Morula (gastropod)

Morula
	A dorsal view of a shell of Morula striata
Scientific classification
Domain:	Eukaryota
Kingdom:	Animalia
Phylum:	Mollusca
Class:	Gastropoda
Subclass:	Caenogastropoda
Order:	Neogastropoda
Superfamily:	Muricoidea
Family:	Muricidae
Subfamily:	Ergalataxinae
Genus:	Morula; Schumacher, 1817
Type species
	Morula papillosa Schumacher, 1817;
Synonyms[1]
	Drupa (Morula) Schumacher, 1817; Habromorula Houart, 1995 (original rank); Morula (Habromorula) Houart, 1995· accepted, alternate representation; Morula (Morula) Schumacher, 1817· accepted, alternate representation; Pentadactylus (Morula) Schumacher, 1817;

Morula is a genus of small, predatory sea snails, marine gastropod mollusks in the subfalily Ergalataxinae of the family Muricidae, the murex snails or rock snails.[1]

Species

Species within the genus Morula include:

Morula aglaos (Houart, 1995)[2]
Morula albanigra (Houart, 2002)[3]
Morula ambrosia (Houart, 1995)[4]
Morula angulata (G.B. Sowerby III, 1894)[5]
Morula aspera (Lamarck, 1816)[6]
Morula bicatenata (Reeve, 1846)[7]
Morula biconica (Blainville, 1832)[8]
Morula cernohorskyi (Houart & Tröndle, 1997)[9]
Morula chrysostoma (Deshayes, 1844)[10]
Morula coronata (H. Adams, 1869)[11]
Morula dichrous (Tapparone-Canefri, 1880)[12]
Morula echinata (Reeve, 1846)[13]
Morula euryspira (Houart, 1995)[14]
Morula fuscoimbricata (G.B. Sowerby III, 1915)[15]
Morula iostoma (Reeve, 1846)
Morula japonica (G.B. Sowerby III, 1903)[16]
Morula lepida (Houart, 1995)[17]
Morula nodicostata (Pease, 1868)[18]
Morula oparensis (Melvill, 1912)
Morula peasei (Houart, 2002)[19]
Morula porphyrostoma (Reeve, 1846)[20]
Morula praecipua (Rehder, 1980)[21]
Morula rodgersi (Houart, 2000)[22]
Morula spinosa (H. Adams & A. Adams, 1853)[23]
Morula striata (Pease, 1868)[24]
Morula uva (Röding, 1798)[25]
Morula variabilis (Pease, 1868)[26]
Morula whiteheadae (Houart, 2004)[27]
Morula zebrina (Houart, 2004)[28]

Species brought into synonymy

Morula (Azumamorula) mutica (Lamarck, 1816):[29] synonym of Azumamorula mutica (Lamarck, 1816)
Morula anaxares (Kiener, 1835):[30] synonym of Muricodrupa anaxares (Kiener, 1836)
Morula cancellata Quoy & Gaimard:[31] synonym of Muricodrupa fenestrata (Blainville, 1832)
Morula concatenata (Lamarck, 1822):[32] synonym of Drupella rugosa (Born, 1778)
Morula consanguinea (Smith, 1890):[33] synonym of Claremontiella consanguinea (E. A. Smith, 1890)
Morula elata Blainville:[34] synonym of Drupella cornus (Röding, 1798)
Morula fenestrata Blainville, 1832:[35] synonym of Muricodrupa fenestrata (Blainville, 1832)
Morula funiculata (Reeve, 1846):[36] synonym of Oppomorus funiculatus (Reeve, 1846)
Morula granulata (Duclos, 1832):[37] synonym of Tenguella granulata (Duclos, 1832)
Morula margariticola (Broderip):[38] synonym of Ergalatax margariticola (Broderip, in Broderip & Sowerby, 1833)
Morula marginalba (Blainville, 1832):[39] synonym of Tenguella marginalba (Blainville, 1832)
Morula marginatra Blainville, 1832:[40] synonym of Semiricinula marginatra (Blainville, 1832)
Morula musiva (Kiener, 1835):[41] synonym of Tenguella musiva (Kiener, 1835)
Morula nodulifera: synonym of Oppomorus noduliferus (Menke, 1829)
Morula nodulosa (C.B. Adams, 1845):[42] synonym of Claremontiella nodulosa (C. B. Adams, 1845)
Morula ocellata:[43] synonym of Ergalatax contracta (Reeve, 1846)
Morula oparense (Melvill, 1912):[44] synonym of Morula oparensis (Melvill, 1912)
Morula palmeri Powell, 1967: synonym of Pascula palmeri (Powell, 1967) (original combination)
Morula parva (Reeve, 1846):[45] synonym of Lauta parva (Reeve, 1846)
Morula purpureocincta (Preston, 1909):[46] synonym of Oppomorus purpureocinctus (Preston, 1909)
Morula rosea (Reeve, 1846):[47] synonym of Muricopsis (Risomurex) roseus (Reeve, 1846)
Morula rumphiusi Houart, 1996:[48] synonym of Murichorda rumphiusi (Houart, 1996)
Morula squamiliratum Smith:[49] synonym of Orania serotina (A. Adams, 1853)
Morula squamosa Pease:[50] synonym of Semiricinula squamosa (Pease, 1868)
Morula tuberculata Blainville:[51] synonym of Morula (Morula) granulata (Duclos, 1832)

Taxonomy

In 2002, Roland Houart described two new species and revised eight others, for a total 23 species of Morula sensu stricto.[52] He had previously (1994–1995) described Habromorula as a separate genus, containing 14 species that were earlier classified with Spinidrupa sensu lato.[53] In 2004, he placed Azumamorula, Oppomorus, and Habromorula into Morula as subgenera, and moved the whole group from subfamily Rapaninae to Ergalataxinae.[54]

Molecular genetics

In 2012–2013, Claremont et al. published a study of sequences for four genes across numerous species of Morula and related molluscs.[55] They found a clade of 11 species, and proposed a restricted definition of Morula to include: M. aspera, M. benedicta, M. (Habromorula) biconica, M. chrysostoma, M. (H.) coronata, M. (H.) japonica, M. nodicostata, M. (H.) spinosa, M. (H.) striata, M. uva (type species), and M. zebrina.

Habromorula did not form a clade in this analysis, however the authors noted that "monophyly of this morphologically distinctive group is not strongly contradicted, so that further sampling and analysis could yet support Habromorula as a monophyletic subgenus of Morula".

Subgenus Morula (Oppomorus) formed a separate clade; the authors proposed returning Oppomorus to genus level, containing O. nodulifera, O. funiculata and O. purpureocincta.

M. ceylonica, M. granulata, M. marginalba, M. musiva, and an undescribed species formed a distinct clade related to Muricodrupa. The authors proposed resurrecting the genus name Tenguella, for which T. granulata is the type species. They note that these species "have not previously been recognized as a morphologically distinctive group".

"Morula" anaxares was weakly related to Muricodrupa fenestrata, and "Morula" nodulosa was sister to the Tenguella–Muricodrupa group. "Morula" parva and "Morula" echinata placed in a third major clade, separate from the two that contain Tenguella and the main Morula group, respectively.

In response to the gene-sequence phylogeny, Houart, Zuccon and Puillandre formally described new genera in 2019: Claremontiella, Murichorda, and Lauta.[56] C. nodulosa, C. consanguinea, Mur. rumphiusi, and L. parva were previously in Morula. They also moved Morula anaxares to Muricodrupa anaxares, and reassigned M. palmeri to Pascula, the latter on basis of morphology rather than gene sequence.

References

Morula Schumacher, 1817. Retrieved through: World Register of Marine Species on 25 April 2010.
Morula aglaos (Houart, 1995). Retrieved through: World Register of Marine Species on 25 April 2010.
Morula albanigra Houart, 2002. Retrieved through: World Register of Marine Species on 25 April 2010.
Morula ambrosia (Houart, 1995). Retrieved through: World Register of Marine Species on 25 April 2010.
Morula angulata (G.B. Sowerby III, 1894). Retrieved through: World Register of Marine Species on 25 April 2010.
Morula aspera (Lamarck, 1816). Retrieved through: World Register of Marine Species on 25 April 2010.
Morula bicatenata (Reeve, 1846). Retrieved through: World Register of Marine Species on 25 April 2010.
Morula biconica (Blainville, 1832). Retrieved through: World Register of Marine Species on 25 April 2010.
Morula cernohorskyi Houart & Tröndle, 1997. Retrieved through: World Register of Marine Species on 25 April 2010.
Morula chrysostoma (Deshayes, 1844). Retrieved through: World Register of Marine Species on 20 November 2020.
Morula coronata (H. Adams, 1869). Retrieved through: World Register of Marine Species on 25 April 2010.
Morula dichrous (Tapparone-Canefri, 1880). Retrieved through: World Register of Marine Species on 25 April 2010.
Morula echinata (Reeve, 1846). Retrieved through: World Register of Marine Species on 25 April 2010.
Morula euryspira Houart, 1995. Retrieved through: World Register of Marine Species on 25 April 2010.
Morula fuscoimbricata (Sowerby, 1915). Retrieved through: World Register of Marine Species on 25 April 2010.
Morula japonica (Sowerby, 1903). Retrieved through: World Register of Marine Species on 25 April 2010.
Morula lepida (Houart, 1995). Retrieved through: World Register of Marine Species on 25 April 2010.
Morula nodicostata (Pease, 1868). Retrieved through: World Register of Marine Species on 25 April 2010.
Morula peasei Houart, 2002. Retrieved through: World Register of Marine Species on 25 April 2010.
Morula porphyrostoma (Reeve, 1846). Retrieved through: World Register of Marine Species on 25 April 2010.
Morula praecipua Rehder, 1980. Retrieved through: World Register of Marine Species on 25 April 2010.
Morula rodgersi Houart, 2000. Retrieved through: World Register of Marine Species on 25 April 2010.
Morula spinosa (H. Adams & A. Adams, 1853). Retrieved through: World Register of Marine Species on 25 April 2010.
Morula striata (Pease, 1868). Retrieved through: World Register of Marine Species on 25 April 2010.
Morula uva (Röding, 1798). Retrieved through: World Register of Marine Species on 25 April 2010.
Morula variabilis (Pease, 1868). Retrieved through: World Register of Marine Species on 25 April 2010.
Morula whiteheadae Houart, 2004. Retrieved through: World Register of Marine Species on 25 April 2010.
Morula zebrina Houart, 2004. Retrieved through: World Register of Marine Species on 25 April 2010.
Morula mutica (Lamarck, 1816). Retrieved through: World Register of Marine Species on 25 April 2010.
Morula anaxares (Kiener, 1835). Retrieved through: World Register of Marine Species on 25 April 2010.
Morula cancellata Quoy & Gaimard. Retrieved through: World Register of Marine Species on 25 April 2010.
Morula concatenata (Lamarck, 1822). Retrieved through: World Register of Marine Species on 25 April 2010.
Morula consanguinea (Smith, 1890). Retrieved through: World Register of Marine Species on 25 April 2010.
Morula elata Blainville. Retrieved through: World Register of Marine Species on 25 April 2010.
Morula fenestrata Blainville. Retrieved through: World Register of Marine Species on 25 April 2010.
Morula funiculata (Reeve, 1846). Retrieved through: World Register of Marine Species on 25 April 2010.
Morula granulata (Duclos, 1832). Retrieved through: World Register of Marine Species on 25 April 2010.
Morula margariticola (Broderip). Retrieved through: World Register of Marine Species on 25 April 2010.
Morula marginalba (Blainville, 1832). Retrieved through: World Register of Marine Species on 25 April 2010.
Morula marginatra Blainville. Retrieved through: World Register of Marine Species on 25 April 2010.
Morula musiva (Kiener, 1835). Retrieved through: World Register of Marine Species on 25 April 2010.
Morula nodulosa (C.B. Adams, 1845). Retrieved through: World Register of Marine Species on 25 April 2010.
Morula ocellata. Retrieved through: World Register of Marine Species on 25 April 2010.
Morula oparense (Melvill, 1912). Retrieved through: World Register of Marine Species on 25 April 2010.
Morula parva (Reeve, 1846). Retrieved through: World Register of Marine Species on 25 April 2010.
Morula purpureocincta (Preston, 1909). Retrieved through: World Register of Marine Species on 25 April 2010.
Morula rosea (Reeve, 1846). Retrieved through: World Register of Marine Species on 25 April 2010.
Morula rumphiusi Houart, 1996. Retrieved through: World Register of Marine Species on 25 April 2010.
Morula sguamiliratum Smith. Retrieved through: World Register of Marine Species on 25 April 2010.
Morula squamosa Pease. Retrieved through: World Register of Marine Species on 25 April 2010.
Morula tuberculata Blainville. Retrieved through: World Register of Marine Species on 25 April 2010.
Houart, Roland (2002). "Comments on a group of small Morula s.s. species (Gastropoda: Muricidae: Rapaninae) from the Indo-West Pacific with the description of two new species". Novapex. 3 (4): 97–118.
Houart, Roland (1994). "Some notes on the genus Spinidrupa Habe and Kosuge, 1966 (Muricidae: Ergalataxinae), with the description of Habromorula gen. nov. (Muricidae: Rapaninae) and four new species from the Indo-West Pacific". Iberus. 12: 21–31. BHL: info digitised copy of printed article
Houart, Roland (2004-11-10). "Review of the Recent species of Morula (Oppomorus), M. (Azumamorula) and M. (Habromorula) (Gastropoda: Muricidae: Ergalataxinae)". Novapex. 5 (4): 91–130.
Claremont, Martine; Houart, Roland; Williams, Suzanne T.; Reid, David G. (2013-02-01). "A molecular phylogenetic framework for the Ergalataxinae (Neogastropoda: Muricidae)". Journal of Molluscan Studies. 79 (1): 19–29. doi:10.1093/mollus/eys028. ISSN 0260-1230.
Houart, Roland; Zuccon, Dario; Puillandre, Nicolas (2019-04-09). "Description of new genera and new species of Ergalataxinae (Gastropoda: Muricidae)". {{cite journal}}: Cite journal requires |journal= (help)

External links

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[WoRMS-1] Morula Schumacher, 1817. Retrieved through: World Register of Marine Species on 25 April 2010.

[2] Morula aglaos (Houart, 1995). Retrieved through: World Register of Marine Species on 25 April 2010.

[3] Morula albanigra Houart, 2002. Retrieved through: World Register of Marine Species on 25 April 2010.

[4] Morula ambrosia (Houart, 1995). Retrieved through: World Register of Marine Species on 25 April 2010.

[5] Morula angulata (G.B. Sowerby III, 1894). Retrieved through: World Register of Marine Species on 25 April 2010.

[6] Morula aspera (Lamarck, 1816). Retrieved through: World Register of Marine Species on 25 April 2010.

[7] Morula bicatenata (Reeve, 1846). Retrieved through: World Register of Marine Species on 25 April 2010.

[8] Morula biconica (Blainville, 1832). Retrieved through: World Register of Marine Species on 25 April 2010.

[9] Morula cernohorskyi Houart & Tröndle, 1997. Retrieved through: World Register of Marine Species on 25 April 2010.

[10] Morula chrysostoma (Deshayes, 1844). Retrieved through: World Register of Marine Species on 20 November 2020.

[11] Morula coronata (H. Adams, 1869). Retrieved through: World Register of Marine Species on 25 April 2010.

[12] Morula dichrous (Tapparone-Canefri, 1880). Retrieved through: World Register of Marine Species on 25 April 2010.

[13] Morula echinata (Reeve, 1846). Retrieved through: World Register of Marine Species on 25 April 2010.

[14] Morula euryspira Houart, 1995. Retrieved through: World Register of Marine Species on 25 April 2010.

[15] Morula fuscoimbricata (Sowerby, 1915). Retrieved through: World Register of Marine Species on 25 April 2010.

[16] Morula japonica (Sowerby, 1903). Retrieved through: World Register of Marine Species on 25 April 2010.

[17] Morula lepida (Houart, 1995). Retrieved through: World Register of Marine Species on 25 April 2010.

[18] Morula nodicostata (Pease, 1868). Retrieved through: World Register of Marine Species on 25 April 2010.

[19] Morula peasei Houart, 2002. Retrieved through: World Register of Marine Species on 25 April 2010.

[20] Morula porphyrostoma (Reeve, 1846). Retrieved through: World Register of Marine Species on 25 April 2010.

[21] Morula praecipua Rehder, 1980. Retrieved through: World Register of Marine Species on 25 April 2010.

[22] Morula rodgersi Houart, 2000. Retrieved through: World Register of Marine Species on 25 April 2010.

[23] Morula spinosa (H. Adams & A. Adams, 1853). Retrieved through: World Register of Marine Species on 25 April 2010.

[24] Morula striata (Pease, 1868). Retrieved through: World Register of Marine Species on 25 April 2010.

[25] Morula uva (Röding, 1798). Retrieved through: World Register of Marine Species on 25 April 2010.

[26] Morula variabilis (Pease, 1868). Retrieved through: World Register of Marine Species on 25 April 2010.

[27] Morula whiteheadae Houart, 2004. Retrieved through: World Register of Marine Species on 25 April 2010.

[28] Morula zebrina Houart, 2004. Retrieved through: World Register of Marine Species on 25 April 2010.

[29] Morula mutica (Lamarck, 1816). Retrieved through: World Register of Marine Species on 25 April 2010.

[30] Morula anaxares (Kiener, 1835). Retrieved through: World Register of Marine Species on 25 April 2010.

[31] Morula cancellata Quoy & Gaimard. Retrieved through: World Register of Marine Species on 25 April 2010.

[32] Morula concatenata (Lamarck, 1822). Retrieved through: World Register of Marine Species on 25 April 2010.

[33] Morula consanguinea (Smith, 1890). Retrieved through: World Register of Marine Species on 25 April 2010.

[34] Morula elata Blainville. Retrieved through: World Register of Marine Species on 25 April 2010.

[35] Morula fenestrata Blainville. Retrieved through: World Register of Marine Species on 25 April 2010.

[36] Morula funiculata (Reeve, 1846). Retrieved through: World Register of Marine Species on 25 April 2010.

[37] Morula granulata (Duclos, 1832). Retrieved through: World Register of Marine Species on 25 April 2010.

[38] Morula margariticola (Broderip). Retrieved through: World Register of Marine Species on 25 April 2010.

[39] Morula marginalba (Blainville, 1832). Retrieved through: World Register of Marine Species on 25 April 2010.

[40] Morula marginatra Blainville. Retrieved through: World Register of Marine Species on 25 April 2010.

[41] Morula musiva (Kiener, 1835). Retrieved through: World Register of Marine Species on 25 April 2010.

[42] Morula nodulosa (C.B. Adams, 1845). Retrieved through: World Register of Marine Species on 25 April 2010.

[43] Morula ocellata. Retrieved through: World Register of Marine Species on 25 April 2010.

[44] Morula oparense (Melvill, 1912). Retrieved through: World Register of Marine Species on 25 April 2010.

[45] Morula parva (Reeve, 1846). Retrieved through: World Register of Marine Species on 25 April 2010.

[46] Morula purpureocincta (Preston, 1909). Retrieved through: World Register of Marine Species on 25 April 2010.

[47] Morula rosea (Reeve, 1846). Retrieved through: World Register of Marine Species on 25 April 2010.

[48] Morula rumphiusi Houart, 1996. Retrieved through: World Register of Marine Species on 25 April 2010.

[49] Morula sguamiliratum Smith. Retrieved through: World Register of Marine Species on 25 April 2010.

[50] Morula squamosa Pease. Retrieved through: World Register of Marine Species on 25 April 2010.

[51] Morula tuberculata Blainville. Retrieved through: World Register of Marine Species on 25 April 2010.

[52] Houart, Roland (2002). "Comments on a group of small Morula s.s. species (Gastropoda: Muricidae: Rapaninae) from the Indo-West Pacific with the description of two new species". Novapex. 3 (4): 97–118.

[53] Houart, Roland (1994). "Some notes on the genus Spinidrupa Habe and Kosuge, 1966 (Muricidae: Ergalataxinae), with the description of Habromorula gen. nov. (Muricidae: Rapaninae) and four new species from the Indo-West Pacific". Iberus. 12: 21–31. BHL: info digitised copy of printed article

[Houart2004-54] Houart, Roland (2004-11-10). "Review of the Recent species of Morula (Oppomorus), M. (Azumamorula) and M. (Habromorula) (Gastropoda: Muricidae: Ergalataxinae)". Novapex. 5 (4): 91–130.

[55] Claremont, Martine; Houart, Roland; Williams, Suzanne T.; Reid, David G. (2013-02-01). "A molecular phylogenetic framework for the Ergalataxinae (Neogastropoda: Muricidae)". Journal of Molluscan Studies. 79 (1): 19–29. doi:10.1093/mollus/eys028. ISSN 0260-1230.

[56] Houart, Roland; Zuccon, Dario; Puillandre, Nicolas (2019-04-09). "Description of new genera and new species of Ergalataxinae (Gastropoda: Muricidae)". {{cite journal}}: Cite journal requires |journal= (help)