characters in guide to species identification in the field.
detail, selective and individual sampling ofsuch fragilejel
lyfishes isthe ideal.
Materials and Methods
Submersible-based studies onjellyfishes in Japanese wa
ters have been mostly observational records (Peres 1959, A living specimen was collected from southeast of Hat
Toyokawa et al. 1998, Lindsay et al. 1999, 2004, Miyake et sushima Island, Sagami Bay (35°00'N, 139°14'E), on No
al. 2002), methodology for faunal studies with associated vember 9, 2002. Sampling was achieved using a suction
observational records (Hunt & Lindsay 1999), and an ob- sampler (Hunt et al. 1997) from the crewed submersible
Shinkai 2000, operated by the Japan Marine Science &
Technology Center (JAMSTEC). The dive number was
Corrcspondingauthor: Minoru Kitamura; e-mail, kitnmura(W-Jamstec.go.jp;
Tel: -81-46-867-9527; lax: I X1-46-S67-9525 2K1409, and the observer was M. Kitamura. Bottom depth
Description of Tiaropsidiumshinkaiin. sp. 10]
at the dive point was 1200m, and the depth of collection
was 439m. Before collection, the swimming behavior of A
the present specimen was observed directly and recorded in
situ on Sony Digital Betacam videotapes, which are stored
at the Computer and Information Department, JAMSTEC
{Dive 2K.1409, i/6-6/6). Physico-chemical environmental
parameters at the sampling depth were determined using a
ScaBird SBE19 CTD attached lo the submersible on the
dive. After collection, the specimen was fixed and pre
served in a 5% buffered formalin-seawatersolution. Forne
matocyst observations, the distal part ofa long tentacle was
cut offbefore fixation anda squash preparation ofthe tenta
cle was made. Preservation liquid for the nematocyst prepa
ration was 5% formalin/5% glycerin-seawater. Morphologi
cal observations and measurements of the medusan body
and nematocysts were carried out under a dissecting and a
phase-contrast microscopes, respectively. Terminology used
in this study follows that of Bouillon (1999) and Mariscal
(1974).
Taxonomic Description
Family Tiaropsidae Boero, Bouillon & Danovaro, 1987 B
Genus Tiaropsidium Torrey, 1909
Tiaropsidiumshinkaii n. sp.
(Figs. 1-6)
Material examined. Holotype, collected from Sagami
Bay (35°00'N, 139°14'E), 439m depth, on Nov. 9, 2002.
The holotype is deposited in the National Science Museum,
Tokyo(NSMT-Co1406).
Description. Umbrella deep (Fig. IA), 24.3 mm in
width and 24.8mm in height. Jelly tender, thick at apex but
thin at margin, 3.2mm and 0.6mm respectively. Four radial
canals straight and narrow (Fig. 2). Manubrium short, about
one fifth ofumbrella height; attached to each of lour radial
canals (Fig. 3A); cruciform in transverse section, with four
recurved lips. Length of attached portion about equal to
half of manubrium height. Stomach pointed centrally at
Fig. I. Tiaropsidium shinkaii n. sp. A: Preserved specimen, in
base. Color of outer surface of manubrium white; interra- the lateral view. 13: Live specimen, filmed from Shinkai 2000(439
dial portion ofinner wall of stomach black (Fig. 3B). Nu m depth).
merous horizontal furrows running along inner wall of
stomach. Gonads, white, elongated and sinusoidal, about
halflength ofradial canals, located on distal halfof radial tentacles not coiled, with slightly wider bases; all similar in
canals but not reaching umbrella margin. Two kinds often length, from 0.4 to 0.6mm. Number oftiny tentacles: three
tacles present, four long and 62 tiny. Former located perra- to five between tentacle bulbs and sense organs, seven to
dially and coiled two or three times at base in preserved nine between successive sense organs, 14 to IS in total in
condition. Each oflong tentacles with large, globular tenta each quadrant Eight compound sense organs, two m each
cle bulb, with associated small baso-abaxial projection quadrant, located on the velar side ofthe umbrella margin.
(Fig. 4A). Tentacle bulbs with Iwo brownish-pigmented Each sense organ with open vesicle and ocellus. Each vesi
areas inside (Fig. 4A). At perradial umbrella margin, jelly cle, a small invagination of velum, opening toward um
projected downward (Fig. 4A, B). Each tentacle bulb lo brella cavity (Fig. 6A-C). Ocellus located on outer side of
cated on adaxial side ofjelly projection (Fig. 4B). Only one vesicle, round in lateral view, while round, triangular or
type ofnematocyst (microbasic mastigophore: Fig. 5) rec square with rounded corners in oral view, black in color.
ognized on long tentacles. Undischarged capsules ellipsoid Ring canal narrow. Velum not overly developed. 1.8mm in
in shape, llftm in length and 2.1—2.8/im in width. Tiny width.
102 M. Kitamura, D. J. Lindsay& H. Miyake
Typelocality. Sagami Bay. ated by the Japan Marine Science & Technology Center
Geographical distribution. Known only from the type (JAMSTEC). Shinkaialsomeans 'deep sea' in Japanese.
locality. Habitat. The major water mass at the depth ofcollec
Etymology. The specific name,shinkaii, is derived from tion was classified as Intermediate Kuroshio Water (IKW;
the name ofthe crewed submersible 'Shinkai 2000* oper Fujimura & Nagata 1991), a subset ofNorth Pacific Inter-
5mm
Fig. 3. Tiaropsidium shinkaii n. sp. Manubrium in the lateral
Fig. 2. Tiaropsidium shinkaii n. sp. Lateral view. Scale
view(A)andtheoralview(B). Scalebar=5mm.
bar=1cm.
tb
5 mm
Fig. 4. Tiaropsidium shinkaiin. sp. Umbrella margin neartentacle bulb in the lateral view (A) and tentacle bulb in the lateral view (B).
Velum isnotdrawn in Fig. 4A. Abbreviations: dpj,downwardprojectionofjelly; cso, compoundsensoryorgan; g, gonad; It, longtentacle;
p,baso-abaxial projectionoftentaclebulb; rac,radial canal;ric,ringcanal;tb,tentaclebulb;tt,tinytentacle;vel, velum. Scalebar=5mm.
Description of Tiaropsidiumshinkaiin. sp. 103
mediate Water (NPIW; Talley 1993). The collection depth
(439m) was located within the salinity minimum layer
formed by the IKW. Physico-chemical environmental para
meters at the collection depth were as follows: temperature,
6.6°C; salinity, 34.29; sigma-t, 26.91.
Macro-morphology and swimming behavior in situ as a
guide to species identification in thefield. The umbrella
was deep, as high as wide. The manubrium, four gonads,
four tentacle bulbs and four long tentacles were prominent
and whitish (Fig. 1B). The manubrium had a short stomach
with slightly recurved lips. The gonads were sinusoidal and
located on the distal halfofthe radial canals. The proximal
portions of the gonads were narrower. The tiny tentacles
and compound sense organs were not recognizable in the
NTSC video record from the Shinkai 2000. Swimming of
the present specimen was slow and continuous with bell
pulsing at0.6Hz. Contraction oftheumbrella during swim
ming occurred primarilynearthe margin. When the medusa
Fig. 5. Tiaropsidium shinkaii n. sp. Nematocyst from long ten was swimming, the long tentacles were very extended, 13
tacle. Scalebar=10fim. times as high as the umbrella height, and were dragged be
hindthe medusa.
Discussion
The present specimen has compound sense organs, each
of which is comprised of an ocellus and a statocyst (only
open vesicles were recognized in the present specimen), in
dicating that it belongs to the family Tiaropsidae, which in
cludes three genera, Tiaropsis, Tiaropsidium, and Octogo-
nade. Of the three genera, Tiaropsis has only one type of
tentacle, while the others have two types, longtentacles and
rudimentary ones. One ofthe characters distinguishing be
ves o tween the genera Tiaropsidium and Octogonade is the num
ber ofradial canals. Species included in the former genus,
vel except T.polyradiatum, have four canals while those in the
lattergenus have eight. Thepresentspecimen has twotypes
of tentacles, and four radial canals, relegating it to the
genus Tiaropsidium.
At present, six species in the genus Tiaropsidium have
been described, T. atlanticum Russell, 1956, T.japonicum
Kramp, 1932, T. kelseyi Torrey, 1909, T. mediterraneum
(Metschnikoff, 1886), T.polyradiatum Kramp, 1965, and T.
roseum (Agassiz & Mayer 1899) (see Kramp 1961, Boero
et al. 1987, Bouillon & Boero 2000). Morphological differ
ences between Tiaropsidium species, including our present
vel specimen, are shown in Table 1. The number oflong tenta
cles is one ofthe more important characters for distinction
of Tiaropsidium species. The present material resembles T.
roseum because ofthe presence offour long tentacles, but
has the following different characters: (1) a deep umbrella,
1 mm
(2) thickjelly at the apex, (3) sinusoidal gonads located on
the distal part ofthe radial canals, (4) baso-abaxial projec
Fig. 6. Tiaropsidiumshinkaiin. sp. Compound sense organ and
tionoftentaclebulbs andperradial gelatinous projections at
row oftiny tentacles inthelateral view(A), the oral view (B) and
schematic drawing ofopeningofthe vesicle viewed from the um umbrella margin near the tentacle bulbs, (5) black pigmen
brellacavity (C). Abbreviations: o, ocellus; oves, opening ofvesi tation in interradial portion of inner wall of stomach. As
cle;ric,ringcanal;tt,tinytentacle; ves,vesicle. Scalebar=1mm. concerns (1), although the specimens of T. roseum de-
Table 1. Comparisonsbetweenthedifferentspeciesofthegenus Tiaropsidium.
Numberoflong Numberof Numberof
Size in tentaclesand tinytentacles compound
Species Umbrella Gonads Color Distribution References
mm shapeof ineach sense
tentaclebulbs quadrant organs
Tiaropsidiumatlanticum Flatterthan 60wide Probably24,with Probably 18 Linear,alongmiddle Probably Interradialwallsof EnglishChannel; 1
Russell, 1956 hemisphere elongatedswollen 3/4ofradialcanals about48 stomachand long dwellingdepth
with fairly bulbs tentaclesblack unknown
thickjelly
Tiaropsidiumjaponicum Watch-glass 18 wide 8,with largeand 12-14 Linear,alongalmost 16 Unknown Sagami Bay,eastof 2,3
Kramp, 1932 shapedwith swollenbulbs entire length ofradia1 CookStrait; midwater
thinjelly canals
Tiaropsidiumkelseyi Somewhat 50wide 8, withelongated About 10 Curtain-like, much 8 Manubrium, canals, WestcoastofNorth 4
Torrey, 1909 conical,three bulbs folded,alongentire gonadsand tentacles America;midwater
timesasbroad lengthoftheradial faintyellow
ashigh canals 2
C
Tiaropsidium Globularwith 5 high. 2 oppositeperradially, 5 Elongated,along 8 Gonads, stomachand Messina; dwelling 5 3)
mediterraneum thickjelly 7 wide another2perradial distal 2/3 ofradial tentacle bulbs depth unknown p
(Metschnikoff, 1886) portionsofumbrella canals yellowish-grey
margin hadsmalt bulbs 1
Tiaropsidium Flattenedwith 30wide Apparently24—32,no Morethan2 Alongentire length More Unknown Nicobars;dwelling 3
-<
polyradiatum thinjelly descriptionabout ofradial canals than8 depthunknown 9?
Kramp, 1965 bulbs X
Viaropsidiumroseum Flattenedwith 15wide 4,withbroadbulbs 7 Elongatedoval, near 8 Entodermofstomach FijiIslands, Malayan 3,6-12
(Agassiz&Mayer, thinjelly stomach, somewhat ocher-yellowto Archipelago,Mauritius,
m
1899) longerthan 1/3 of dull-red Nicobars,northof
radial canals PapuaNew-Guinea,off
Sydney,castofNew
Zealand, Benguela
Current,California;
surfacelayer
Tiaropsidiumshinkaii Deep,jelly 24.8 high, 4,eachbulbwith 13-19 Sinusoidal,distalpart 8 Outerwall of SagamiBay;midwater Present
n. sp. thickatapex 24.3 wide baso-abaxial ofradial canals manubrium,gonads. (439m) study
projection tentacleswhite;
interradial portionsof
innerwall ofstomach
black
References: 1) Russell (1956); 2) Kramp (1932); 3) Kramp(1965); 4) Torrey (1909); 5) Metschnikoff(1886); 6) Agassiz & Mayer(1899); 7) Maas (1909); 8) Brown(1916); 9) Boero,
Bouillon& Danovaro(1987); 10)Bouillon & Barnett(1999); 11) Pages,Gili & Bouillon(1992); 12)Alvarifio& Kimbrell(1987)
Descriptionof Tiaropsidiumshinkaiin. sp. 105
scribed by Agassiz & Mayer (1899), Brown (1916), Kramp script. We also thank Drs. Hiroyuki Yamamoto, Katsunori
(1958) and Boero et al. (1987) had a hemispherical um Fujikura, Shinji Tsuchida, and Yoshihiro Fujiwara for their
brella, theyweretooyoungtobecompared with thepresent comments and advice. Comments of two anonymous re
material. These specimens were 2.5mm in umbrella height, viewers helped to improve the manuscriptsignificantly.
3mm, 5-8mm and 1.5mm in umbrella diameter, respec
tively.
Literature Cited
Tiaropsidium roseum has previously been collected from
surface waters in the Indian and Pacific Oceans, Malayan Agassiz, A & A. G. Mayer 1899. Acalephs from the Fiji Islands.
Archipelago (surface, Maas 1905), Mauritius (128-0 fath Bull. Mus. Comp. Zool. Han'ardColl.32: 157-189. pis. 1-17.
oms, Brown 1916), Nicobars (surface, Kramp 1958), Alvarino, A. & C. A. Kimbrell 1987. Abundance ofzooplankton
Benguela Current (93-0m, Pages et al. 1992), Fiji (surface, species in California coastal waters during April 1981, Febru
Agassiz & Mayer 1899), a coralline island off the north ary 1982, March 1984, March 1985. NOAA Technical Memo
coast of Papua New-Guinea (Boero et al. 1987), and Cali randum NMFS, NOAA-TM. NMFS. SWFC. 74: 1-59.
fornia (70-0m, Alvarino & Kimbrell 1987). There have Bigelow, H. B. 1913. Medusaeandsiphonophoraecollectedbythe
been two otherreports ofcollection from the Pacific Ocean U. S. fisheries steamer 'Albatross' in the north-western Pacific,
(off Sydney, 3000m wire out, Kramp 1965; east of New 1906.Proc. U. S. Nat. Mus. 44: 1-119.
Zealand, sampling details unknown, Bouillon & Barnett Boero, F., J. Bouillon & R. Danovaro 1987. The life cycle of
Tiaropsidium roseum (Tiaropsidae fam. nov., Leptomedusae,
1999), but the exact depths of collection are unclear. Al
Cnidaria).Indo-MalaycmZool. 4: 293-302.
though there are two records from ambiguous depths, it is
Bouillon, J. 1999. Hydromedusae, p. 385-465. In South Atlantic
probably safe to assume that T. roseum is a surface species.
Zooplankton, vol. /, (ed. D. Boltovskoy). Backhuys Publishers,
The present material, however, was collected from 439m
Leiden,The Netherlands.
depth. The black pigmentation in the stomach wall, which
Bouillon, J. & T. J. Barnett 1999. The marine fauna of New
may function to mask any bioluminescence produced by in
Zealand: Hydromedusae(Cnidaria: Hydrozoa). NIWA Biodiver
gested prey, allows us to infer that the present species is a
sityMem. 113: 1-136.
true member ofthe midwater fauna. Among the previously
Bouillon, J. & F. Boero 2000. Synopsis ofthe familiesandgenera
described species of the genus Tiaropsidium, only T. at-
ofthe hydromedusae ofthe world, with a list ofthe worldwide
lanticum that probably dwells the midwater had black pig
species. ThalassiaSalentina24: 47-296.
mentation inthe stomachwall.
Brown, E. T. 1916. Medusae from the Indian Ocean. Trans. Linn.
Uchida (1965) described Tiaropsis rosea (=T. roseum) Sot: London (Zool.) 17: 169-211.
collected from Misaki, Sagami Bay, but his description and Fujimura, M. & Y. Nagata 1991. Mixing process in the mixed
figure seems only to be a quotation of Mayer (1910). We water and Kuroshio extension regions and modification ofthe
hesitate to classify his specimen as T. roseum because his Intermediate Kuroshio Water. UmitoSora67: 75-84.(inJapan
description was very briefand the material has been lost. esewithEnglishabstract)
Kramp (1932) described T. japonicum based on two Gili, J. M., J. Bouillon & F. Pages 1998. A new species ofKram-
specimens collected from Sagami Bay, at about 600m pella (Hydrozoa, Hydroidomedusae, Tiarannidae) from the
depth. Although both our present specimen and T. japon deep waters of Antikythira Strait (Cretan Sea, North East
icum were collected from the midwater layer of Sagami Mediterranean).Sci. Mar. 62: 135-139.
Bay, the former is easily distinguishable from the latter by Gili, J. M., J. Bouillon, F. Pages, A. Palanques & P. Puig 1999.
having (1) eight compound sense organs, (2) four long ten Submersible canyons as habitats of prolific plankton popula
tacles, (3) a deep umbrella, and (4) sinusoidal gonads lo tions: three new deep-sea Hydroidomedusae in the western
Mediterranean.Zool. J. Linn. Soc. 125: 313-329.
cated on the distal half of the radial canals. The present
Hunt, J. C, J. Hashimoto, Y. Fujiwara. D. J. Lindsay, K. Fujikura,
specimen is 24.3mm in width, while Kramp's T.japonicum
S. Tsuchida & T. Yamamoto 1997. The development, imple
materials were nearly the same size, 18mm. The above-
mentation, and establishment of a Meso-pelagic and Bentho-
mentioned morphological differences are, therefore, consid
pelagic biological survey program using submersibles in the
ered to be interspecificratherthan ontogenetic.
seasaroundJapan.JAMSTECJ. DeepSeaRes. 13: 675-685.
From the above-mentioned morphological and distribu
Hunt, J. C. & D. J. Lindsay 1998. Observations onthe behaviorof
tional differences, we conclude that a new species in the
Atolla (Scyphozoa: Coronatae) and Nanomia (Hydrozoa:
genus Tiaropsidium should be established to accomodate
Physonectae): use ofthe hypertrophiedtentacle in preycapture.
thepresent material.
Plankton Biol. Ecol.45: 239-242.
Hunt,J. C. & D. J. Lindsay 1999. Methodology forcreatinganob
Acknowledgements servational database ofmidwater fauna using submersibles: re
sults fromSagami Bay, Japan. PlanktonBiol. Ecol. 46: 75-87.
We sincerely wish to thank the captain and crew of the Kramp, P. L. 1932. A revision ofthe medusae belonging to the
R/V Natsushima and the operations team of the Shinkai family Mitrocomidae. Vidensk Medd. j'ra Dansk naturhist.
2000 fortheir dedicatedefforts. We thank Drs. Shin Kubota Foren. Kjobenhavn92: 305-384.
and Takashi Okutani fortheircritical reviews ofthis manu Kramp, P. L. 1958. Hydromedusae inthe Indian Museum. Rec. In-
106 M. Kitamura, D.J. Lindsay& H. Miyake
dianMus. 53: 339-376. waters off Kushiro, Hokkaido Japan. Plankton Biol. Ecol. 49:
Kramp, P. L. 1961. Synopsis ofthe medusae ofthe world../. Mar. 44-^6.
Biol.Ass. U. K.40: 1-469. Pages, F, J. M. Gili & J. Bouillon 1992. Planktonic cnidarians of
Kramp, P. L. 1965. The hydromedusae ofthe Pacific and Indian the BenguelaCurrent.Sci. Mar. 56(suppl. 1): 1-64.
Oceans.DanaRep. 63: 1-161. Peres, J. M. 1959. Deux plongees au large du Japon avec bathy
Larson, R. J., C. E. Mills & G. R. Harbison 1991. Western At scaphe francais F.N.R.S. III. Bull. Inst. Oceanogr. Monaco
lantic midwater hydrozoan and scyphozoan medusae: in situ 1134: 1-28.
studies using a manned submersible. Hydrobiologia 216/217: Pugh, P. R. & G. R. Harbison 1987. Three new species ofprayine
311-317. siphonophore (Calycophorae, Prayidae) collected by a sub
Lindsay, D., J. Hunt, J. Hashimoto, K. Fujikura, Y. Fujiwara, S. mersible, with notes on related species. Bull. Mar. Sci. 41:
Tsuchida & K. Itoh 1999. The benthopelagic community of 68-91.
Sagami Bay. JAMSTEC J. Deep Sea Res. 14: 493-499. (in Pugh, P. R. & M. J. Youngbluth 1988. Two new species ofprayine
Japanesewith Englishabstract) siphonophore (Calycophorae, Prayidae) collected by the sub-
Lindsay, D., Y. Furushima, H. Miyake, M. Kitamura & J. Hunt mersibles Johnson-Sea-Link I and //. J. Plankton Res. 10:
2004. The scyphomedusan fauna ofthe Japan Trench: prelimi 637-657.
nary results from a remotely-operated vehicle. Hydrobiologia Russell, F. S. 1956. On two new medusae, Merga reesi n. sp. and
530/531: 537-547. Tiaropsidium atlanticum n. sp. J. Mar. Biol. Ass. V. K. 35:
Maas, O. 1905. Die craspedoten medusen der Siboga-Expedition. 493^98.
SibogaExped. Monogr. 10: 1-85. pis. 1-14. Talley, L. D. 1993. Distribution and formation ofNorth Pacific In
Mariscal, R. N. 1974. Nematocysts, p. 129-178. In Coelenterate termediateWater.J. Phys. Oceanogr. 23: 517-537.
biology (ed. L. Muscatine & H. M. Lenhoff). Academic Press, Torrey, H. B. 1909. The Leptomedusae ofthe San Diego region.
N. Y. Univ. CaliforniaPubl. Zool. 6: 11-31.
Matsumoto, G. I., K. A. Raskoff& D. J. Lindsay 2003. Tiburonia Toyokawa, M, T. Toda, T. Kikuchi & S. Nishida 1998. Cnidarians
granrojo n. sp., a mesopelagic scyphomedusa from the Pacific and ctenophores observed from the manned submersible
Ocean representing the type ofa new subfamily (class Scypho- Shinkai 2000 in the midwater ofSagami Bay, pacific coast of
zoa: order Semaeostomeae: family Ulmaridae: subfamily Japan.PlanktonBiol. Ecol.45: 61-74.
Tiburoniinaesubfam. nov.)Mar. Biol 143: 73-77. Uchida, T. 1928. Studies on Japanese hydromedusae. 2. Tra-
Mayer, A. G. 1910. Medusae oftheworld. Hydromedusae, vols. I, chymedusaeandNarcomedusae.Jpn.J. Zool. 2: 73-97.
II,pp. 1-498, pis. 1-76. Carnegie Inst. Washington. Uchida, T. 1947. Medusae in the vicinity ofShimoda.J. Fac. Sci.
Metschnikoff, E. 1886. Medusologische Mittheilungen. Arb. zool. Hokkaido Univ.9: 331-343.
hist. Univ. Wien. 6: 237-266. Uchida, T. 1965. 79. Tiaropsis rosea Agassiz et Mayer, p. 192. In
Miyake, H., D. J. Lindsay, J. C. Hunt & T. Hamatsu 2002. New encyclopedia ofthefauna ofJapan (ed. Okada, Uchida&
Scyphomedusa Aurelia limbata (Brandt, 1838) found in deep Uchida). Hokuryukan,Tokyo, (inJapanese)