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Ernst Haeckel's Mysterious Species, Part I: the Validity of Carybdea

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Ernst Haeckel's Mysterious Species, Part I: the Validity of Carybdea Plankton Benthos Res 15(1): 1–29, 2020 Plankton & Benthos Research © The Plankton Society of Japan Ernst Haeckel’s mysterious species, Part I: the validity of Carybdea murrayana Haeckel, 1880 (Cubomedusae) and revisional notes on Haeckel’s other Carybdeidae ILKA STRAEHLER-POHL* Medusa(‘)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 Darwin’s 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 & Haeckel’s 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 Tableobserved 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 in their Carybdeidae species list. Morphological comparison 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
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