Figure 1.
Rhizoids found in Candidae species.
Rhizoids found in Candidae species (arrows). A, NHMUK 1926.9.6.84, Licornia jolloisii (Audouin, 1826); interconnective rhizoids (white arrow). B–D, Holdfast rhizoids (white arrows). B, NHMUK 1963.3.6.7a, Cradoscrupocellaria ellisi (Vieira & Spencer Jones, 2012); smooth rhizoids. C, NHMUK 2010.12.6.1, Cradoscrupocellaria arisaigensis Vieira, Spencer Jones & Winston, 2013; rhizoids with retroussé hooks. D, NHMUK 1899.6.1.340, Licornia cyclostoma (Busk, 1852); ringed rhizoids. E, USNM 8426, Paralicornia sinuosa (Canu & Bassler, 1927) n. comb.; colony without rhizoid on abfrontal surface. F, NHMUK 1887.12.9.83, Notoplites clausus (Busk, 1884); colony with adjacent rhizoids on abfrontal surface (black arrow).
Figure 2.
Notation for the order of zooids at branch bifurcations.
Abfrontal surface of colony showing the uniform notation for the order of zooids at branch bifurcations proposed by Harmer [25] and the position of the joints (modified from Harmer [25]). A, Joints passing across zooids FD and GC (type 15). B, Joints passing across zooids FJ and GK (type 8).
Figure 3.
Branch and zooid morphologies.
A, NHMUK 1890.1.30.25-27, Canda retiformis (Pourtalès, 1867); adjacent zooids sharply inclined in relation to the axis, characteristic of Canda. B–C, Position of the joints in zooids C and D at bifurcation; note the development of cryptocyst (white arrows) and two distinct opercula (rectangle). B, NHMUK 1936.12.30.146, Licornia jolloisii (Audouin, 1826); the joints (j) pass across the opesiae in zooids C and D; note the membranous operculum with its distinctly chitinous distal rim. C, NHMUK 1899.7.1.780, Scrupocaberea maderensis (Busk, 1860) n. comb.; the joints (j) are passing across the gymnocysts in C and D zooids; note the entirely chitinous operculum placed in an obliquely truncate distal area. D–F, Distal edge of autozooid (black arrow). D, NHMUK 1987.1.18.41, Paralicornia limatula (Hayward, 1988) n. comb.; smooth distal edge. E, NHMUK 1996.4.26.5, Scrupocaberea sp.; corrugated distal edge in ovicelled zooids. F, NHMUK 1934.10.8.1, Scrupocellaria minuta (Kirkpatrick, 1888); corrugated distal edge in non-ovicelled zooids.
Figure 4.
Oral spines and scutum. A, NHMUK 1911.10.1.367, Scrupocellaria inermis Norman, 1867; zooid without distal spines and scutum. B, NHMUK 1882.5.24.8-12, Cradoscrupocellaria gautieri Vieira, Spencer Jones & Winston, 2013; zooid with unbranched oral spines. C, MZUSP 266, Aspiscellaria sp.; zooid with proximal-most outer spine branched (cervicorn) and additional unbranched spines; note the single ovoid plate of scutum. D, MZUSP 532, Licornia drachi (Marcus, 1955) n. comb.; zooid with proximal-outer, outer and inner spines bifurcated, and distal unbranched spines. E–L, Distinct morphologies of scutum at the inner edge of opesia, arising at the median region of opesia—Character 16: E, state 0, NHMUK 1936.12.30.146, Licornia jolloisii (Audouin, 1826); F, state 1, NHMUK 2010.10.6.1, Cradoscrupocellaria tenuirostris (Osburn., 1950); G, state 2, NHMUK 1911.10.1.353, Cradoscrupocellaria ellisi (Vieira & Spencer Jones, 2012); H, state 3, NHMUK 2010.12.6.22, Cradoscrupocellaria odonoghuei Vieira, Spencer Jones & Winston, 2013—or from its distal third—Character 17: I–J, state 0 (I, NHMUK 1968.1.18.110, Pomocellaria californica (Trask, 1857); J, NHMUK 1911.10.1.373, Scrupocellaria delilii (Audouin, 1826)); K, state 2, NHMUK 1928.9.13.98, Scrupocaberea dongolensis (Waters 1908); L, state 3, NHMUK 1934.10.8.1, Scrupocellaria minuta (Kirkpatrick, 1888).
Figure 5.
Frontal and lateral avicularia.
Frontal and lateral avicularia. A, NHMUK 1888.4.16.200, Aspiscellaria frondis (Kirkpatrick, 1890) n. comb.; zooids without lateral avicularium. B–H, Zooids with sessile lateral avicularium (white and black arrows). B, NHMUK 2010.12.6.22, Cradoscrupocellaria odonoghuei Vieira, Spencer Jones & Winston, 2013; zooid with avicularium positioned at outer distal corner and directed obliquely distolaterally. C, NHMUK 1899.7.1.804, Scrupocaberea dongolensis (Waters 1908) n. comb.; zooid with avicularium positioned lateral to the opercular area and directed laterally. D–F, Rostrum of lateral avicularium. D, NHMUK 1936.12.30.146, Licornia gaspari (Thornely 1907); serrated lateral edge, straight at its tips. E, NHMUK 1968.1.18.110, Pomocellaria californica (Trask, 1857) n. comb.; smooth lateral edge, with curved tips. F, NHMUK 1899.7.1.804, Scrupocaberea dongolensis (Waters 1908) n. comb.; serrated lateral edge, strongly hooked. G–I, Dimorphic lateral and frontal avicularia (black arrows). G, NHMUK 1968.1.18.110, Pomocellaria californica (Trask, 1857) n. comb.; dimorphic lateral avicularium with triangular mandible. H, NHMUK 1998.9.29.6, Paralicornia sp.; dimorphic lateral avicularium with trifoliate mandible. I, NHMUK 1968.1.18.104, Licornia diegensis (Robertson, 1905) n. comb.; dimorphic frontal avicularium.
Figure 6.
Sessile ooecia found in Candidae.
Sessile ooecia found in Candidae. A, NHMUK 1966.1.10.9, Scrupocellaria scruposa (Linnaeus, 1758); ooecium with a small drop-shaped pore with a distal avicularium (white arrow). B, NHMUK 1968.1.18.111, Pomocellaria varians (Hincks, 1882) n. comb.; ooecium with single, large fenestra, without avicularium. C, NHMUK 1899.7.1.736, Cradoscrupocellaria bertholletii (Audouin, 1826); ooecium with porous ectooecium.
Figure 7.
Heterozooids on abfrontal surface of the colony.
Heterozooids on abfrontal surface of the colony. A, NHMUK 1911.10.1.385, Tricellaria congesta (Norman, 1903); colony without abfrontal heterozooids. B, NHMUK 1911.10.1.376, Aquiloniella scabra (van Beneden, 1848) n. comb.; abfrontal avicularia. C, NHMUK 1929.4.26.17, Cradoscrupocellaria galapagensis Vieira, Spencer Jones & Winston, 2013; abfrontal vibracula, with transverse setal groove. D, lateral rhizoidal chamber associated with outer zooids at the bifurcation in Tricellaria sp.; note the absence of abfrontal avicularia. E–I, Abfrontal vibracula with oblique setal groove (axial vibracula shown by black arrow and other vibracula shown by white arrow). E, NHMUK 1987.1.18.41, Paralicornia limatula (Hayward, 1988) n. comb.; vibracula with straight setal groove occupying less than half the length of the vibracular chamber; the setal groove of the axial vibraculum is laterally placed. F, NHMUK 1936.12.30.146, Licornia gaspari (Thornely 1907); vibracula with straight setal groove occupying two thirds of the length of the vibracular chamber; the palate is complete and with a small foramen. G, NHMUK 1928.9.13.105, Licornia curvata (Harmer, 1926) n. comb.; vibracula with straight setal groove occupying the entire length of the vibracular chamber; note the presence of single axial vibraculum. H, NHMUK 1911.10.1.373, Scrupocellaria delilii (Audouin, 1826); vibracula with curved setal groove occupying two thirds of the total length of the vibracular chamber; note the presence of two axial vibracula (black arrows). I, MOM 420193, Scrupocellaria incurvata Waters 1897; vibracula with curved setal groove occupying the entire length of the vibracular chamber; note the presence of two axial vibracula (black arrows).
Figure 8.
Semi-strict consensus tree based on Notoplites as outgroup. New combinations are marked with an asterisk (*). Values below branches refer to Bremer supports (>1).
Figure 9.
Scanning electron micrographs of Scrupocellaria scruposa (Linnaeus, 1758).
Scrupocellaria scruposa (Linnaeus, 1758). A,C. NHMUK 1966.1.10.9, English Channel, United Kingdom; B. NHMUK 1847.9.15.28, British coast. A, Frontal surface of the bifurcation. B, Close-up of the frontal surface of the ovicelled zooid. C, Abfrontal surface of the colony.
Figure 10.
Species assigned to Scrupocellaria van Beneden, 1845 s. str.
Species assigned to Scrupocellaria van Beneden, 1845 s. str. A–B, Scrupocellaria aegeensis Harmelin, 1969, NHMUK 2010.12.7.3, Aegean Sea. C–D, Scrupocellaria delilii (Audouin, 1826), NHMUK 1911.10.1.373, Adriatic. E–F, Scrupocellaria incurvata Waters, 1897, NHMUK 1899.7.1.797, Crete, Mediterranean. G–H, Scrupocellaria inermis Norman 1867, NHMUK 1911.10.1.367, syntype, British coast. I–J, Scrupocellaria intermedia Norman, 1896, NHMUK 1911.10.1.369, syntype, Norway. K–L, Scrupocellaria minuta (Kirkpatrick, 1888), NHMUK 1934.10.8.1, Mauritius. M–N, Scrupocellaria scrupea Busk, 1851, NHMUK 2010.12.8.5, Guernsey.
Figure 11.
Species assigned to Aquiloniella n. gen.
Species assigned to Aquiloniella n. gen. A–B, Aquiloniella americana (Packard, 1863) n. comb., MCZ 134, syntype, ?Labrador. C–D, Aquiloniella orientalis (Kluge, 1955) n. comb., Svalbard (P. Kuklinski coll.). E–F, Aquiloniella paenulata (Norman, 1903) n. comb., MCZ 523, Gulf of Maine. G–H, Aquiloniella scabra (van Beneden, 1848) n. comb., NHMUK 1911.10.1.376, United Kingdom.
Figure 12.
Species assigned to Aspiscellaria n. gen.
Species assigned to Aspiscellaria n. gen. A–B, Aspiscellaria bellula (Osburn 1947) n. comb., AMNH 1518.1, Jamaica. C–D, Aspiscellaria carmabi (Fransen, 1986) n. comb.; C, VMNH 13699, Belize; D, NHMUK 2012.7.1.10, paratype, Curaçao. E–F, Aspiscellaria cornigera (Pourtalès, 1867) n. comb., NHMUK 1911.10.1.368, Barbados. G–H, Aspiscellaria frondis (Kirkpatrick 1890) n. comb., LM Vieira coll., Brazil. I–J, Aspiscellaria hildae (Fransen, 1986) n. comb., NHMUK 2012.7.1.9, paratype, Curaçao. K–L, Aspiscellaria piscaderaensis (Fransen, 1986) n. comb., NHMUK 2012.7.1.8, paratype, Curaçao.
Figure 13.
Species assigned to Paralicornia n. gen.
Species assigned to Paralicornia n. gen. A–B, Paralicornia limatula (Hayward, 1988) n. comb., NHMUK 1987.1.18.41, Mauritius. C–D, Paralicornia obtecta (Haswell, 1880) n. comb., NHMUK 1897.5.1.209, Australia. E–F, Paralicornia sinuosa (Canu & Bassler, 1927) n. comb., USNM 8426, Hawaii. G–L, Undescribed species of Paralicornia: (G–H) NHMUK 1996.4.26.8, Mauritius; (I–J) NHMUK 1936.12.30.177, Mauritius; (K–L) NHMUK 2000.4.11.1546, Australia.
Figure 14.
Species assigned to Pomocellaria n. gen.
Species assigned to Pomocellaria n. gen. A–B, Pomocellaria californica (Trask, 1857) n. comb., NHMUK 1968.1.18.110, California. C–D, Pomocellaria inarmata (O'Donoghue & O'Donoghue, 1926) n. comb., NHMUK 1964.4.2.10, holotype, Pacific coast of United States. E–F, Pomocellaria talonis (Osburn, 1950) n. comb., VMNH 13690, Pacific coast of United States. G–H, Pomocellaria varians (Hincks, 1882) n. comb., NHMUK 1968.1.18.111, California.
Figure 15.
Species assigned to Scrupocaberea n. gen.
Species assigned to Scrupocaberea n. gen. A–C, Scrupocaberea maderensis (Busk, 1860) n. comb. A, NHMUK 1899.7.1.780, syntype, Madeira; B–C, specimen from Azores. D–E, Scrupocaberea dongolensis (Waters, 1909) n. comb., NHMUK 1928.9.13.98, syntype, Sri Lanka. F–G, Scrupocaberea ornithorhynchus (Wyville Thomson, 1858) n. comb., NHMUK 1899.7.1.783, syntype, Australia. H–L, Undescribed species of Scrupocaberea: (H–I) NHMUK 1887.12.9.103, Cape Verde, (J–K) NHMUK 1961.11.2.42, Gulf of Mexico; (L) NHMUK 1928.3.6.169, Malay Archipelago.