Research Article
Research Article
JellyWeb: an interactive information system on Scyphozoa, Cubozoa and Staurozoa
expand article infoStefano Martellos, Luca Ukosich, Massimo Avian
‡ University of Trieste, Department of Life Sciences, Trieste, Italy
Open Access


Identification of organisms is traditionally based on the use of “classic” identification keys, normally printed on paper. These keys have several drawbacks: they are mainly based on the systematics, requiring identification of orders, families and genera at first; they are written by experts for other experts, in a specific scientific jargon; they have a “frozen” structure (sequence of theses/antitheses); once published, they cannot be changed or updated without printing a new edition. Due to the use of computers, it is now possible to build new digital identification tools, which: 1) can be produced automatically, if the characters are stored in a database; 2) can be freed from the traditional systematics, giving priority to easy-to-observe characters, incl. those usually uncommon to the classical keys, such as ecology and distribution; 3) can be updated in real time once published on-line; 4) can be available on different media, and on mobile devices. An important feature of these new digital tools is their “collaborative” nature. They can be enriched by the contribution of several researchers, which can cooperate while maintaining rights and property of the resources and data they contribute to the system. JellyWeb, the information system on Scyphozoa, Cubozoa and Staurozoa has been developed in Trieste since 2010. The system was created with the aim of – potentially – becoming a starting point for a wide collaborative effort in developing a user-friendly worldwide digital identification system for jellyfishes.


Biodiversity informatics, Cnidaria , FRIDA, identification, jellyfish, Medusozoa


Since the Rio Earth Summit in 1992, access to biodiversity information has become a fundamental task. Biodiversity data are targeted by several efforts of digitalization and aggregation, most of which focus on primary biodiversity data, i.e. natural history collection specimens and field records. Some of these efforts produced wide global networks, e.g. the GBIF (Global Biodiversity Information Facility; Berendsohn et al. 2010, King et al. 2010), which, together with the BioCASE (Biodiversity Collection Access for Europe, Holetschek et al. 2012), is mobilizing ca. 600 millions of records. Primary biodiversity data are mostly used in modeling the distribution of the taxa, and in predicting the effect of climate changes and anthropic pressure on endangered or alien invasive taxa. Taxon related information (nomenclature, auto-ecology, etc.) become the focus of similar large scale efforts only in the last years (Martellos and Attorre 2012, Martellos 2014). The GBIF itself is starting to aggregate checklist data (GBIF 2010), while other efforts are focused on molecular data (Field et al. 2011, Holetschek et al. 2012, Wieczorek et al. 2012), and to ecological information (Fegraus et al. 2005). In the field of hydrobiology, some recent examples can be Fish-SPRICH (Brosse et al. 2013) and Fish-AMAZBOL (Carvajal-Vallejos et al. 2014). In the case of jellyfishes, online resources are however scarce, but some relevant exceptions (e.g., the Jellyfish Dataset Initiative,

Digital identification keys are a particular case in the world of biodiversity informatics. Since the development of the DELTA language (Dallwitz 1980), efforts aiming at creating online digital identification keys followed several approaches. The resulting products differ in usability, accessibility, size, etc. (Nimis and Martellos 2009, Hagedorn et al. 2010, Randlane et al. 2010, Martellos and Nimis 2015). With the development of FRIDA (FRiendly IDentificAtion, Martellos 2010), the researchers of the Department of Life Sciences, University of Trieste, aimed at producing a simple but effective instrument for the development of digital identification keys in collaborative efforts. This led to the publication – in the framework of project Dryades, and of the EU projects KeyToNature (, SiiT ( and CSMON-LIFE (LIFE13 ENV/IT/842, – of ca. 600 different digital identification keys for several groups of organisms.

As far as Scyphozoa, Cubozoa and Staurozoa are concerned, there are digital databases hosting taxonomic information, such as WoRMS (World Register of Marine Species,, as well as paper printed keys to genera (as an example, see Cornelius 1997; other keys are listed in Morandini et al., 2005). Few examples of digital resources are available in the web, often limited to specific geographic regions, as the Cubozoan and scyphozoan key of the Carolinian Biogeographic Province (Calder 2009), the key to the Scyphozoa and Cubozoa of the South Atlantic Bight (Calder and King 2008), and, as far as the Mediterranean is concerned, the web site MeteoMeduse (Boero 2013, The latter, however, is an example of citizen science observatory, and does not provide an identification key. To our knowledge, no comprehensive digital identification tools to species of these taxa exist.

By combining taxonomical, ecological, and morphological and anatomical features into an information system, we developed the so called JellyWeb, a simple tool which allow to researchers and laypersons to identify Scyphozoa, Cubozoa and Staurozoa to the species level. This paper presents the results of this effort, available online at the URL


Data were collected from several sources in literature. The most relevant are Kramp (1961), WoRMS (, the Scyphozoan Wiki (, and Mills (1999-). Further sources are listed in Balboni 2008, Benci 2008, Sarto 2009, Sola 2009, Coral 2012, Benci 2012, Savonitto 2012, Ukosich 2014. Other paper are under consideration, and will lead to adding to the database other species for several genera, such as Atolla (A. russelli, A. gigantea, A. chuni), Aurelia (A. marginalis), Chironex (C. yamaguti), Cyanea (C. lamarkii, C. rosea, C. annaskala, C. tzetlinii, and several other species), Desmonema (D. comatum, D. scoresbyanna), Drymonema (D. gorgo, D. larsoni), Nausithoe (N. marginata), Pelagia (P. benovici) Tripedalia (T. binata).

The information system is freely available online at the URL It organizes data collected in the last five years by the research unit headed by Massimo Avian, at the Dept. of Life Sciences of the University of Trieste. The researchers which contributed to the project agreed on distributing the data under a Creative Commons, share alike, by attribution 3.0 (CC 3.0 by-sa) license.

The software of the information system has been developed in PHP language. The data are stored in a MySQL database. The system is equipped with a multi-entry query interface (Hagedorn et al. 2010), which operates on both a taxonomic database, and a database of nine easily recognizable morphological characters (see below). The multi-entry interface allows complex queries, which can be a first step in the identification of an organism. The multi-entry query system returns lists of taxa, on which the identification process can continue by using a digital identification system. The latter has been developed by using the FRIDA (FRiendly IDentificAtion) package (Martellos 2010). It operates on a morpho-anatomical database, which hosts ca. 200 characters for several infra-generic taxa of Scyphozoa, Cubozoa and Staurozoa (a revision of the content of the database due to recent taxonomic advancements is ongoing). The output of the digital identification system is a digital identification key to the remaining taxa, which can be used by an interactive interface, or printed out as a dichotomous, illustrated key. The whole key can also be exported in a stand-alone version for mobile devices (Nimis et al. 2012).

The query interfaces have been developed according to the results of several usability tests, conducted in the framework of projects KeyToNature and SiiT, as detailed in Martellos and Nimis (2015). The system is under continuous development, following users’ input.


JellyWeb hosts several information pages and two query system. The home page ( provides access to several sections: information, describing how the system works; survey area; query (detailed below); checklist, listing all taxa alphabetically by genus and species name, and providing access to their taxon pages; credits.

The query system is made of two parts.

  1. Multi-entry interface (Fig. 1). The first interface of the query system provides the users with the opportunity of specifying a set of nine easily observable characters, and/or scientific name and family. The morphological characters are:

    • Jellyfish sessile / swimming;

    • Umbrella shaped like a cube or a box / not shaped like a cube or a box;

    • Tentacles present / absent;

    • Tentacles isolated / grouped in clusters;

    • Umbrella with a coronal groove / without a coronal groove;

    • Umbrella flat / not flat;

    • Oral arms absent / 4 / more than 4;

    • Jellyfish with filaments (oral arm appendages) / without filaments;

    • Jellyfish with scapulae / without scapulae.

    For each character, an information popup window with images and text detailing the most relevat features is accessible by clicking on the question mark button. The result of a query is a list of taxa (Fig. 2). For each taxon an image is displayed (if available, see below). A link provides access to the taxon page (Fig. 3), which displays a description, as well as all the images available in the system, with credits and metadata, and other information (when available). Taxon pages can host a virtually unlimited amount of information and images, and/or provide access to external resources through HTML links.

  2. Digital identification key. The results page of the multi-entry interface allows to generate an interactive identification key to remaining taxa. The key can be used through a simple single entry interface (Fig. 4, Hagedorn et al. 2010), or printed out as a textual, illustrated dichotomous key. At each step of the identification process users can list out the remaining taxa, or print an illustrated key. By clicking on a taxon name, the corresponding taxon page is shown (Fig. 3). Each key generated by this system is different from the others, since they contain a different number of infra-generic taxa. Normally, the lower the number of taxa is, the easier the resulting key. A key to all the taxa currently included in our databases can also be generated, and is provided below.

Figure 1. 

Multi-entry interface. The multi-entry interface allows to combine the states of several morphologic and anatomic characters, together with taxonomic information, to query the database.

Figure 2. 

List of taxa. The result of a query made by using the multi-entry interface is an illustrated list of infra-generic taxa.

Figure 3. 

Taxon page. A typical taxon page displays an image, a description, as well as all the other images available in the system, together with credits and metadata. Taxon pages can host a virtually unlimited amount of data, links and media.

Figure 4. 

Single entry digital identification key. The digital identification key to remaining taxa is generated from the results of the multy-entry query system. It is used through a single entry interface, and can be printed out as a textual, illustrated dichotomous key as well.

Dichotomous key to all taxa

This key was automatically generated by the system, and contains all the infra-generic taxa currently stored in our databases at the date October 30, 2015. When a taxon is added to the system the key automatically changes. Hence, the key an user will obtain in the future will be slightly – or completely – different. The keys are not the transposition of an existing paper printed key, but are automatically generated by the system from a database for morphological and anatomical characters by using the package FRIDA (Martellos 2010).

1 Medusa sessile 2
Medusa swimming 34
2 (1) Medusa without aboral peduncle Lucernariopsis vanhoeffeni (Browne, 1910)
Medusa with aboral peduncle 3
3 (2) Medusa with sense organs: rhopalioids (anchors) 4
Medusa without sense organs 15
4 (3) Coronal muscle divided 5
Coronal muscle unbroken 10
5 (4) Calyx not conical 6
Calyx conical 7
6 (5) Calyx quadro-pyramidal Haliclystus borealis Uchida, 1933
Calyx pyramidal, octangular Haliclystus salpinx Clark, 1863
7 (5) Marginal anchors fairly large, egg-shaped Haliclystus stejnegeri Kishinouye, 1899
Not as above 8
8 (7) Marginal anchors very large, biscuit-shaped Haliclystus antarcticus Pfeffer, 1889
Not as above 9
9 (8) Marginal anchors kidney-shaped, with a short, cylindric stalk Haliclystus auricula (Rathke, 1806)
Marginal anchors small, oval Haliclystus kerguelensis Vanhöffen, 1908
10 (4) Peduncle single-chambered Manania hexaradiata (Broch, 1907)
Peduncle with 4 perradial chambers 11
11 (10) Gonads not united by a transverse circumferential membrane (claustrum) which divide each of the 4 perradial stomach pouches into an outer and an inner space Stenoscyphus inabai (Kishinouye, 1893)
Gonads united by a transverse circumferential membrane (claustrum) which divide each of the 4 perradial stomach pouches into an outer and an inner space 12
12 (11) Calyx as long as wide Manania gwilliami Larson & Fautin, 1989
Calyx longer than wide 13
13 (12) Calyx with dark herringbone pattern Manania distincta (Kishinouye, 1910)
Calyx without dark herringbone pattern 14
14 (13) Arms twice as long as broad Halimocyathus platypus Clark, 1863
Arms short Manania handi Larson & Fautin, 1989
15 (3) Peduncle with 4 perradial chambers 16
Peduncle single-chambered 22
16 (15) Peduncle with muscle in the septa 17
Peduncle without muscle in the septa 18
17 (16) On each arm about 9 tentacles Depastrum cyathiforme (M. Sars, 1846)
On each arm about 25 tentacles Depastromorpha africana Carlgren, 1935
18 (16) Gonads united by a transverse circumferential membrane (claustrum) which divide each of the 4 perradial stomach pouches into an outer and an inner space 19
Gonads not united by a transverse circumferential membrane (claustrum) which divide each of the 4 perradial stomach pouches into an outer and an inner space 20
19 (18) On each arm 60–80 tentacles Craterolophus convolvulus (Johnston, 1835)
On each arm about 30 tentacles Craterolophus macrocystis von Lendenfeld, 1884
20 (18) Arms adradial Kishinouyea nagatensis (Oka, 1897)
Arms interradial 21
21 (20) Arms larger at base than S. tsingtaoensis Sasakiella cruciformis Okubo, 1917
Arms narrower at base than S. cruciformis Sasakiella tsingtaoensis Ling, 1937
22 (15) Peduncle without muscle in the septa Lucernariopsis campanulata (Lamouroux, 1815)
Peduncle with muscle in the septa 23
23 (22) Marginal lobes (arms) faintly developed 24
Marginal lobes (arms) well developed 26
24 (23) Tentacles reduced Lipkea stephensoni Carlgren, 1933
Not as above 25
25 (24) Tentacles not true Lipkea ruspoliana Vogt, 1886
Tentacles rudimentary Lipkea sturdzi (Antipa, 1893)
26 (23) Tentacles up to 60 on each arm 27
Tentacles more than 60 on each arm 28
27 (26) Subumbrellar margin with 4 perradial pigment spots Stylocoronella riedli Salvini-Plawen, 1966
Subumbrellar margin without 4 perradial pigment spots Stylocoronella variabilis Salvini-Plawen, 1987
28 (26) Peduncle rudimentary Lucernaria australis Vanhöffen, 1908
Peduncle true 29
29 (28) Peduncle as long or longer than height of calyx 30
Peduncle shorter than height of calyx 31
30 (29) Tentacles 100–140 on each arm Lucernaria quadricornis O.F.Müller, 1776
Tentacles 700–850 on each arm Lucernaria walteri (Antipa, 1892)
31 (29) Tentacles 80 or less on each arm Lucernaria infundibulum Haeckel, 1880
Tentacles more than 80 on each arm 32
32 (31) Peduncle 1/3 as long as height of calyx Lucernaria haeckeli (Antipa, 1892)
Not as above 33
33 (32) Peduncle less than 1/3 of the height of calyx Lucernaria bathyphila Haeckel, 1880
Peduncle about half as long as height of calyx Lucernaria sainthilairei (Redikorzev, 1925)
34 (1) Medusa with calix Tesserantha connectens, Haeckel, 1880 – Warning: some authors debate on the validity of swimming Stauromedusae (see Rodriguez et al. 2011)
Medusa with umbrella 35
35 (34) Exumbrella divided by a circular and deep coronal groove 36
Exumbrella not divided by a circular and deep coronal groove 64
36 (35) Tentacles from 4 to 6 37
Tentacles 8 or more 42
37 (36) Rhopalia 4 38
Rhopalia 6 39
38 (37) Gonads almost equidistant Pericolpa campana (Haeckel, 1880)
Gonads in 4 pairs Pericolpa quadrigata Haeckel, 1880
39 (37) Gonads 6 Atorella arcturi Bigelow, 1928
Not as above 40
40 (39) Gonads 8 Atorella octogonus Mills, Larson & Young, 1987
Gonads 4 41
41 (40) Gonads sac-like, swollen Atorella subglobosa Vanhöffen, 1902
Gonads leaf-shaped Atorella vanhoeffeni Bigelow, 1909
42 (36) Rhopalia up to 6 43
Rhopalia more than 6 48
43 (42) Rhopalia perradial, 4 44
Rhopalia interradial, 4 45
44 (43) Coronal muscle divided Paraphyllina intermedia Maas, 1903
Coronal muscle unbroken Paraphyllina ransoni Russel, 1956
45 (43) Marginal lappets 32 Nauphantopsis diomedeae Fewkes, 1885
Not as above 46
46 (45) Gonads 4 Periphyllopsis galatheae Kramp, 1959
Gonads 8 47
47 (46) Marginal lappets 16 Periphylla periphylla (Péron & Lesueur, 1809)
Marginal lappets 24 Periphyllopsis braueri Vanhöffen, 1902
48 (42) Gonads 4 or 4 pairs 49
Gonads 8 53
49 (48) Stomach pouches break up into numerous ragged-edged branches in the marginal lappets 50
Stomach pouches simple, radiating 51
50 (49) Subumbrellar protuberances in 2 circles Linuche aquila Mayer 1910
Subumbrellar protuberances in 3 circles Linuche unguiculata (Schwartz, 1788)
51 (49) Gonads bean-shaped Palephyra indica Vanhöffen, 1902
Gonads crescent-shaped 52
52 (51) Gonads with horns recurved Palephyra antiqua Haeckel, 1880
Gonads consisting of 3 swellings Palephyra pelagica Haeckel, 1880
53 (48) Rhopalia > 8 54
Rhopalia 8 (Genus Nausithoe. The key refers to free-swimming stages only) 56
54 (53) Gastric ostia with two pigmented spots Atolla vanhoeffeni Russell, 1957
Gastric ostia without pigmented spots 55
55 (54) Species with 20–24 tentacles Atolla parva Russell, 1958
Species with usually 22, sometimes up to 32 tentacles Atolla wyvillei Haeckel, 1880
56 (53) Central disk with large pits 57
Central disk without pits 58
57 (56) Central disk with radiating furrows Nausithoe rubra Vanhöffen, 1902
Central disk without radiating furrows Nausithoe atlantica Broch, 1914
58 (56) Gonads very small Nausithoe clausi Vanhöffen, 1892
Not as above 59
59 (58) Gonads of normal dimensions Nausithoe albatrossi (Maas, 1897)
Gonads large 60
60 (59) Central disk not thick nor finely punctured Nausithoe globifera Broch, 1914
Central disk thick, finely punctured 61
61 (60) Central disk with radiating furrows Nausithoe challengeri (Haeckel, 1880)
Central disk without radiating furrows 62
62 (61) Medusa with chocolate brown or carmine gonads and blue gastric cirri Nausithoe picta Agassis & Mayer, 1902
Medusa without chocolate brown or carmine gonads and blue gastric cirri 63
63 (62) Gastric cirri not grouped in clusters Nausithoe punctata (Kölliker, 1853)
Gastric cirri grouped in clusters Nausithoe limpida Hartlaub, 1909
64 (35) Opening of the subumbrellar cavity partly closed by an annular diaphragm (velarium) 65
Opening of the subumbrellar cavity not closed by an annular diaphragm (velarium) 88
65 (64) Tentacles 8 or more 66
Tentacles from 4 to 6 76
66 (65) Stomach pouches without diverticula Tripedalia cystophora Conant, 1897
Stomach pouches with 8 diverticula 67
67 (66) Gonads not four-leaved Chirodectes maculatus (Cornelius, Fenner & Hore, 2005)
Gonads four-leaved 68
68 (67) Medusa with nematocysts on bell 69
Medusa without nematocysts on bell 71
69 (68) Each pedalium with more than 4 fingers and tentacles Chiropsalmus quadrumanus Müller, 1859
Each pedalium with 4 or less fingers and tentacles 70
70 (69) Each pedalium with 2 fingers and tentacles Chiropsalmus zygonema Haeckel, 1880
Each pedalium with 3–4 fingers and tentacles Chiropsalmus alipes Gershwin, 2006
71 (68) Medusa with mesenteries poorly developed Chiropsella bronzie Gershwin, 2006
Not as above 72
72 (71) Gastric saccules are functioning gonads Chironex fleckeri Southcott, 1956
Gastric saccules are not functioning gonads 73
73 (72) Stomach pouches with 2 branched or feathered saccules 74
Stomach pouches with 2 unbranched saccules 75
74 (73) Each pedalium with 9–11 fingers and tentacles Chirodropus gorilla Haeckel, 1880
Each pedalium with 21 fingers and tentacles Chirodropus palmatus Haeckel, 1880
75 (73) Tentacles and fingers irregularly placed Chiropsoides buitendijki (van der Horst, 1907)
Tentacles and fingers not irregularly placed Chiropsoides quadrigatus (Haeckel, 1880)
76 (65) Tentacles branched Manokia stiasnyi Bigelow, 1938
Tentacles simple 77
77 (76) Stomach with weakly developed mesenteries 78
Not as above 80
78 (77) Sensory niches without well developed covering scale Copula sivickisi Stiasny, 1926
Sensory niches with covering scale above 79
79 (78) Velarial canals 3–4 per octant Carybdea marsupialis (Linnaeus, 1758)
Velarial canals 2 per octant Carybdea rastonii Haacke, 1886
80 (77) Stomach without mesenteries 81
Stomach with well developed mesenteries 84
81 (80) Exumbrella without nematocyst-warts Alatina moseri (Mayer, 1906)
Exumbrella with nematocyst-warts 82
82 (81) Velarial canals 3 per octant Alatina rainensis Gershwin ,2005
Velarial canals 4–5 per octant 83
83 (82) Medusa with 6 eyes per rhopalium Alatina madraspartana Menon, 1930
Medusa with 1 eye per rhopalium Alatina tetraptera (Haeckel, 1880)
84 (80) Medusa with phacellae Tamoya haplonema Müller, 1859
Medusa without phacellae 85
85 (84) Velarial canals 1 per octant Carukia shinju Gershwin, 2005
Not as above 86
86 (85) Velarial canals 2 per octant Carukia barnesi Southcott, 1966
Not as above 87
87 (86) Velarial canals 4–5 per octant Malo maxima Gershwin, 2005
Velarial canals more than 5 per octant Gerongia rifkinae Gershwin & Alderslade, 2005
88 (64) Medusa with a permanent primary mouth opening in adult specimens 89
Medusa without a permanent primary mouth opening in adult specimens 135
89 (88) Medusa without tentacles 90
Medusa with tentacles 94
90 (89) marginal lappets very shallow, or entirely lacking 91
marginal lappets evident 92
91 (90) Exumbrella transparent white, sometimes with brown nuances on margins Deepstaria enigmatica (Russel, 1967)
Exumbrella reddish-brown, with stomach margin lighter brown Deepstaria reticulum (Lerson, Madin & Harbison, 1988)
92 (90) Rhopalia from 24 to more than 50, one in every cleft between the lappets Tiburonia granrojo (Matsumoto, Raskoff & Lindsay, 2003)
Not as above 93
93 (92) Rhopalia 20 Stygiomedusa gigantea (Browne, 1910)
Rhopalia 8 Stellamedusa ventana (Raskoff & Matsumoto, 2004)
94 (89) Medusa with ring-canal 95
Medusa without ring-canal 113
95 (94) Tentacles arising from umbrella’s margin 96
Tentacles not arising from umbrella’s margin 104
96 (95) Marginal lappets 48 Undosa undulata (Stiasny, 1935) – Warning: dubious species, some authors suggest it is a juvenile stage of Discomedusa lobata Claus 1877
Not as above 97
97 (96) Marginal lappets 16 98
Not as above 99
98 (97) Oral arms broad, egg-shaped Ulmaris prototypus (Haeckel, 1880)
Oral arms narrow and pointed Ulmaris snelliusi (Stiasny, 1935)
99 (97) Marginal lappets 32 100
Marginal lappets 64 102
100 (99) Tentacles 32 or 48 Discomedusa lobata (Claus, 1877)
Tentacles 24 101
101 (100) Perradial canals branched Discomedusa philippina (Mayer, 1910)
Perradial canals not branched Floresca parthenia (Haeckel, 1880)
102 (99) Tentacles 24 Parumbrosa polylobata (Kishinouye, 1910)
Tentacles 16 103
103 (102) Anastomoses absent Diplulmaris antarctica (Maas, 1908)
Anastomoses present Diplulmaris malayensis (Stiasny, 1935)
104 (95) Tentacles arising from subumbrella 105
Tentacles arising from exumbrella 107
105 (104) Gonads 8 Poralia rufescens (Vanhöffen, 1902)
Gonads 4 106
106 (105) Rhopalia 8 Sthenonia albida (Eschscholtz, 1829)
Rhopalia 16 Phacellophora camtschatica (Brandt, 1835)
107 (104) Oral arms bifurcated Aurosa furcata (Haeckel, 1880)
Oral arms not bifurcated 108
108 (107) Marginal lappets 16 Aurelia labiata (Chamisso & Eysenhardt, 1821)
Marginal lappets 8 109
109 (108) Oral arms short, thick and curved, much folded, extending laterally against subumbrellar surface Aurelia limbata (Brandt, 1835)
Not as above 110
110 (109) Oral arms linear, thick and stiff, with densely crenulated margins, as long as bell’s radius Aurelia aurita (Linnaeus, 1758)
Not as above 111
111 (110) Oral arms narrow and thin, with slightly folded margins only in proximal part Aurelia solida (Browne, 1905)
Oral arms long and broad, curtain-like, with densely crenulated margins 112
112 (111) Adradial canals not branched Aurelia maldivensis (Bigelow, 1904)
Adradial canals branched Aurelia colpata (Brandt, 1838)
113 (94) Tentacles arising from the subumbrella at some distance from the margin 114
Tentacles arising from the exumbrellar margin 122
114 (113) Tentacles not arranged in tufts Drymonema dalmatinum (Haeckel, 1880)
Tentacles arranged in tufts 115
115 (114) Medusa without radial muscolature in the subumbrella 116
Medusa with radial muscolature in the subumbrella 118
116 (115) Medusa with few broad canals in the lappets Desmonema gaudichaudi (Lesson, 1830)
Medusa with numerous narrow canals in the lappets 117
117 (116) Tentacles not ribbon-like Desmonema chierchianum (Vanhöffen, 1888)
Tentacles ribbon-like Desmonema glaciale (Larson, 1986)
118 (115) Rhopalar and tentacular stomach pouches completely separated 119
Rhopalar and tentacular stomach pouches connected by anastomoses 120
119 (118) Peripheral canals without, or with few anastomoses Cyanea capillata (Linnaeus, 1758)
Peripheral canals with numerous anastomoses Cyanea purpurea (Kishinouye, 1910)
120 (118) Peripheral canals with numerous anastomoses Cyanea nozakii (Kishinouye, 1891)
Peripheral canals without, or with few anastomoses 121
121 (120) Radial muscles originating from the outer side of coronal muscle Cyanea buitendijki (Stiasny, 1919)
Radial muscles originating from the middle of coronal muscle Cyanea mjobergi (Stiasny, 1921)
122 (113) Stomach pouches 32 123
Stomach pouches 16 124
123 (122) Subgenital pits heart-shaped Sanderia malayensis (Goette, 1886)
Subgenital pits horseshoe-shaped Sanderia pampinosus (Gershwin & Zeidler, 2008)
124 (122) Marginal lappets 16 125
Not as above 126
125 (124) Nematocyst warts about as long as wide Pelagia noctiluca (Forsskål, 1775)
Nematocyst warts highly protrusive, more long than wide Pelagia flaveola (Eschscholtz, 1829)
126 (124) Marginal lappets 48 127
Marginal lappets 32 129
127 (126) Tentacles all alike Chrysaora fulgida (Reynaud, 1830)
Tentacles different in length 128
128 (127) Tentacles usually 5 per octant, 1 central primary, 2 lateral secondary about half in length, 2 tertiary, between former two types, about 1/4 as long as the median Chrysaora lactea (Eschscholtz, 1829)
Tentacles 5 per octant, 3 primary arising from deep cleft between tentacular lappets and 2 lateral and shorter secondary, arising from subumbrellar side of rhopalar lappets Chrysaora quinquecirrha (Desor, 1848)
129 (126) Tentacles 8 Chrysaora colorata (Russel, 1964)
Tentacles 24 130
130 (129) Exumbrella yellowish-brown or reddish-yellow with 32-rayed chestnut-brown star Chrysaora fusescens (Brandt, 1835)
Not as above 131
131 (130) Exumbrella reddish-brown or purplish-pink with 16 broad, darker radial bands and numerous light spots Chrysaora plocamia (Lesson, 1830)
Not as above 132
132 (131) Oral arms extremely large with frilly margins, hardly coiled to form a dense mass Chrysaora achlyos (Martin, Gershwin, Burnett, Cargo & Bloom, 1997)
Oral arms linear, with broad frilly margins, more or less coiled around central body 133
133 (132) Exumbrella golden-brown, with darker margins, sometimes with 16-32 lighter radial stripes Chrysaora fuscescens (Brandt, 1835)
Not as above 134
134 (133) Stomach pouches all-alike Chrysaora hysoscella (Linnaeus, 1766)
Stomach pouches unequal, tentacular ones slightly broader proximally and distally than rhopalar ones Chrysaora melanaster (Brandt, 1838)
135 (88) Umbrella with papillar knobs 136
Umbrella without papillar knobs 138
136 (135) Oral arms without filaments Lobonemoides sewelli Rao, 1931
Oral arms with filaments 137
137 (136) Intracircular anastomosing network in communication with the inter-rhopalar canals Lobonema smithii Mayer, 1910
Intracircular anastomosing network not in communication with the inter-rhopalar canals Lobonemoides robustus Stiasny, 1920
138 (135) Oral arms dichotomous 139
Oral arms three-winged 159
139 (138) Medusa with 4 completely separated subgenital cavities 140
Medusa with 4 not completely separated subgenital cavities 147
140 (139) Oral arms 3/4 the lenght of bell radius Cassiopea frondosa (Pallas, 1774)
Not as above 141
141 (140) Oral arms cylindrical, slender, somewhat longer than bell radius Cassiopea ornata Haeckel, 1880
Not as above 142
142 (141) Oral arms very large, flat, with 6-8 short, wide-spreading main branches Cassiopea depressa Haeckel, 1880
Not as above 143
143 (142) Oral arms 1 1/4 times the lenght of bell radius, triangular in cross-section, aboral surface broad and flat, with 10–15 alternate primary branches Cassiopea xamachana Bigelow, 1892
Not as above 144
144 (143) Oral arms wide, flat, with 4–6 flat, short tree-shaped side branches Cassiopea andromeda (Forskål, 1775)
Not as above 145
145 (144) Oral arms with numerous small lateral branches in their proximal portion Cassiopea medusa Light, 1914
Oral arms cylindrical, 1 1/2 times as long as bell radius, branched tree-like 146
146 (145) Species with numerous large club-shaped vesicles Cassiopea mertensi Brandt, 1838
Species without ribbon-like filaments Cassiopea ndrosia Agassiz & Mayer, 1899
147 (139) Oral arms without filaments 148
Oral arms with filaments 150
148 (147) Exumbrella without a central rised dome Marivagia stellata (Galil & Gershwin, 2010)
Exumbrella with a central rised dome 149
149 (148) More than 1 cupolar warts Netrostoma dumokuroa (Agassiz & Mayer, 1899)
1 cupolar wart Netrostoma nuda (Gershwin & Zeidler, 2008)
150 (147) In each octant 3 radial canals 151
In each octant more than 3 radial canals 153
151 (150) Between the mouths two kinds of appendages Netrostoma coerulescens Maas, 1903
Between the mouths numerous appendages 152
152 (151) Exumbrella with a central rised dome Netrostoma setouchianum (Kishinouye, 1902)
Exumbrella without a central rised dome Cephea octostyla (Forskål, 1775)
153 (150) Exumbrella without a central rised dome Polyrhiza vesiculosa (Agassiz, 1862)
Exumbrella with a central rised dome 154
154 (153) Medusa with warts on the central portion of the exumbrella 155
Medusa without warts on the central portion of the exumbrella 156
155 (154) Radial canals 5–6 per ottante Cephea cephea (Forskål, 1775)
Radial canals 7 per ottante Cephea coerulea Vanhöffen, 1902
156 (154) In each octant 4–6 radial canals Cotylorhiza erythraea Stiasny, 1920
Not as above 157
157 (156) In each octant 7–9 radial canals Cotylorhiza tuberculata (Macri, 1778)
In each octant more than 11 radial canals 158
158 (157) Radial canals 11–13 per ottante Cotylorhiza ambulacrata Haeckel, 1880
Radial canals 16–17 per ottante Cotylorhiza pacifica (Mayer, 1915)
159 (138) Oral arms triangular 160
Oral arms not triangular 162
160 (159) Oral arms terminate in a short, oval knob Thysanostoma loriferum (Ehrenberg, 1835)
Not as above 161
161 (160) Oral arms terminate in a long, tapering filament Thysanostoma flagellatum (Haeckel, 1880)
Oral arms without a terminal portion Thysanostoma thysanura Haeckel, 1880
162 (159) Oral arms not pyramidal 163
Oral arms pyramidal 181
163 (162) Oral arms broad 164
Oral arms of normal width 169
164 (163) Oral arms without filaments 165
Oral arms with filaments 166
165 (164) Oral arms without terminal clubs Lychnorhiza malayensis Stiasny, 1920
Oral arms with terminal clubs Pseudorhiza aurosa von Lendenfeld, 1882
166 (164) Oral arms with terminal clubs 167
Oral arms without terminal clubs 168
167 (166) Medusa without a single filament at the distal end of one of the oral arms Anomalorhiza shawi Light, 1921
Medusa with a single filament at the distal end of one of the oral arms Pseudorhiza haeckeli Haacke, 1884
168 (166) In each octant 8 velar lappets Lychnorhiza arubae Stiasny, 1920
In each octant 4 velar lappets Lychnorhiza lucerna Haeckel, 1880
169 (163) Oral arms coalesced throughout their entire length 170
Oral arms coalesced in proximal portion only 171
170 (169) Velar lappets about 14 in each octant Stomolophus meleagris Agassiz, 1862
Velar lappets about 24 in each octant Stomolophus fritillaria Haeckel, 1880
171 (169) Oral arms with filaments 172
Oral arms without filaments 175
172 (171) Umbrella more than 100 cm wide Nemopilema nomurai (Kishinouye, 1922)
Umbrella less than 100 cm wide 173
173 (172) Velar lappets 14–20 in each octant Rhopilema esculentum Kishinouye, 1891
Velar lappets 8 in each octant 174
174 (173) Exumbrella with sharply conical warts Rhopilema hispidum (Vanhöffen, 1888)
Exumbrella with blunt tuberculation Rhopilema nomadica Galil, Spanier & Ferguson, 1990
175 (171) Oral arms with clubs 176
Oral arms without clubs 177
176 (175) Velar lappets 14–20 in each octant Rhopilema rhopalophorum Haeckel, 1880
Velar lappets 6 in each octant Rhopilema verrilli (Fewkes, 1887)
177 (175) Oral arms without terminal clubs 178
Oral arms with terminal clubs 179
178 (177) Umbrella ca. 150 mm wide; marginal lobes rectagular in shape Eupilema scapulare Haeckel, 1880
Umbrella ca. 400 mm wide; marginal lobes triangular in shape Eupilema inexpectata (Pages, Gili & Bouillon, 1992)
179 (177) Proximal portion of oral arms considerably longer than distal portion Rhizostoma luteum (Quoy & Gaimard, 1827)
Proximal portion of oral arms about as long as distal portion 180
180 (179) Taxon present in the Mediterranean and in the Atlantic Ocean Rhizostoma pulmo (Macri, 1778)
Taxon present in the North Sea only Rhizostoma octopus (Macri, 1778)
181 (162) Oral arms shorter than usual 182
Oral arms of normal lenght 195
182 (181) Oral arms without terminal appendages 183
Oral arms with terminal appendages 184
183 (182) Medusa with rhopalar canals with anastomoses throughout thier length Mastigietta palmipes (Haeckel, 1880)
Medusa with perradial rhopalar canals without anastomoses, interradial canals with anastomoses Versuriga anadyomene (Maas, 1903)
184 (182) Intracircular mesh-work of canals never communicating with the rhopalar canals 185
Intracircular mesh-work of canals usually communicating with the rhopalar canals 187
185 (184) Terminal appendages nearly as long as the oral arms Phyllorhiza pacifica (Light, 1921)
Terminal appendages very long, with distal expansion 186
186 (185) Oral filaments without a triple heart-shaped knob; bell diameter far larger than 25 cm Phyllorhiza punctata (von Lendenfeld, 1884)
Oral filaments with a triple heart-shaped knob; bell of ca. 25 cm of diameter Phyllorhiza peronlesueuri (Goy, 1990)
187 (184) Mouth arms twice as long as disk radius 188
Not as above 189
188 (187) In each octant more than 10 canal-roots Mastigias pantherinus Haeckel, 1880
In each octant up to 10 canal-roots Mastigias siderea Chun, 1896
189 (187) Mouth arms shorter than disk radius 190
Mouth arms long as disk radius 192
190 (189) In each octant more than 10 canal-roots Mastigias ocellatus (Modeer, 1791)
In each octant up to 10 canal-roots 191
191 (190) Vaulted bell, thin at margin but very thick at apex Mastigias gracilis (Vanhöffen, 1888)
Doubtful species, flat and hat-shaped bell, average size unknown Mastigias roseus (Reynaud, 1830)
192 (189) In each octant up to 10 canal-roots 193
In each octant more than 10 canal-roots 194
193 (192) Umbrella not flat Mastigias papua (Lesson, 1830)
Umbrella flat, disk-shaped Phyllorhiza luzoni Mayer, 1915
194 (192) Perradial rhopalar canals not bottle-shaped Mastigias albipunctatus Stiasny, 1920
Perradial rhopalar canals bottle-shaped Mastigias andersoni Stiasny, 1926
195 (181) Oral arms with filaments 196
Oral arms without filaments 199
196 (195) Intracircular anastomosing network not in communication with the rhopalar canals 197
Intracircular anastomosing network in communication with the rhopalar canals 198
197 (196) Distal three-winged portion of oral arms about twice as long as proximal simple portion Crambione bartschi (Mayer, 1910)
Distal three-winged portion of oral arms as long as proximal simple portion Crambione mastigophora Maas, 1903
198 (196) Oral arms narrow with short filaments Acromitus flagellatus (Maas, 1903)
Oral arms thick and broad with long filaments Acromitus maculosus Light, 1914
199 (195) Oral arms with terminal clubs 200
Oral arms without terminal clubs 202
200 (199) In each octant 10 velar lappets Leptobrachia leptopus (Chamisso & Eysenhardt, 1821)
Not as above 201
201 (200) In each octant 16 velar lappets Crambionella orsini (Vanhoffen, 1888)
In each octant 12 velar lappets Crambionella stuhlmanni (Chun, 1896)
202 (199) Intracircular anastomosing network not in communication with the rhopalar canals 203
Intracircular anastomosing network in communication with the rhopalar canals 204
203 (202) In each octant 4 cleft velar lappets Acromitoides purpurus (Mayer, 1910)
In each octant at least 5 cleft velar lappets Acromitoides stiphropterus (Schultze, 1897)
204 (202) Distal three-winged portion of oral arms 1/6 as long as proximal simple portion Catostylus mosaicus (Quoy & Gaimard, 1824)
Not as above 205
205 (204) Distal three-winged portion of oral arms 6 times as long as proximal simple portion Catostylus perezi Ranson, 1945
Not as above 206
206 (205) Distal three-winged portion of oral arms 5 times as long as proximal simple portion Catostylus viridescens (Chun, 1896)
Not as above 207
207 (206) Distal three-winged portion of oral arms half as long as proximal simple portion Catostylus tripterus (Haeckel, 1880)
Not as above 208
208 (207) Distal three-winged portion of oral arms as long as proximal simple portion Catostylus ornatellus (Vanhöffen, 1888)
Distal three-winged portion of oral arms 2–4 times as long as proximal simple portion 209
209 (208) Oral arms 2/3 the length of bell diameter Catostylus townsendi Mayer, 1915
Not as above 210
210 (209) Oral arms 1–1,5 times the length of bell radius Catostylus cruciatus (Lesson, 1830)
Oral arms as long as bell diameter Catostylus tagi (Haeckel, 1869)


Digital resources on biodiversity can be relevant not only to researchers, but also to laypeople, such as tourists or citizen scientists. The importance of involving citizens in understanding, monitoring and protecting biodiversity has been recently expressed by the European Commission, in the document “Establishing Horizon 2020” (EU Regulation no. 1291/2013). However, most of the biodiversity-related resources available in the Web – especially the ones dedicated to to “difficult” groups, such as jellyfish – are normally devoted almost exclusively to experts (Martellos and Nimis 2015). Exposing scientific information in a form which can be accessible to everybody – without losing its content and informative value – can be a true revolution. Many citizens, especially if already interested in nature and aware of environmental issues (e.g. the presence of invasive alien species), are potentially interested in similar resources. Hence digital resources can be used to involve a wider amount of citizens in scientific tasks, such as the collection of those “big data” which are nowadays fundamental to researchers. The examples of OPAL initiative in the British Isles (; accessed 08 August 2015) or, in the field of jellyfish, of MeteoMedusa (Boero 2013, Boero et al. 2013), and JellyWatch (; accessed 08 August 2015) are demonstrating the effectiveness of a citizen science approach in collecting scientific data.

JellyWeb is based on morpho-anatomic and taxonomic data, collected and organized in ca. 10 years of research. The development of the portal (Martellos and Nimis 2015) was based upon the experience of the European project KeyToNature (mainly devoted to digital identification) and of the project Dryades (devoted to the pubblication of biodiversity data in the web). This is the first portal devoted to organisms other than vascular plants developed by the research unit of the Dept. of Life Science of the University of Trieste. During its development, a particular attention was paid to user interfaces, in order to provide high quality scientific information in the most straightforward way, and to make it useable by the wider audience as possible.

The multi-entry interface can be useful to both researchers (whom can simply type the name of a taxon to retrieve related information or generate an identification key), and laypeople (whom can use it to start the identification of a jellyfish they have just seen on the seashore). As a further help, interactive keys are enriched by images and drawings of the most relevant characters. Since digital keys are generated in real time, on the basis of the list of remaining organisms, each query produces a different identification key.

Since identification is nowadays often based on molecular analysis, the system has been developed to host molecular data as well. In fact, several attempts to revise the taxonomy of the various taxa like the Discomedusae on the basis of morphological observations integrated with genetic analysis are underway, highlighting several critical points, such as the recognition of cryptic species in the Aurelia complex within the “traditional” species Aurelia aurita (Dawson and Jacobs 2001, Dawson and Martin 2001, Dawson 2003, Dawson et al. 2005, Ramšak et al. 2012), or even at higher taxonomic levels like the proposition of at least two new families within the Semaeostomeae(Bayha and Dawson 2010, Straehler-Pohl et al. 2011). The integration of molecular information in a digital identification system by using the FRIDA software was studied by Bruni et al. (2012) for vascular plants.


JellyWeb is an accumulative system, which can potentially host all data on Scyphozoa, Cubozoa and Staurozoa, and even extend its aim to other groups of the phylum Cnidaria. However, a research group alone can hardly complete such a challenging task. The research unit at the University of Trieste plans to maintain and enrich JellyWeb, but its growth could be faster, if other research groups join this effort. A researcher, or a research group, can contribute to the system by:

  • Fostering a taxon (such as a genus, or a family). This can be done by managing an instance of the FRIDA system. FRIDA allows to different authors to independently manage separate instances, while at the same time contributing to the same database of morphological ans anatomical data, hence, generating updated multi-authored keys to any subset of taxa in the whole system (for a complete description see Martellos 2010). All the digital keys which are generated by the system give credit to the authors of all the data. The keys and all the data and images in JellyWeb are always distributed under a Creative Commons share alike, by attribution 3.0 license (CC 3.0 by-sa).

  • Contributing to the image archive. High quality images of morphological and anatomical characters and of the whole organisms are probably the most relevant bottlenecks in the process of creating a portal such as JellyWeb. Especially when identifing a taxon, digital images are of capital relevance, both for choosing among the leads of each choice, and as visual census when an identification has been achieved. Several species of Scyphozoa, Cubozoa and Staurozoa are known for one or few specimens, and, even when the taxa are well known, high quality images are, however, scarce. JellyWeb was developed to host a virtually unlimited number of images for each taxon. Each image is displayed with credits to the author(s) and owner(s), institution(s), other metadata, and license.

  • Producing descriptions. Another relevant bottleneck in developing digital identification keys and portals to one or more groups of organisms are their descriptions. While taxonomic descriptions can be found in books and papers, descriptions which could be actually useful to people other than researches are difficult to produce. In our experience, to be appreciated by a wider audience, they should mix different sources of information, from ecology to taxonomy, from distribution to human uses, relevance for economy, etymology of the name, etc. Hence, their production is not a simple cut and paste, but a relevant effort of analysis and synthesis.

Potential contributor can contact Massimo Avian (, to define the extent of their participation.


The authors are grateful to all the researchers and students (Balboni, Benci, Coral, Sarto, Savonitto, Sola) who contributed to this study. We are also grateful to Dr. Rodolfo Riccamboni, who developed the code of the multi-entry interface of the portal, and to Prof. Pierluigi Nimis, for his suggestions and comments. We are also grateful do Mrs. Sara Triulzi for English language check.


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