Ernst Haeckel s mysterious species, Part I: the validity of Carybdea murrayana Haeckel, 1880 (Cubomedusae) and revisional notes on Haeckel s other ...
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Plankton Benthos Res 15(1): 1–29, 2020
Plankton & Benthos
Research
© The Plankton Society of Japan
Ernst Haeckelʼs mysterious species, Part I:
the validity of Carybdea murrayana Haeckel,
1880 (Cubomedusae) and revisional notes
on Haeckelʼs other Carybdeidae
Ilka Straehler-Pohl*
Medusa(ʻ)s nursery, Private Laboratory of Developmental, Evolutionary Biology and Life Cycle Research, Altmarkstr. 25,
21864 Stade-Hagen, Germany
Received 26 February 2019; Accepted 30 September 2019 Responsible Editor: Dhugal Lindsay
doi: 10.3800/pbr.15.1
Abstract: The type material of the species Carybdea murrayana Haeckel, 1880 was rediscovered in the Cubozoa
collection of the Natural History Museum in London. A comparison of C. murrayana with Carybdea marsupialis (Lin-
naeus, 1758) and with Carybdea branchi Gershwin & Gibbons, 2009 was performed because the validity of the species
has been doubted for over a century and because C. murrayana had been declared a synonym of C. marsupialis by au-
thors like Mayer, Bigelow and Kramp, due to an apparent overlapping distribution range. The results demonstrate that
C. murrayana is different from C. marsupialis but identical to C. branchi. Therefore, C. branchi is declared a junior
synonym of the valid species C. murrayana Haeckel, 1880 according to the International Code of Zoological Nomen-
clature.
Additionally, the line drawings and/or descriptions of the carybdeid species Procharagma prototypus Haeckel, 1880
and Procharybdis cuboides Haeckel, 1880 were translated, diagnosed, compared to well-known carybdeid species and
revised.
Key words: box jellyfish, carybdeid species, Carybdea marsupialis, museum collection, John Murray Expedition
which might have led also to general doubt concerning his
Introduction
descriptions of new species.
Ernst Haeckel is one of the most disputed scientists of Haeckel described about 18 new cubomedusan species
the 19th century due to his partial support of Darwinʼs the- in his 2nd part of “System der Acraspeden, Medusen” at the
ory and their friendship, of further developing his own end of the 19th century (Haeckel 1880) (see Table 1).
recapitulation theory (“ontogeny recapitulates phylogeny”) Nearly all of the species newly described by Haeckel
of evolution and for establishing the disputed and nowa- (1880) were doubted or declared invalid by many authors
days disproved biogenetic law or embryological parallel- in the past into the present (e.g. Mayer 1910, Kramp 1961,
ism (Haeckel 1899, Blechschmidt 1977, Krauße 1987). Gershwin 2005a+b, 2006a, Collins et al. 2011 Bentlage &
Haeckelʼs ideas were often speculative and mostly lacked Lewis 2012). The main reasons were: nearly all described
empirical support, which tarnished his scientific creden- species seemed to have never turned up again; some were
tials (Haeckel 1899, Bowler 1989, Milner 1990, Krauße based on doubtful material (e.g. juvenile, partly destroyed
1987). Additionally, he was accused of falsification con- specimens); and others were seen as synonyms of already
cerning drawings (Rütimeyer 1868, Teudt 1909, Richards described species from other locations.
2008) commented on and denied several times by Haeck- Carybdea murrayana Haeckel, 1880 is one example of
el himself (Haeckel 1891, Haeckel 1910, Richards 2008), such a doubtful species. It was sampled by the naturalist
John Murray during the H.M.S. Challenger Expedition in
* Corresponding author: Ilka Straehler-Pohl; E-mail, I.Straehler-Pohl@ 1876, named by Haeckel in 1880 (described and drawn
web.de in more detail by him in 1882) but was never officially
2 I. Straehler-Pohl
Table 1. List of cubomedusan species identified and described by Haeckel (1880), including sampling locations and collectors–species
observed for the present study are marked in bold.
Original species Size:
Reference for Synonym (change)/ Sampling location/ Actual
No. name (according BH×BW Actual Family
synonyms Status Collector Suborder
to Haeckel 1880) (mm)
1 Charybdea Bentlage & Lewis Alatina obeliscus 35×20 West coast of Africa; Cape Carybdeida Alatinidae
obeliscus (2012): (Haeckel, 1880) / Verde lslands / Museum
nomen dubium Godeffroy, Germany
2 Charybdea Bentlage & Lewis Alatina philippina 30×20 Pelew Islands (Palau), Carybdeida Alatinidae
philippina (2012): (Haeckel, 1880) / The Philippines / Semper
nomen dubium
3 Charybdea Gershwin (2005a, b): Alatina pyramis 30×20 Tropic belt of Atlantic Carybdeida Alatinidae
pyramis (Haeckel, 1880) / valid Ocean; Antilles / Museum
Godeffroy, Germany
Bentlage & Lewis Alatina pyramis (Haeckel,
(2012): 1880) / nomen dubium
4 Procharybdis Mayer (1910): Maybe juvenile of 40×40 Pacific Coast of Central Carybdeida Alatinidae
securigera Carybdea rastonii America / Fuchs
Haacke, 1887 / not valid
5 Procharybdis Mayer (1910): Procharybdis tetraptera / 30×20 Indian Ocean; Carybdeida Alatinidae
tetraptera maybe a Carybdea Sunda-Archipelago
species and/or immature (Sunda Strait)-Java/
Sumatra/ Rabbe
Gershwin (2005a, b): Alatina tetraptera
(Haeckel 1880) / valid
6 Procharybdis Bentlage & Lewis Alatina turricula (Haeckel, 170×70 Pelew Islands (Palau), Carybdeida Alatinidae
turricula (2012): 1880) / nomen dubium The Philippines/ Semper
7 Charybdea Kramp (1961): Carybdea marsupialis 60×50 West coast of Africa, Carybdeida Carybdeidae
murrayana (Linnaeus, 1758) / not far from Sierra
not valid Leone/John Murray
Gershwin & Gibbons Carybdea murrayana
(2009): Haeckel, 1880 / valid
8 Procharagma Mayer (1910): Carybdea rastonii 8×8 Chinese Sea/Weber Carybdeida Carybdeidae
prototypus (Haacke, 1887) / not valid
Bentlage & Lewis Carybdea prototypus
(2012): (Haeckel, 1880),
nomen dubium
9 Procharybdis Mayer (1910): Carybdea rastonii 35×35 Sandwich Islands/n/a Carybdeida Carybdeidae
cuboides Haacke, 1887 / not valid
10 Tamoya Mayer (1910): Tamoya haplonema / 80×40 West Indian Ocean, Carybdeida Tamoyidae
prismatica not valid Antilles / Schnehagen
Straehler-Pohl Tamoya prismatica /
(2019): species inquirenda
11 Procharybdis Mayer (1910), Procharybdis flagellata / 40×20 North coast of Carybdeida ?
flagellata Southcott (1956): not valid Australia;Torres-Strait /
Weber (New-Guinea /
Lesson?)
12 Procharagma Gershwin 2005a Copula aurea / ? 10×? Pelew Islands (Palau), Carybdeida Tripedaliidae
aurea The Philippines / Semper
13 Tamoya Bentlage & Lewis Tamoya bursaria (Lesson, 100×70 New-Guinea, Rawack, Carybdeida ?
bursaria (2012): 1829) / nomen dubium Waigiou / Lesson
14 Tamoya Bentlage & Lewis Tamoya haeckeli 160-200 Tropical Part of Pacific Carybdeida ?
gargantua (2012): Southcott, 1967= ×80-100 Ocean; Tahiti / Lesson;
Tamoya gargantua / valid Samoa / Weber
Straehler-Pohl Tamoya haeckeli
(2019): Southcott, 1967 /
species inquirenda
15 Chirodropus Uchida (1929): Chirodropus gorilla / valid 150×120 Coast of Lower Guinea Chirodropida Chirodropidae
gorilla (Equatorial Guinea),
Chinchozo, Loango /
Falkenstein
Validity of Carybdea murrayana Haeckel, 1880 3
Table 1. List of cubomedusan species identified and described by Haeckel (1880), including sampling locations and collectors–species
observed
Table 1. forContinued.
the present study are marked in bold.
Original species Size:
Reference for Synonym (change)/ Sampling location/ Actual
No. name (according BH×BW Actual Family
synonyms Status Collector Suborder
to Haeckel 1880) (mm)
Thiel (1936): Chiropsalmus
quadrumanus / not valid
Kramp (1961): Chirodropus gorilla/valid
16 Chirodropus Mianzan & Cornelius Maybe conspecific with 100×70 South Atlantic Ocean, Chirodropida Chirodropidae
palmatus (1999): Chirodropus gorilla/ Not far from Island St.
uncertain status Helena/Levasseur
Thiel (1936): Chiropsalmus
quadrumanus/not valid
Gershwin (2006a): Chirodropus palmatus/
provisionally valid
Straehler-Pohl Chirodropus palmatus/
(2019): valid
17 Chiropsalmus Gershwin (2006a): Chiropsoides 50×45 Indian Ocean, Rangoon/ Chirodropida Chiropsalmidae
quadrigatus* quadrigatus/valid Thallitzer
18 Chiropsalmus Thiel (1928): Chiropsalmus quadrigatus 60×40 South Atlantic Ocean, on Chirodropida Chiropsalmidae
zygonema (=Chiropsoides the Argentinean coast/
quadrigatus)/not valid Smith
Uchida (1929): Chiropsalmus zygonema/
uncertain species
Thiel (1936): Chiropsalmus
quadrumanus/not valid
Southcott (1956): Chiropsalmus zygonema/
probably immature form,
status unsure
Gershwin (2006a): Chiropsalmus
zygonema/valid
*confused with other chirodropid species from Asian waters that belonged to other families and genera by e.g. Mayer (1910), Southcott
(1956), Kramp (1961).
sighted again and was, therefore, doubted and seen as a
Material and Methods
probable variety of Carybdea marsupialis (Linnaeus, 1758)
(Mayer 1910, Bigelow 1938, Kramp 1961). Gershwin & All medusae specimens observed were preserved in 5%
Gibbons (2009) were the first to argue for the validity of formalin and water.
the species. Bentlage & Lewis (2012) listed C. murrayana
Morphological comparison
in their Carybdeidae species list.
The type material of C. murrayana was rediscovered The rediscovered type specimens of Carybdea mur-
in 2016 in the cubomedusan collection of the Natural His- rayana were preserved in 5%–7% formalin; therefore, no
tory Museum of London. It consisted of two mature female molecular analysis could be performed. However, as all
medusae, one was dissected and broken into three pieces, anatomical structures were excellently preserved a direct
and the other one was only cut open on one side. These comparison with the structures of the Carybdea species
specimens show all the features that Haeckel (1880, 1882) from Africa and Spain was possible. Therefore, only mor-
drew and described. As there was always doubt concern- phological characters were used to identify the species, a
ing the validity of this species, the goal of this paper is to valid approach even in the age of molecular phylogenies
provide more information on C. murrayana by comparing (Páll-Gergely 2017).
it with specimens of C. marsupialis and Carybdea branchi
Species observed
Gershwin & Gibbons, 2009 this last one with overlapping
distribution range. Project 1: Carybdea murrayana:
Additionally, two other species of the familiy Carybdei- As the statement that Carybdea murrayana from West
dae described by Haeckel (1880) as Procharagma proto- Africa is considered to be a synonym of “Carybdea marsu-
typus Haeckel, 1880 and Procharybdis cuboides Haeckel, pialis” by Mayer (1910), Bigelow (1938) and Kramp (1961)
1880 are diagnosed and revised. was based on descriptions by Linnaeus (1758) and Claus
(1878), specimens of C. marsupialis from the Mediterra-
nean Sea were also examined. As the distributional range
of Carybdea branchi overlaps with that of C. murrayana,
4 I. Straehler-Pohl
Table 2a. Unregistered specimens examined for this study.
No. of
Provider Species ID on label Sampling location** Collector Sampling date
specimens
André C. Morandini 7 Carybdea branchi South Africa, Hout Bay A.C. Morandini & 05 May 2013
Gershwin & (34°03′4.8″S, 18°20′53.99″E), S.N. Stampar
Gibbons, 2009* swimming at surface near pier
Melisa J. Acevedo 5 Carybdea marsupialis Spain, Denia, Almadrava Beach M. J. Acevedo 10, 29 Sep 2010
(Linnaeus, 1758) (38°51′50″N, 0°1′24″E), surface,
dipnet
Angel Yanagihara 10 Carybdea arborifera Hawaii, Kewalo Basin A. Yanagihara 29 May 2013
Maas, 1897 (21°17.564′N, 157°51.462′W)
Sho Toshino 3 Carybdea brevipedalia Japan, Kanagawa Prefecture, S. Toshino 24 Oct 2011
Kishinouye, 1891 Aburatsubo Bay (35°09′35.2″N,
139°36′56.0″E)
Sho Toshino 3 Carybdea brevipedalia Japan, Oita Prefecture, Oita City, S. Toshino Sep 2017
Kishinouye, 1891 Hoso fishing port (33°14′37″N,
(juvenile medusae of 131°46′34″E)
different sizes and
developmental stages)
Sho Toshino 1 Carybdea brevipedalia Japan, Wakayama Prefecture, S. Kubota 1995
Kishinouye, 1891 Shirahana (33°40′54″N,
135°20′40″E)
Jamie Seymour 1 Carybdea rastonii Australia, Victoria, Mirimbula G. Hood 03 Mar 2000
Haacke, 1887 (36°53′53″S, 149°54′04″E)
Jamie Seymour 1 Carybdea rastonii South Australia, Waterloo Bay J. Seymour Feb 1999
Haacke, 1887 (33 39′ 00″S, 134 54′ 00″E)
George I. Matsumoto 8 Carybdea confusa USA, California, Santa Barbara, S. Anderson 21 Oct 1998
Straehler-Pohl, 20 m West of Goleta Pier, 5 m
Matsumoto & depth (34°24′58″N, 119°49′43″W)
Acevedo, 2017*
*Identifications were done in 2013, **longitudes and latitudes taken from: http://latitude.to or http://google.de/maps
the holotype, paratypes, additional registered museum ma- Maas, 1903, Carybdea brevipedalia Kishinouye, 1910, and
terial and specimens of C. branchi recently sampled in Carybdea rastonii Haacke, 1887 due to type localities and
South Africa were also examined. proposed synonymy by Mayer (1910).
All information about the specimens observed and the All information about the specimens observed and the
referring museum collections are listed in Tables 2a, b. museum collections are listed in Tables 2a, b.
Additionally, all data were compared to the original line
Measurements
drawings made by Haeckel in 1879 (with courtesy of the
collections of Ernst-Haeckel-Haus, Friedrich-Schiller-Uni- Standard measurements were used (Gershwin & Gib-
versität Jena Institut für Geschichte der Medizin, Natur- bons 2009, Straehler-Pohl 2014, Acevedo et al. 2019): bell
wissenschaft und Technik) and to the detailed descriptions height (BH) as length between bell turn-over (velarium
and line drawings of Haeckel (1880, 1882). excluded from measurement) and top of apex; interpedalial
diameter (IPD) as distance between opposite pedalia (outer
Project 2: Procharagma prototypus and Procharybdis pedalial wing edges) at the level of the bell turn-over; in-
cuboides terrhopalial width (IRW) was measured between adjacent
As the type materials of Procharagma prototypus and rhopalia, with the specimen flattened; pedalia length (PL)
Procharybdis cuboides could not be found in the European was measured from attachment to bell (pedalial base) to
museum collections that were visited, only data from the the tentacle insertion, as a proportion in relation to bell
literature (Haeckel 1880, 1904) and the original line draw- height.
ings of Haeckel (between 1877 and 1880, with courtesy of Photographs were taken under the same conditions with
the collections of the Ernst-Haeckel-Haus, Friedrich-Schil- digital cameras (Canon Powershot G12 and Canon Eos
ler-Universität Jena Institut für Geschichte der Medizin, 550D).
Naturwissenschaft und Technik) were used to diagnose the
“Gonads” in Cubozoa
validity and identity of the species.
Species used for comparison were Carybdea arborifera The study follows Acevedo et al. (2019) in using the
Validity of Carybdea murrayana Haeckel, 1880 5
Table 2b. Specimens from museum collections examined for this study.
No. Species Original Sampling Sampling
Collection Code No. Collector
specimens identification identification location date
NHM 1882.10.9.2a+b 2 Carybdea Charybdea West Africa, John Murray 04 Apr 1876
(lecto- & murrayana murrayana Haeckel, 20°10′N, 14°51′W
paralectotype) Haeckel, 1880*** 1880 (type species)
NHM 2000.1800–1803 4 Carybdea 1938: Medusae; South Africa, Discovery 17 Nov 1938
murrayana Gershwin 2000: Simonstown docks; Expedition
Haeckel, 1880*** Carybdea robsonae Hand net, surface
Gershwin, Paratypes;
Gershwin &
Gibbons 2009:
Carybdea branchi
NHMD ̶ 1 Carybdea Carybdea alata South Africa, G.F. Papenfuss 27 Oct 1937
murrayana Reynaud 1830 False Bay (“Africana
Haeckel, 1880*** Expedition”:
University of
Cape Town
Ecological
Survey 1937-40)
RBINS I.G. 11204 1 Carybdea Dr. G. Ranson, 1945: 4 miles off Lüderitz Cruise of 18 Jan 1938
murrayana Tamoya haplonema Bay (Namibia, Africa) “Mercator”
Haeckel, 1880*** Müller, 1859
NHM 1983.4.25.1 1 Carybdea Carybdea Durban Museum E.T. Brown beq. Dec 1914
arborifera marsupialis (South Africa)
Maas 1897** (Linnaeus, 1758)
MCNB MZB 2015-4807 1 Carybdea Carybdea branchi South Africa, Hout A.C. Morandini 05 Ma y 2013
murrayana Gershwin & Bay (34°03′4.8″S, & S.N. Stampar
Haeckel, 1880*** Gibbons, 2009 18°20′53.99″E),
swimming at surface
near pier
SAM H4863 (holotype 1 Carybdea Carybdea branchi SE corner of Alfred L. Gershwin & 18 Jan 2001
C. branchi) murrayana Gershwin & Basin, in front of L. Hoensen
Haeckel, 1880**** Gibbons, 2009 the Two Oceans
Aquarium, V&A
Waterfront, Cape
Town [33°54.527′S,
018°25.074′E], surface
SAM H4864a+b 2 Carybdea Carybdea branchi SE corner of Alfred L. Gershwin & 18 Jan 2001
(paratypes murrayana Gershwin & Basin, in front of L. Hoensen
C. branchi) Haeckel, 1880**** Gibbons, 2009 the Two Oceans
Aquarium, V&A
Waterfront, Cape
Town [33°54.527′S,
018°25.074′E], surface
NHM 21.11.16.14 3 Carybdea Carybdea rastonii Japan, Mesaki ?, collected for 1921
brevipedalia Haacke, 1887 Exhibition
Kishinouye, 1891**
CAS CASIZ 197981 1 Carybdea confusa Carybdea California, Santa Shane Anderson 21 Oct 1998
Straehler-Pohl, marsupialis Barbara, 20
Matsumoto & (Linnaeus, 1758) meters west of
Acevedo, 2017 Goleta Pier
(34°24′58″N,
119°49′43″W),
5 m depth
CAS CASIZ 197982 2 Carybdea confusa Carybdea California, Santa Shane Anderson 21 Oct 1998
Straehler-Pohl, marsupialis Barbara, 20
Matsumoto & (Linnaeus, 1758) meters west of
Acevedo, 2017 Goleta Pier
(34°24′58″N,
119°49′43″W),
5 m depth
6 I. Straehler-Pohl
Specimens from museum collections examined for this study.
Table 2b. Continued
No. Species Original Sampling Sampling
Collection Code No. Collector
specimens identification identification location date
BNHM 1997.768-781 14 Carybdea rastonii Carybdea rastoni Australia, New South P.F.S. Cornelius 18 Feb 1994
Haacke 1887** Haacke 1887 Wales, Cabbage Tree
Bay, 33°48.08′S,
151°17.58′E
BNHM 1972.5.24.1 1 Carybdea Carybdea S. Italy, Gargano P. R. Laming 1972
marsupialis marsupialis Peninsula, Bay
(Linnaeus, 1759)** (Linnaeus, 1758) of Campi
MCNB MZB 2015-1701 1 Carybdea Carybdea Spain, Denia, Melisa J. 06 Oct 2010
marsupialis marsupialis Almadrava Beach Acevedo
(Linnaeus, 1758)* (Linnaeus, 1758) (38°51′50″N,
0°1′24″E), surface,
dipnet
MCNB MZB 2015-4806 10 Carybdea Carybdea Spain, Denia, Melisa J. 10, 29 Sep 2010
marsupialis marsupialis Almadrava Beach Acevedo
(Linnaeus, 1758)* (Linnaeus, 1758) (38°51′50″N,
0°1′24″E), surface,
dipnet
*Identifications were done 2015, **Identifications were done in 2016, ***Identifications were done in 2017, ****Identifications were done
in 2019 NHM: British Museum of Natural History, London NHMD: Natural History Museum of Denmark, Copenhagen MCNB: Museu de
Ciències Naturals de Barcelona RBINS: Royal Belgium Institute of Natural Science, Brussels SAM: Iziko South African Museum, Cape
Town.
term gonads to refer to areas where gametes are formed. 2019)
With heart-shaped rhopalial niche openings showing a
single upper covering scale and no lower covering scale;
Results
lacking rhopalial horns. Gastric phacellae either brush to
epaulette-shaped or brush-shaped filaments growing in
Project 1̶Carybdea murrayana
horizontal rows. Polypoid stage and medusa production:
There were two errors noted in Acevedo (2016: pp. 30, only known for two species, Carybdea xaymacana Conant,
32, 41, 42, 43) and Straehler-Pohl et al. (2017: p. 131, 136) 1897 and Carybdea brevipedalia. Medusa production: only
concerning the structures of the velarial canal system in C. known from C. xaymacana and C. brevipedalia.
branchi and C. murrayana, partly corrected by Acevedo et Type species: Carybdea marsupialis (Linnaeus, 1758)
al. (2019: 526–527, 539)̶both were described to possess
3 velarial canal roots per octant. Additional inspections Carybdea murrayana Haeckel, 1880
confirmed the results of Gershwin & Gibbons (2009) that Type locality: West Coast of Africa, not far from Sierra
C. branchi possesses only 2 velarial canal roots per octant Leone (30°10′N, 14°51W), Challenger Expedition 1872-76
and revealed that C. murrayana also possesses only 2 ve- Station 348, from 400 m depth (Haeckel 1880, 1882).
larial canal roots per octant. Herewithin, the errors by Ace-
vedo (2016) and Straehler-Pohl et al. (2017) are corrected Inspection of Syntypes
within Table 3 ʻTable of Identificationʼ and Figures 4S, T, The “holotype” consisted of 2 specimens (=syntypes)
U, 5G, M. of which one was dissected by Haeckel for his descrip-
tion and line drawings of structural details (Haeckel 1880,
Systematics (based on Straehler-Pohl (2017), Jarms & Morandini 1882) while the other was just cut open for the habitus line
(2019)) drawings by Haeckel (1882, 1904) but was not further dis-
sected. Even if the first specimen is broken into 3 parts, the
Phylum Cnidaria Verrill, 1865
Subphylum Medusozoa Petersen, 1979 structures are in better condition than in the other one. I
Class Scyphozoa Goette, 1887 designate it as the lectotype (1882.10.9.2a) also because the
Order Cubomedusae Haeckel, 1880 first description is mainly based on this specimen, while
Suborder Carybdeida Gegenbaur, 1857 the other is designated as a paralectotype (1882.10.9.2b)
Family Carybdeidae Gegenbaur, 1857 according to the International Code of Zoological Nomen-
Genus Carybdea Péron & Lesueur, 1810
clature (ICZN 1999).
The lectotype (NMH 1882.10.9.2a, Fig. 1B) was in good
Genus Carybdea Péron & Lesueur, 1810 condition even if completely dissected; the bell was bro-
Definition: (after Acevedo et al. 2019, Straehler-Pohl ken into three pieces (Fig. 1B) but the internal structures
Validity of Carybdea murrayana Haeckel, 1880 7
Table 3. Identification Table: Comparison of characters of specimens of Carybdea marsupialis from Spain, Carybdea murrayana from
West Africa and Carybdea branchi from South Africa.
Carybdea marsupialis Carybdea murrayana Carybdea branchi
(literal citation) (literal citation) (literal citation)
MZB 2015-1701 (neotype); MZB 2015- NMH 1882.10.9.2a+b (lecto- & paralec- SAM-H4863 (holotype); SAM-H4864a+b
4806; BNHM 1972.5.24.1; 5 unregistered totype); Haeckel# 1880, 1882; Figs. 1A–Q, (paratypes); NMH 2000.1800–1803; I.G.
specimens from Denia; Spain; Acevedo et 2A–F, 3A, B, 5H–M, 6 of present study 11204; RBINS IG 11204; NHMD unregis-
al. 2019; Figs. 5N–R tered specimen; 7 unregistered specimens
from Hout Bay, South Africa; MZB 2015-
4807; unregistered living specimen from
Lüderitz Bay, Namibia (Simon Elwen
(Namibian Dolphin Project)), Gershwin
& Gibbons 2009 (measurements of 28
specimens used for statistics); Figs. 4A–U,
5A–G
Bell
• highly transparent, colourless, base • transparent to translucent, colourless to • highly transparent, base of gastric
of gastric phacellae, brownish-orange reddish brown (preserved; Figs. 1B, I, J) phacellae, yellowish to reddish-brown,
(freshly preserved, in-life; Fig. 5O, S); pedalium, outer wing base, light brown,
translucent whitish (preserved; Fig. 5N) outer wing distal end, dark reddish brown
(in-life (Figs. 4A, M): colouration fades
fast after preservation); transparent to
translucent, colourless to yellowish (pre-
served; Figs. 4B–E, 5A)
(Acevedo et al. 2019, p. 521: highly trans- (Gershwin & Gibbons 2009, p. 47: slightly
parent with few whitish nematocyst warts translucent whitish with conspicuous pig-
sparsely scattered on bell from apex (very mentation comprising a single dark red
small warts) to bell margin (big warts blotch on abaxial corner of pedalium at
along interradial furrows); [...] phacellae base of each of the 4 tentacles (Plates 1A,
brownish-orange in colour, colour remains B, 2C), a single small red blotch on apical
after preservation (Figs. 5C, D)) portion of exumbrella above each of the
four phacellae (Plate 1A), and a single
faint brownish blotch at “shoulder” of each
of the four pedalia)
• mesogloea, thin • mesogloea, firm, thick at interradial • mesogloea, firm, thick at apex and in-
corners (Fig. 1I, 2A, 3A2–3) terradial furrows (Figs. 4C–E, 5A)
(Haeckel 1882, p. 94: The gelatinous (Gershwin & Gibbons 2009, p. 43: with
substance of the umbrella shows a consid- thick, rigid mesoglea, especially apically)
erable degree of firmness, [...], thickest at
the two sides of the pillars [...])
• cylindrical to pyramidal with rounded • almost cubical to truncate pyramidal • cubic to pyramidal with rounded edges
edges (Fig. 5N) with rounded edges (Figs. 1I, 3A1–3) (Figs. 4A–E, 5A)
(Haeckel 1882, p. 93: [...] nearly cubical,
[...] shape of a truncated, regular quadrilat-
eral pyramid.)
• apex, very thick mesogloea, domed, • apex, mesogloea, slightly thicker than • apex, thick mesogloea, slightly arched,
with horizontal constriction (Fig. 5N) in other parts, slightly arched, no horizon- with slight horizontal constriction (Fig.
tal constriction visible (Figs. 2A, 3A1–4) 4C–E)
(Haeckel 1882, p. 94: The gelatinous (Gershwin & Gibbons 2009, pp. 43, 44:
substance of the umbrella [...] thickest [...] [...] with thick, rigid mesoglea, especially
above in the cap-shaped apical cover of apically; [...] Coronal indentation shallow
the umbrella) just below apex. [...] Fig. 1A)
8 I. Straehler-Pohl
Table 3. Identification Table: Comparison of characters of specimens of Carybdea marsupialis from Spain, Carybdea murrayana from
West Africa
Table and Carybdea branchi from South Africa.
3. Continued.
Carybdea marsupialis Carybdea murrayana Carybdea branchi
(literal citation) (literal citation) (literal citation)
• nematocyst warts, small, densely scat- • nematocyst warts, none visible in NMH • nematocyst warts, different sizes (in-
tered, flat on bell sides, prominent on 1882.10.9.2a+b (according to Haeckel terradial furrows, bell margin, bigger
apex, round to oval, whitish to brownish 1882, pp. 93–94: The exumbrella appears nematocyst warts, bell sides, apex smaller
(in-life; Fig. 5S), from apex to velarium finely granulated, numerous urticating nematocyst warts), densely scattered from
warts or round groups of thread cells are apex to bell-turnover (Fig. 4A), flat to
scattered freely over it) prominent, irregularly shaped, white (Fig.
4L, M); easily rubbed off when handled,
preserved and sent to other labs (own ob-
servation; Figs. 4E, 5A)
(Gershwin & Gibbons 2009, p. 43: [...]
with numerous oblong to amorphous
small unraised or slightly raised nemato-
cyst warts scattered densely over entire
exumbrella)
Size
• BH: up to 30 mm (preserved; Fig. 5N) • BH: up to 60 mm (preserved: Haeckel • BH: up to 82 mm (preserved)
1882)
(mean: 28.2 mm, SD: 2.49, n=5) (actual mean: 53.0, SD: 4.243, n=2) (mean: 45.58, SD: 10.498, n=35)
• IRW: no data • IRW: 29 mm (preserved: Haeckel • IRW: up to 51 mm
1880)
(actual mean: 46.5, SD: 4.95, n=2) (mean: 28.55, SD: 7.566, n=35)
• IPD: up to 35 mm • IPD: up to 68 mm • IPD: up to 99 mm (preserved)
(mean: 32.6 mm, SD: 2.51, n=5) (actual mean: 65.00, SD: 4.243, n=2) (mean: 57.53, SD: 14.410, n=35)
• IPD/BH: 1.16 (mean, SD: 0.099, n=5) • IPD/BH: 1.23 (mean, SD: 0.018, n=2) • IPD/BH: 1.26 (mean; SD: 0.076,
n=35)
• IRD/BH: no data • IRW/BH: 0.66 (mean; SD: 0.011, • IRW/BH: 0.62 (mean; SD: 0.053,
n=2) n=35)
• PL: up to 13 mm (mean: 11.4 mm, SD: • PL: 32 mm, 25 mm (n=2) • PL: up to 40 mm (mean: 31.20, SD:
1.14, n=5) 7.079, n=5)
• PW: up to 7 mm (mean: 6.6 mm, SD: • PW: 18 mm, 14 mm (n=2) • PW: up to 23 mm (mean: 17.60, SD:
0.89, n=5) 3.286, n=5)
(Haeckel 1882, p. 93: Horizontal diameter (Gershwin & Gibbons 2009, p. 43: Bell to
of the umbrella, 50 mm; vertical diameter, about 68 mm in height (after several days
60 mm.) in formalin), about 80 mm live [...])
Gonads
• Interradial • interradial • interradial
• paired, 4 • paired, 4 • paired, 4
• size, covering area of gastric pouch • size, length covering entire area of • size, length covering entire area of
from bell margin to 3/4 of bell height be- gastric pouch from stomach to bell margin gastric pouch from stomach to bell margin
low stomach (Fig. 5N)
• narrow leaf-shaped, very thin tissue, • broad leaf-shaped (Figs. 1E, J), very • narrow to broad leaf-shaped, thin tis-
separated by perforated interradial sep- thin tissue, pleated, separated by unperfo- sue, pleated, separated by unperforated
tum, attached at entire length of septum rated septum, attached along entire length septum, attached at entire length of sep-
of septum tum, outer margins of adjacent gonad
leaves might touch or overlap (Figs. 4E)
Validity of Carybdea murrayana Haeckel, 1880 9
Table 3. Identification Table: Comparison of characters of specimens of Carybdea marsupialis from Spain, Carybdea murrayana from
West Africa
Table and Carybdea branchi from South Africa.
3. Continued.
Carybdea marsupialis Carybdea murrayana Carybdea branchi
(literal citation) (literal citation) (literal citation)
(Haeckel 1882, p. 100: eight broad, thin, (Gershwin & Gibbons 2009, p. 47: Go-
semi-oval leaves which are fastened in nads attached along entire length of inter-
pairs along the four interradial septal sel- radial septa; narrowly leaf-shaped, typi-
vages, and project freely from these into cally not overlapping along the interradius
the four radial pouches; they occupy the in present collection, pleated or simple.
greater part of their hollow space so that Interradial septa lacking perforations.)
the two reproductive leaves of each pouch
touch each other or even overlap with their
free margins in its middle; (present study
Fig. 3A6))
• sexes unimorph; opaque white in pre- • sexes̶both specimens female (Fig. • sexes unimorph; opaque yellowish
served specimens (7% formalin) 1E), males unknown; opaque flesh co- white to brownish flesh-coloured in pre-
loured in preserved specimens (7% for- served specimens (7% formalin)
malin)
Pedalia
• 4, single, simple (Fig. 5Q) • 4, single, simple • 4, single, simple
• slightly stalked • slightly stalked • slightly stalked
• non-keeled at midline • keeled at midline • keeled at midline
• irregularly, broad leaf-shaped • leaf-shaped (Figs. 1F, M, 2A2–3, 3A3) • leaf-shaped (Figs. 4M–P, 5E)
(Haeckel 1882, p. 95: [...] shaped like a
thin longish oval leaf)
• PL ca. 40% of BH • PL ca. 57% of BH (mean, SD: 0.002, • PL ca. 55% of BH (mean, SD: 0.039,
n=2) n=7)
(Haeckel 1882, p. 95: [...] nearly a third as (Gershwin & Gibbons 2009, p. 43: Pedalia
long as the height of the umbrella) about 1/2 bell height)
Inner wing
• flattened, nearly scalpel-shaped (Fig. • flattened, semi-oval shaped • flattened, semi-oval
5Q)
(Gershwin & Gibbons 2009, p. 43: [...]
greatly rounded, nearly hemispherical)
• not overhanging tentacle insertion; in • no incision above tentacle insertion, • no incision above tentacle insertion,
some mature medusae margin sometimes slightly overhanging tentacle insertion in slightly overhanging tentacle insertion in
undulated one specimen (Fig. 5K, arrow) large specimens (Figs. 4M, O, 5E, arrow)
(Gershwin & Gibbons 2009, p. 43: [...]
without tentacular overhang)
• nematocyst warts, none • nematocyst warts, none • nematocyst warts, none
Outer wing
• straight, narrower than inner wing, not • semi-oval shaped, narrower and longer • narrow semi-oval shaped, narrower
overhanging tentacle insertion than inner wing and longer than inner wing
• nematocyst warts, irregular white • nematocyst warts, none visible in • nematocyst warts, irregularly shaped
nematocyst bands on outer keel, smaller NMH 1882.10.9.2a+b warts/bands from keel to pedalial canal
warts scatter outer wing (Fig. 5Q) border, scattered, (partly rubbed off in un-
registered specimens from Hout Bay); in-
life: light brown pigmentation at pedalial
base, dark brown pigmentation above ten-
tacle insertion (Fig. 4A (white arrows), M)10 I. Straehler-Pohl
Table 3. Identification Table: Comparison of characters of specimens of Carybdea marsupialis from Spain, Carybdea murrayana from
West Africa
Table and Carybdea branchi from South Africa.
3. Continued.
Carybdea marsupialis Carybdea murrayana Carybdea branchi
(literal citation) (literal citation) (literal citation)
Pedalial canal
• cross section, diamond-shaped at base • cross section, triangular at base, • cross section, triangular at base,
and midsection, flattened oval/ellipsoid diamond-shaped from below knee bend diamond-shaped from below knee bend
towards distal end towards distal end towards distal end
(Gershwin & Gibbons 2009, p. 43: Peda-
lial canal strongly quadratic in cross sec-
tion throughout length)
• same diameter from knee bend towards • same diameter from knee bend towards • same diameter from knee bend towards
tentacle insertion, tentacle diameter wider tentacle insertion, distal end broadly flared tentacle insertion, distal end broadly flared
than insertion (Figs. 1F, 5K (arrow)) (Figs. 4N–P, 5E)
(Gershwin & Gibbons 2009, p. 43: flared
slightly at tentacle insertion)
• straight after knee bend, after 1/3 • straight • straight
curved towards inner wing, after 2/3
straight towards distal end
• very slightly, laterally keeled • smooth edged/keeled at midline (Fig. • smooth edged/keeled at midline (Figs.
1F, 5K) 4O, Q, 5E)
• knee bend rounded to rectangular, knee bend bulged to triangular volcano- • knee bend bulged to triangular vol-
without any appendages (Fig. 5Q) shape (Figs. 1G, L, 5L) cano-shape (Figs. 4Q, white arrow, 5F,
white arrow)
(Gershwin & Gibbons 2009, p. 43: [...]
broadly rounded bulge or lateral-pointing
thorn)
Tentacles
• 4, single • 4, single • 4, single
• light brownish pink, with white nema- • flesh coloured (preserved: Fig. 1I, J) • flesh coloured (preserved: Figs. 4B–E,
tocyst-batteries (in-live: Fig. 5S); white 5A), light pink (in-life: Figs. 4A, M, V)
(preserved: Fig. 5N)
• width, tapering slightly from below • width, broadly flared at base, tapering • width broadly flared at base, tapering
base to distal end slightly below base to distal end (Figs. below base to distal end (Figs. 4A, J, 5A)
1I, J)
• broad, cross-section, round (in-life, • filiform, broad, round in cross-section, • filiform, broad, round in cross section
preserved) (preserved) (preserved and in-life)
• “pearl-string”, “string” bearing series of • bearing series of dense nematocyst • bearing series of dense, narrow nema-
“nematocyst-pearls” bands tocyst rings
Rhopalial niches
• 4, perradial, cavity, heart-shaped • 4, perradial, cavity, oval heart-shaped • cavity, oval heart-shaped
• orifice, rounded heart-shaped (Fig. 5P) • orifice, oval heart-shaped (Figs. 1D, • orifice, oval heart-shaped (Figs. 4J–M,
K, 3A8, 5J) 5D)
• covering scales (preserved: Fig. 5P): • covering scales (preserved: Fig. 5J): • covering scales (in-life: Fig. 4K; pre-
upper, 1, triangular with sharp longish tip, upper, 1, broad triangular, lower, 0, orifice served: 4J, L, M, 5D): upper, 1, broad tri-
lower, 0, orifice extends as narrow furrow extends as a very broad furrow running angular, lower, 0, orifice extends as a very
running from cavity base to bell margin from cavity base to bell margin broad furrow running from cavity base to
bell margin
(Haeckel 1882, p. 96: Each sense club [...] (Gershwin & Gibbons 2009, p. 43: 1 shal-
is partly covered externally by the protec- low covering scale above and 2 nearly
tive scale, which projects like a roof over imperceptible scales below)
the ectodermal aperture of the rhopalial
niche)
• 1/5 up from margin • 1/6 up from margin • 1/6 up from margin
• number of eyes per rhopalium, 6 • number of eyes per rhopalium, 6 • number of eyes per rhopalium, 6Validity of Carybdea murrayana Haeckel, 1880 11
Table 3. Identification Table: Comparison of characters of specimens of Carybdea marsupialis from Spain, Carybdea murrayana from
West Africa
Table and Carybdea branchi from South Africa.
3. Continued.
Carybdea marsupialis Carybdea murrayana Carybdea branchi
(literal citation) (literal citation) (literal citation)
Rhopalial horns
• none • none • none
Velarium
• narrow (≤1/4 of bell diameter) • broad (≥1/4 of bell diameter) • broad (≥1/4 of bell diameter)
(Gershwin & Gibbons 2009, p. 45: Ve-
larium narrow)
• nematocyst warts, some, small, loosely • nematocyst warts, none • nematocyst warts, none
scattered
(Gershwin & Gibbons 2009, p. 45: [...]
lacking nematocyst warts or freckles)
• canal roots, 3 per octant (Fig. 5R) • canal roots, 2 per octant (Figs. 1H, • canal roots, 2 per octant (Figs. 4S–U,
N–Q; 2C, F, 3B, 5M) 5G)
(Gershwin & Gibbons 2009, p. 45: [...]
Velarial canals 2 per octant)
• canals, slim; next to frenulum, 1, sim- • canals, broad, 2–4 per root, canals den- • canals, broad, 2–4 per root, canals den-
ple to slightly forked; middle, 1–2, only dritic, some side branches tend to grow in dritic, some side branches tend to grow in
single side branches; next to pedalium, centripetal direction (Figs. 1H, O, Q) centripetal direction (Figs. 4S, T, 5G)
3–4, most complex, several side branches
(Fig. 5R)
(Gershwin & Gibbons 2009, p. 45: [...]
dendritic (Plate 2D),with edges of branch-
es bearing lateral lobations)
• sharply pointed tips, deeply forked, • very complexly patterned (Figs. 1H, • very complexly patterned (Figs. 4K,
slightly lobed • with smooth margin (Fig. 2C, F, 3B, 5M), canals flanking frenulum L), canals flanking frenulum are as com-
5R) are as complexly branched as the ones plexly branched as the ones flanking the
flanking the pedalia; dendritic, lobed main pedalia; dendritic, lobed main branches
branches and several dendritic, lobed side and several dendritic, lobed side branches,
branches, all octants show a different pat- all octants show a different pattern (Figs.
tern (Figs. 1 N–Q, 3B); not anastomosing 4U, 5G); not anastomosing
(Haeckel 1882, pp. 99–100: dendritic, (Gershwin & Gibbons 2009, p. 45: [...]
caecal, velar canals run from their lower with edges of branches bearing lateral
or distal margin into the “velarium”. lobations, non-anastomosing)
[...] Their ramification is delicately den-
dritic and is weaker towards the perradius,
stronger towards interradius. There are
forty-eight velar canals on the whole, so
that twelve of them come on each quad-
rant. The largest velar canal lies nearest
the interradial pedalia and shows 6 to
8 pairs of side branches, partly simple,
partly cleft. [...])
Adradial lappets
• none • none • none
Perradial lappets
• none • none • none
Structures of digestive systems
• manubrium, short in length (1/4 of BH) • manubrium, short in length (1/5 of • manubrium, intermediate in length
not reaching lower bell half, mouth arms, BH) not reaching into lower bell half (pre- (1/5–1/4 of BH) not reaching into lower
4, narrow, no nematocyst warts on mouth served), mouth arms, 4, broad, large/long bell half (preserved), mouth arms, 4,
tube and lips with rounded tips (Fig. 3A9) broad, long, rounded tips (Fig. 4R)12 I. Straehler-Pohl
Table 3. Identification Table: Comparison of characters of specimens of Carybdea marsupialis from Spain, Carybdea murrayana from
West Africa
Table and Carybdea branchi from South Africa.
3. Continued.
Carybdea marsupialis Carybdea murrayana Carybdea branchi
(literal citation) (literal citation) (literal citation)
(Haeckel 1882, p. 97: [...] four lanceolate (Gershwin & Gibbons 2009, p. 47: Manu-
or oval “oral lobes.”) brium long, with large, narrowly rounded
lips reaching to at least halfway down bell
cavity in life)
• stomach, small, flat, square • stomach, wide but flat (Figs. 2A, 3A) • stomach, large but flat
• mesenteries, narrow, small • mesenteries, well developed (Fig. 2A, • mesenteries, well developed
3A)
• gastric phacellae, 4, interradial, epau- • gastric phacellae, 4, interradial, bushy, • gastric phacellae, 4, interradial, bushy,
lette-shaped (Fig. 5O), mounted on four, epaulette-shaped (Figs. 1C, 2B, E, 3A, 5I), epaulette-shaped (Fig. 4F–H), gastric
conspicuously raised stomach corners gastric filaments, single-rooted, multiple filaments, single-rooted, multiple-stalked,
(Fig. 5S); filaments brush-like, tightly stalked, tree-like branched (Figs. 1C, 2B, tree-like branched (Fig. 4I), 10 to 20 fila-
bundled, originating from single root, 3A, 5I), 10 to 12 filaments per single stalk, ments per single stalk, 1 to 3 branches per
deeply branched up from the root, with 1 to 3 branches per filament filament
numerous short gastric filaments; phacel-
lae, base, brownish-orange in colour, also
for some time after preservation
(Haeckel 1882, p. 98: [...] bush, composed (Gershwin & Gibbons 2009, p. 47: greatly
of ten to twelve larger and several smaller bushy, arising from up to about 20 closely
branches. The stems of these branches are pressed stalks in a single tight bundle;
connected below at the root, [...], and so each stalk dendritically branched numer-
actually represent the principal branches ous times to endings of single or paired
of a single, very short, powerful stem,) short, solid cirri)
Distribution (recorded sightings)
• Mediterranean Sea, Spain, Italy, Tune- • East Atlantic coast of West Africa (not • East Atlantic coast of Southwest Africa
sia, Malta far from Sierra Leone) (Lüderitz Bay, Namibia, present study) to
South Africa (Soldhana Bay; Langebaan;
Cape Town; Hout Bay; Port Elisabeth)
(Gershwin & Gibbons 2009, p. 47: Pres-
ently known only from South Africa, from
Port Elisabeth along the south coast, to
Laangebaan, Soldhana Bay, on the west
coast)
were in very good condition. The paralectotype (NMH Carybdea branchi Gershwin & Gibbons, 2009
1882.10.9.2b, Figs. 1I, J) was in good condition, the bell Type locality: South east corner of Alfred Basin, in front
complete, only cut open at one side, but the internal struc- of the Two Oceans Aquarium, V&A Waterfront, Cape
tures were in worse condition than those of the lectotype. Town [33°54.527′S, 018°25.074′E] (Gershwin & Gibbons
The internal structures of the lectotype and paralecto- 2009).
type were similar in their main characters (Fig. 1) but the Inspection of holotype, paratypes and other registered ma-
velarial canal pattern varied for each specimen (Figs. 1H, terial
N) as it was from octant to octant within one specimen The holotype (SAM-H4863, Fig. 4B) was in quite good
(Figs. 1N–Q). condition even if one pedalium was ripped off (but still
Next to the characters listed by Haeckel (1880, 1882; see present in the jar) and mostly opaque. The paratypes (Ta-
Table 3), four additional morphological features concern- bles 2b, Fig. 4C) and other registered and unregistered ma-
ing the pedalium and velarium were noted: terial (Tables 2a, b, Figs. 4D, E, 5A) were in equally good
(1) pedalial canal keeled at midline (Fig. 1F), (2) pedalial or even better condition.
canal knee-bend of bulged volcano-shape (Figs. 1G, L), In the paratypes and additional material, the main char-
(3) inner wing of pedalium slightly overhanging tentacle acters were similar to those observed in the holotype by
insertion (Fig. 5K, white arrow) and (4) each octant of the Gershwin & Gibbons (2009) (Table 3). They were also
velarium shows 2 velarial canal roots from which various identical to the ones described above for Carybdea mur-
numbers of complexly branching velarial canals arise. rayana (Table 3) and are not repeated here. The domed
apex, due to thick mesogloea with a circular constriction
below as described (Gershwin & Gibbons 2009) and foundValidity of Carybdea murrayana Haeckel, 1880 13 Fig. 1. Type material of Carybdea murrayana, collected by John Murray during the Challenger Expedition in 1876: A: Flask with type samples with original hand writing of Ernst Haeckel; Lectotype (NMH 1882.10.9.2a): B: habitus, broken in three pieces due to dissection of Haeckel; C: gastric phacellus, note bush-like shape; D: heart-shaped rhopalial niche with 1 large upper scale and 2 narrow lower scales, note broad niche channel running from niche towards bell rim; E: ripe female gonads; F: pedalium with laterally keeled pedalial canal; G: pedalial canal knee bend (dotted line) with volcano-shaped appendage (arrow); H: velarium, full octant between frenulum (f) and pedalium (p), half octant right of pedalium (p), note 2 velarial canal roots per octant with complexly branched velarial canals; Paralectotype (NMH 1882.10.9.2b): I: habitus, cut open by Haeckel; J: opened umbrella with ripe, female gonads; K: rhopalial niche with broad niche channel; L: pedalial canal knee bend (dotted line) with volcano-shaped appendage (arrow); M: pedalium; N–Q: velarium, octants, showing the diverse patterns of the complexly branched velarial canals of a single specimen. in C. branchi (Fig. 4C, E), is not visible anymore in the on the exumbrella/pedalia, the number of gastric filaments, type material of C. murrayana (Fig. 2D) but that might be and the distribution. due to the long time of preservation and the withdrawal of Otherwise, the characters listed in Table 3 show no sig- water content by the preservation medium. Haeckel (1882) nificant differences in the morphological structures of C. described and presented in his line drawings an obvious murrayana compared to the ones of C. branchi. domed apex with a circular constriction at its base (Figs. 2A, 3A1–4). Slight differences between both species can Carybdea marsupialis (Linnaeus, 1758) also be found in the original bell sizes, nematocyst warts Type locality: Mediterranean Sea (Linnaeus 1758).
14 I. Straehler-Pohl
Fig. 3. Line drawings by Haeckel (1882): A: Plate 26 of Chal-
lenger Report by Haeckel (1882); B: Enlarged part showing the
velarium, note the number of canal roots per octant marked by
Fig. 2. Comparison of type material with original pencil line
brackets and numbers; f: frenulum; p: pedalium.
drawings, partly coloured, by Haeckel (1877, with courtesy from
the collections of the Ernst-Haeckel-Haus, Friedrich-Schiller-
Universität Jena Institut für Geschichte der Medizin, Naturwis-
es in the morphological structures of Carybdea marsupia-
senschaft und Technik): A: partly coloured line drawing of habitus,
left: interradial section, middle: full outer habitus, right: perradial lis when compared to the observed West African species
section; B: line drawing of gastric phacellus and filaments; C: co- C. murrayana and C. branchi:
loured line drawing of one and a half velarial octants; D: photo of Adult medusae of C. marsupialis are much smaller than
habitus of paralectotype (NHM 1882.10.9.2b); E: photo of bush- the ones of C. murrayana or C. branchi. The rhopalial
shaped gastric phacellum of lectotype (NHM 1882.10.9.2a); F: niche openings in all three species are heart-shaped and
photo of lectotype showing the same 1.5 velarial octants that were possess just 1 upper covering scale but in C. marsupia-
drawn by Haeckel (Fig. C). lis the bottom niche channel, which runs towards the bell
rim, is narrow while it is very broad in both African spe-
As before the 19th Century the declaration of a holo- cies. While in C. branchi and C. murrayana the pedalial
type was not usual (Daston 2004), there exists no holo- canal knee bend shows a bulged, volcano-shaped append-
type, therefore, a neotype (MZB 2015-1701) was declared age (Figs. 5F, L, arrows), the pedalial canal knee bend
in 2015 (Acevedo 2016, Acevedo et al. 2019). of C. marsupialis is rounded without any appendage (Fig.
The neotype and other registered collection material 5Q, arrow). The velarium shows 3 velarial canal roots per
(Table 3) of C. marsupialis examined for this study were octant with 1–3 branched velarial canals per root, with
all in good condition. In the registered collection material sharp tips, in C. marsupialis (Fig. 5R), while the velarium
the main characters were similar to those observed in the in C. murrayana and C. branchi shows 2 velarial canal
neotype (Acevedo 2016, Acevedo et al. 2019) and identi- roots with 2–4 branched velarial canals per root and with
cal to the ones described in Table 3. They are therefore not rounded tips.
repeated here.
The characters listed in Table 3 show distinct differenc-Validity of Carybdea murrayana Haeckel, 1880 15 Fig. 4. Carybdea branchi: A: adult medusa (in-life; photo by AC Morandini); B: Holotype SAM-H4863; C: Paratype SAM-H 4864b; D: NHM 2000.1800; E: adult medusa (preserved; photo by AC Morandini); F: apex (SAM-H 4863) with gastric phacellae (white arrows); G: apex (SAM-H 4864b) with gastric phacellae (white arrows); H: epaulette-shaped gastric phacellus; I: close up on phacellus base, note clus- tered, multiple stems, originating from 1 root; J: rhopalial niche (SAM-H 4863), note large, irregularly shaped nematocyst warts; K: rhopalial niche (in-life; photo by AC Morandini); L: rhopalial niche (NHM 2000.1800), note large, irregularly shaped nematocyst warts; M: rhopalial niche (preserved), note sham “lower scales” which are a preservation artefact; N: pedalium (SAM-H 4863); M: pedalium (in-life, blue colour due to background; photo by AC Morandini), note brownish coloured spots on base and tentacle insertion, note distinct pedalial knee bend with volcano-shaped appendage (arrow), note inner wing overhanging tentacle insertion; O: pedalium (NHM 2000.1800), note inner wing overhanging tentacle insertion; P: pedalium (SAM-H 4864b); Q: pedalial canal knee bend, note volcano-shaped appendage (arrow); R: ma- nubrium (SAM-H 48064a), note long, broad, slightly frilled mouth arms (“frill” due to preservation); S: octant of velarium (SAM-H 4863; frenulum on the left border), numbers mark canal roots; T: octant of velarium (SAM-H 4864b; frenulum on the left border), numbers mark canal roots; U: quadrant of velarium (NHM 2000.1800; frenulum in the middle), numbers mark canal roots; V: C. branchi swimming next to Chirodropus gorilla in Lüderitz Bay, southern Namibia (after Straehler-Pohl (2019), p. 771, photo: Simon Elwen, Namibian Dolphin Project). f: frenulum; p: pedalium.
16 I. Straehler-Pohl Fig. 5. Comparison of Carybdea branchi (A), Carybdea murrayana (H) and Carybdea marsupialis (N). Carybdea branchi: A: habitus, sampled by AC Morandini; B: gastric phacellus, note bush-like shape. C: gastric filaments (photo by AC Morandini), note tree-like shape with fused stalks originating from one root, branching several times towards the distal end; D: heart-shaped rhopalial niche, note long and broad niche channel running from niche towards bell rim; E: pedalium, note inner pedalial wing overhanging tentacle insertion (arrow; photo by AC Morandini); F: pedalial canal knee bend with volcano-shaped appendage (white arrow); G: velarium, octant, note 2 canal roots with up to 5 velarial canals of diverse shapes, lobations and branching, canal tips rounded; Carybdea murrayana: H: habitus of paralectotype; I: gastric phacellus, note bush-like shape, note single rooted attachment, and multiple branching of filaments; J: heart-shaped rhopalial niche, note broad niche channel running from niche towards bell rim; K: pedalium, note inner pedalial wing overhanging tentacle insertion (ar- row); L: pedalial canal knee bend (dotted line) with volcano-shaped appendage (arrow); M: velarium, octant, note 2 canal roots with up to 5 velarial canals of diverse shapes, lobations and branching, canal tips rounded; Carybdea marsupialis: N: habitus, sampled by MJ Acevedo; O: gastric phacellus, note epaulette-shape; P: heart-shaped rhopalial niche (after Acevedo et al. 2019), note short, narrow niche channel; Q: pedalium, note that inner wing does not overhang tentacle insertion, note also rounded pedalial canal knee bend without appendage (arrow); R: velarium, octant, note 3 canal roots with 1 branching canal per root (after Acevedo et al. 2019); S: adult specimen, in-live, note brownish bases of gastric phacellae and nematocyst warts (photo by Eduardo Obis). f: frenulum; p: pedalium.
Validity of Carybdea murrayana Haeckel, 1880 17
Fig. 6. A: Original German text of the species description of Procharagma prototypus by Haeckel (1880, pp. 436–437). B: Original pencil
sketch by Haeckel in 1877; C: Immature medusa of Carybdea brevipedalia Kishinouye, 1891, bell height 15 mm (photo taken and provided
by Sho Toshino); D: Mature medusa of Carybdea brevipedalia, bell height 40 mm;: Translation from German to English: see main text.
Project 2: Procharagma prototypus and Procharybdis 8 sections), 4 sense clubs small, in heart-shaped nichse,
cuboides short way from bell margin. 4 tentacles simple, cylindrical,
nearly as long as bell height.
Procharagma prototypus Haeckel, 1880: (Figs. 6, 7) Special description: Procharagma prototypus, which I
Translation of original description (Haeckel 1880, pp. could examine in two well-preserved spirit (Comment by
436–437; Fig. 6A) from German to English: ISP: ethanol) samples, is of high interest as being the cubo-
“425. Species: Procharagma prototypus, Haeckel; nova medusa which shows, beyond all other known forms of this
species. order, the simplest organization in form, and which can
easily be phylogenetically derived from Tessera. The bell
Plate XXV, Figures 1–2. possesses a nearly perfect cubic shape, therefore, the flat-
Species diagnosis: Bell cubic-shaped, equally high as tened vent area has the same size and square shape as the
wide; vent area (Comment by ISP: apex) as flattened as 4 lateral planes; the sixth, oral plane of the cube is taken
the 4 lateral planes, square. Stomach forms a flat square up by the square-shaped bell opening. The 4 lateral planes
pouch, offset from the four-lobed mouth tube by a deep are marginally arched at the perradial midline and are off-
palatin-constriction; 4 linear phacellae, each split into 8 set from the rounded interradial edges only by a very flat,
filaments. Bell margin with 8 flat adradial gelatin lobes furrow-like depression.
(Comment by ISP: bell turn-over, velarium separated in The exumbrella is smooth, the gelatin of the bell is thin18 I. Straehler-Pohl Fig. 7. Original pencil line drawings (A–C), printed line drawings (D–E) and figure caption (F, in German) of Procharagma prototypus by Haeckel (1880). F: Figure caption (translated from German to English): Figure 1, 2. Procharagma prototypus, Haeckel (System, p. 436). Procharagmid of the Chinese Sea, drawn after a spirit sample from Weber. ̶Figure 1. The whole medusa, in profile, 6 times enlarged. In the middle of the figure an interradial bell edge protrudes so that 2 sides of the cubic-shaped umbrella are visible at the same time. ̶ Figure 2. Horizontal cross-section through the bell below the stomach so that the bottom of the stomach (gw = subumbrellar wall of the stomach) with the 4 perradial mouth arms (al = mouth arms (perradial)) are visible from below in their full dimension; 5 times enlarged. but solid. The subumbrella shows clearly the 4 perradial set, each split into 8 digiform filaments that decrease in longitudinal muscles which divide the 4 square-panels of size from midline of the palm towards the borders on each the ring muscles into halves and which rise from the sense side. The gonads are 8 rather narrow longish panels, their niches to the mesogonia (Comment by ISP: gonads). The free margin irregularly lobate (Comment by ISP: fold- stomach forms a flat, square pouch at the base of the bell ed); both samples were males. The bell margin bulgingly cavity; a deep palatine constriction separates it from the thickened and split into 8 flat, marginally salient, adra- short, four-sided pyramidal mouth tube, which is drawn dial lobes due to 8 shallow incisions. A short way up the 4 out into 4 short, triangular, perradial mouth arms. Inside perradial incisions, 4 sense clubs are set into flat, heart- the 4 interradial stomach corners 4 linear phacellae are shaped niches; each seems to possess next to the small,
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