Research Article |
Corresponding author: Stefano Martellos ( martelst@units.it ) Academic editor: Leen van Ofwegen
© 2016 Stefano Martellos, Luca Ukosich, Massimo Avian.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Martellos S, Ukosich L, Avian M (2016) JellyWeb: an interactive information system on Scyphozoa, Cubozoa and Staurozoa. ZooKeys 554: 1-25. https://doi.org/10.3897/zookeys.554.6745
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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;
Digital identification keys are a particular case in the world of biodiversity informatics. Since the development of the DELTA language (
As far as Scyphozoa, Cubozoa and Staurozoa are concerned, there are digital databases hosting taxonomic information, such as WoRMS (World Register of Marine Species, http://www.marinespecies.org/), as well as paper printed keys to genera (as an example, see
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 http://dryades.units.it/jelly.
Data were collected from several sources in literature. The most relevant are Kramp (1961), WoRMS (http://www.marinespecies.org/), the Scyphozoan Wiki (http://thescyphozoan.ucmerced.edu/), and
The information system is freely available online at the URL http://dryades.units.it/jelly. 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 (
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
JellyWeb hosts several information pages and two query system. The home page (http://dryades.units.it/jelly) 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.
Multi-entry interface (Fig.
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.
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.
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 (
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 |
– | 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 (
JellyWeb is based on morpho-anatomic and taxonomic data, collected and organized in ca. 10 years of research. The development of the portal (
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 (
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
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 (avian@units.it), 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.