Nitscheina spiculata Canu, 1928: 262 Nitscheina seurati Canu, 1928: 263 Nitscheina fluviatilis Canu, 1928: 263 Conopeum seurati : Bobin and Prenant 1962: 381 Prenant and Bobin 1966: 127 Jebram 1968: 90 Jebram, 1969a: 225 Jebram, 1969b: 341 Ryland and Hayward 1977: 62 Jebram and Rummert 1978: 502 Poluzzi 1980: 101 Occhipinti Ambrogi and d’Hondt 1981: 192 Pisano and Matricardi 1983: 311 Poluzzi and Forti 1983: 54 Occhipinti Ambrogi 1985: 223 Poluzzi and Sabelli 1985: 269 Zabala 1986: 244 Poluzzi and Agnoletto 1988: 7 Zabala and Maluquer 1988: 77 Read and Gordon 1991: 270 Hayward and Ryland 1998: 124 Rosso 2003: 243 De Blauwe 2009: 172 Rosso et al. 2010: 596 Ayari and Taylor 2014: 18 Chimenz Gusso et al. 2014: 89 Rosso and Di Martino 2016: 573 Rouse et al. 2018: 143 The bryozoan genus Conopeum (Electridae) in New Zealand, with description of a new species and discussion of the morphological and genetic characters of Conopeum seurati (Canu, 1928) Gordon, Dennis P. Sutherland, Judy E. Perez, Brenda A. Waeschenbach, Andrea Taylor, Paul D. Di Martino, Emanuela Journal of Natural History 2020 J. Nat. Hist. 2020-09-30 54 15 - 16 947 970 XRGR (Canu, 1928) Canu 1928 [369,778,574,601] Gymnolaemata Electridae Conopeum Animalia Cheilostomatida 9 955 Bryozoa species seurati      Nitscheina spiculata Canu, 1928: 262, pl. 30, figs 1, 2.     Nitscheina seurati Canu, 1928: 263, pl. 30, figs 3–6.     Nitscheina fluviatilis Canu, 1928: 263, pl. 30, fig. 7.     Conopeum seurati: Bobin and Prenant 1962: 381, figs 2, 3;  Prenant and Bobin 1966: 127, figs 33, 34 (cum syn.);  Jebram 1968: 90, fig. 3,  Jebram, 1969a: 225;  Jebram, 1969b: 341;  Ryland and Hayward 1977: 62, fig. 21;  Jebram and Rummert 1978: 502;  Poluzzi 1980: 101; Occhipinti Ambrogi 1981: 80, fig. 26;  Occhipinti Ambrogi and d’Hondt 1981: 192;  Pisano and Matricardi 1983: 311;  Poluzzi and Forti 1983: 54, pl. 1, fig 2, pl. 2, figs 1, 2;  Occhipinti Ambrogi 1985: 223;  Poluzzi and Sabelli 1985: 269, figs 5, 8, 9;  Zabala 1986: 244, fig. 55;  Poluzzi and Agnoletto 1988: 7, fig. 4A;  Zabala and Maluquer 1988: 77, fig. 66;  Read and Gordon 1991: 270, fig. 1A;  Hayward and Ryland 1998: 124, fig. 24C, D, 25;  Rosso 2003: 243;  De Blauwe 2009: 172, figs 150–152;  Rosso et al. 2010: 596;  Ayari and Taylor 2014: 18;  Chimenz Gusso et al. 2014: 89, fig. 28a–d;  Rosso and Di Martino 2016: 573;  Rouse et al. 2018: 143.   Material examined   Mediterranean: NHMUK 2020.1. 3.1, 36.7505° N, 10.5317° E, 0 m, topotypic specimen from Wadi Bezirk, Tunisia, collected by Rakia Ayari; NHMUK 2020.1. 3.2, Wadi Tinja, Tunisia, collected by Rakia Ayari; NMNH A66832, labelled  Nitscheina spiculata, Wadi Abid, Tunisia, collected by Prof . Léon Gaston Seurat( Universityof Algiers); NIWA 134508, StnZ18077, 42.9495° N, 10.5439° E, 0 m, Piombino, Tuscany, Italy, coll.   26 July 2016; NIWA 134512, StnZ18078, 44.8435° N, 12.2898° E, 0 m, on  Xenostrobus securis, Goro, Po Riverdelta, Italy, coll.   23 September 2016; NIWA 134513, StnZ18079, 44.8190° N, 12.3468° E, 0 m, on  Xenostrobus securis, Gorino, Po Riverdelta, Italy, coll. 23 September 2016by Agnese Marchini and Jasmine Ferrario.   New Zealand: NIWA 98801, StnZ18071, 35.3124° S, 174.1210° E, 0 m, Opua, Bay of Islands, New Zealand, coll. 31 July 1988by D.P.  Gordonand S.F.  Mawatari; NIWA 98802, StnZ18614, ~ 43.6108° S, 172.7048 E, Lyttelton, coll. 17 August 1988by D.P.  Gordonand S.F.  Mawatarifrom vessel being cleaned; NIWA 98946, StnZ18081, ~ 40.5569° S, 172.6315° E, Whanganui Inlet, 0 m, on  Xenostrobus securis, coll. January 1963by O.J.  Marston.  MITS 1863, 2WRE105, 35.7321° S, 174.3258° E, no depth data, Whangarei,  3 November 2005; MITS 1925, 2WRE116, 35.7255° S, 174.3271° E, no depth data, Whangarei,  3 November 2005; MITS 66480, WRE048, 35.7233° S, 174.3255° E,  2 m, Whangarei,  14 November 2002; MITS 66498, WRE050, 35.7238° S, 174.3253° E,  1 m, Whangarei,  14 November 2002; MITS 66516, WRE053, 35.7233° S, 174.3255° E,  2 m, Whangarei,  14 November 2002; MITS 66550, WRE061, 35.7242° S, 174.3270° E,  2 m, Whangarei,  14 November 2002; MITS 66578, WRE069, 35.7254° S, 174.3285° E,  2 m, Whangarei,  14 November 2002; MITS 66588, WRE072, 35.7254° S, 174.3285° E,  3 m, Whangarei,  14 November 2002; MITS 71244, OPX20201, 35.3143° S, 174.1223° E,  6 m, Opua,  18 February 2015.   Description Colony encrusting, typically unilamellar but sometimes self-overgrowing and multiserial. Mature autozooids in zone of astogenetic repetition arranged quincuncially, often twice as long as wide and rounded-subrectangular to elongate-oval, but shape variable depending on position in colony; lateral walls of adjoining zooids closely abutted. Entire frontal area covered by membrane-like cuticularised frontal wall; proximal gymnocyst absent ( Figure 3(a)) to little-developed ( Figure 3(d,e)), 1–2 rare spine bases in distolateral corners of some zooids ( Figure 3(d)). No marginal spines. Basal wall wholly calcified, no window. Operculum at distal end of frontal wall, differentiated within it as a distally rounded flap a little longer than semicircular.   Figure 4.  Conopeum seurati( Canu, 1928): (a, c), NIWA 98946, Whanganui Inlet, South Island, New Zealand; (b), NIWA 134508, Piombino, Tuscany, Italy. (a), Autozooids showing the peripheral cryptocystal groove, and a small kenozooid. (b), Lateral view of the cryptocystal rim of an autozooid, showing part of the peripheral grove, granules, minute pores (in the groove and lateral interior wall, and multiporous septula). (c), Lateral view of an autozooid and its proximal neighbour with mural septula. Scalebars: (a, c), 200 μm; (b), 100 μm. Opesia entirely surrounded by narrow sloping cryptocyst ( Figure 3(b)) that is generally of more or less equal width throughout, its surface with conspicuous rounded or pointed granules in 1–4 discontinuous series depending on cryptocyst width; granular part of cryptocyst almost completely surrounded by very narrow furrow ( Figures 3(b). 4(a, b)) inside thin zooid rim; furrow constricted or absent mid-distally. In older (ephebic) zooids, granular cryptocyst thicker, with more pustules, and furrow more obvious. Kenozooids sporadic, small and adventitious ( Figures 3(a), 4(a)) or large ones interzooidal ( Figure 3(e)), typically inserted in corners between autozooids, small, round, elongate-oval, bean-shaped or spindle-shaped, opesia unevenly surrounded by granular cryptocyst and either no gymnocyst or a tiny exposure. Lateral interzooidal communications via three multiporous mural septula along each side ( Figure 4(c)); larger septula occur in lower half of transverse walls, more or less as a crescentic cluster of pores. Sparse, thin calcareous spinules on inner face of distal transverse wall above septula ( Figure 3(b)) or lateral walls either side of septula ( Figure 4(b)), often associated with tiny pores; these pores can also occur within the cryptocystal furrow ( Figure 4(b)) and in basal walls.  Measurements Separate measurements were made of specimens from Wadi Bezirk ( typelocality), the Po River Delta ( Italy) and Whanganui Inlet ( New Zealand): Wadi Bezirk: ZL432–722 (574) [ N= 28]; ZW 150–501 (331) [ N= 36]; OpL350–653 (520) [ N= 18]; OpW101–357 (266) [ N= 24]; PCrL3–62 (28) [ N= 18]; KL149–471 (289) [ N= 10]; KW84–182 (120) [ N= 10] Po Delta: ZL342–580 (480) [ N= 49]; ZW260–373 (313) [ N= 38]; OpL274–480 (400) [ N= 43]; OpW181–278 (227) [ N= 34]; PCrL3–62 (28) [ N= 18]; ZD112–184 (143) [ N= 4] Whanganui Inlet: ZL412–533 (478) [ N= 31]; ZW223–361 (295) [ N= 30]; OpL296–445 (382) [ N= 31]; OpW 167–268 (225) [ N= 32]; PCrL 20–65 (40) [ N= 11]; ZD133–156 (145) [ N= 4]   Remarks The topotypic specimen described above lacked an ancestrula, but a well-developed specimen from Wadi Tinja has a well-preserved ancestrular region. The ancestrular opesia is elongate-oval with a thin non-granular cryptocystal rim and there are no spines. The ancestrula is surrounded by five zooids. There is a mid-distal daughter zooid, which itself has budded one mid-distal and two distolateral zooids. The original daughter zooid and its two distolateral zooids are connected to two other zooids, directed proximally, that flank the ancestrula; they in turn have produced two proximal zooids that abut the proximolateral corners of the ancestrula. Bobin and Prenant (1962)show the isolated ancestrula (based on a specimen from Essex, England) as subcircular with a pair of distolateral spines, having a more-developed cryptocystal rim than in  C. antipodum n. sp. In comparing  C. seuratiand  C. antipodum, the most obvious distinction, especially evident in SEM micrographs, is the cryptocystal furrow in  C. seurati( Figure 3(b)). This furrow is consistent in all Mediterranean specimens that we have seen but completely absent in  C. antipodum. The furrow is mostly not apparent in previously published line drawings, exceptions being Jebram (1968, fig. 3) and Poluzzi (1980, fig. 1 inset; reproduced in Poluzzi and Sabelli 1985, fig. 1 inset). It is seen, but has not been especially highlighted, in published SEM micrographs (e.g. Poluzzi 1980, pl. 2, figs 5, 6; Poluzzi and Agnoletto 1988, fig. 4A). It is apparent in specimens from New Zealand( Figure 4(a,b)), confirming the presence of naturalised populations. Excluding material collected from a vessel hull in Lyttelton,  C. seuratihas been found on anthropogenic substrata at Opua in Northland, in the Town Basin of Whangarei Harbour and in Whanganui Inlet, South Island.  Conopeum antipodum n. sp.is by far the commoner species in New Zealand. The typesof the three species from Wadi Bezirk (  N. seurati, N. spiculata, N. fluviatilis) described by Canu (1928)seem to be no longer extant in the collection of Recent Bryozoa in the MNHM, Paris. Inasmuch as Bobin and Prenant (1962)did not mention having examined them when accepting Borg’s (1931)conclusion that they were all the same species,they may have been missing by that time. However,one of us (EDM) found a historic specimen labelled  Nitscheina spiculata( Figure 3(c)) from Wadi Abid, Tunisia, in the palaeontological collection. This was scanned by SEM for the present study ( Figure 3(d,e)).  Canu’s (1928)illustrations (light micrographs) are very poor, and inadequate to see much in the way of discriminating details. Borg (1931, p. 24) requested, and was sent by Canu, all three species of  Nitscheina(presumably all topotypic) and concluded that they constituted the same variable species. Borg noted that the specimen labelled  N. spiculatahad zooids in the same colony with and without lateral spines. It is unclear whether Borg (1931)was including the distolateral spines in his description of ‘here and there also rather long lateral spines’. He described  N. seuratias being ‘only sparsely provided with lateral spines’, whereas  N. fluviatiliswas ‘provided with lateral spines that are rather variable in length and sometimes in their upper part more or less curved, but many of the zoids [sic] lack spines’. Borg (1936)redescribed a specimen of  N. spiculatafrom Wadi Abid (probably the one examined for this study, collected in June 1928by Prof. Seurat). Again, he treated all three species as synonymous, calling them ‘  Membranipora( Electra) crustulenta(Pallas) var. spiculataCanu’. Bobin and Prenant (1962)showed conclusively that this entity (  spiculata/ seurati/ fluviatilis) was not conspecific or congeneric with  Electra crustulenta(now  Einhornia crustulenta( Pallas, 1766)). Bobin and Prenant (1962)were the first to use the combination  Conopeum seurati, applying it to all three species described by Canu (1928). They noted that  N. spiculatahad page priority in Canu’s paper (published in November 1928) but rejected use of this species name on the grounds that the combination  Nitscheina spiculatawas preoccupied – ”le même nom  N. spiculataavait été utilise déjà par Canu et Bassler (1923, p. 21) pour une éspèce fossile du Tertiaire américain”. In fact, Canu and Bassler (1923)named the species  Membranipora spiculata(which, judging from their photos, is not  Membraniporabut a species of Electridaeor perhaps Calloporidae). Borg (1931, 1936) was correct in retaining  spiculatafor the Tunisian- Mediterranean species but so many authors have followed Bobin and Prenant (1962)and Prenant and Bobin (1966)that the combination  Conopeum seuratiis entrenched in the literature and there seems little point in changing it. [Note that the synonymy given above is deliberately not exhaustive for such a well-known species; additional earlier literature is cited in Prenant and Bobin (1966).] The contemporary topotypic material from Wadi Bezirk examined by SEM shows no obvious frontal gymnocyst or spines ( Figure 3(a)), whereas a historic museum specimen from Wadi Abid labelled  Nitscheina spiculatahas in the same colony many zooids identical to those from Wadi Bezirk. There are also many zooids in which the cystid walls are raised frontalwards around all or part of the zooid, being developed more distolaterally and/or proximolaterally. A distolateral pair of spines is frequently present, being either very short, having the form of pointed tubercles, or much longer and tapering, with well-developed bases that remain when the spines are broken off. Neither the specimens from Wadi Bezirk nor Wadi Abid seen by us have lateral spines, but they were present on some of the original specimens described from this locality, as elsewhere in the Mediterranean and Western European coasts today. Following Borg (1931, 1936), Bobin and Prenant (1962)and Prenant and Bobin (1966), we accept the synonymy of the three species described by Canu (1928), a conclusion supported by gene sequencing of morphologically variable colonies from elsewhere (see below). Variations in Mediterranean  C. seuratihave been documented by different authors, the most comprehensive studies being those of Poluzzi and Forti (1983)and Poluzzi and Sabelli (1985), who described self-overgrowth and boundary interactions, with the formation of erect lobes, some bilamellar. Colonies could also grow around thin algal filaments, forming hollow tubes. Poluzzi and Sabelli (1985)termed some expressions of self-overgrowth as ‘celleporoid layers’ – in which regenerated zooids act as pseudoancestrulae, budding heaps of zooids in loose massive layers of randomly orientated polygonal zooids with smaller polypides. Additionally, both basal and celleporoid layers can include autozooids with closure plates – non-feeding zooids filled with what appeared to be nutritive material. It is noteworthy that  Conopeum seuratiin New Zealandhas been found associated with  Xenostrobus securis, as in Whanganui Inlet; this association was reported for the Italian species sent from the Po River delta for our genetic analysis. Equally, some  C. seuratiin the Town Basin, Whangarei Harbour, was found encrusting tubes of the naturalised-alien polychaete  Ficopomatus enigmaticus( Read and Gordon 1991), an association also seen in Tunisia( Canu 1928), France( Prenant and Bobin 1966), Italy( Pisano and Matricardi 1983) and Turkey( Gugel 1997). [160,504,1149,1174] NHMUK Mediterranean 9 955 NHMUK 2020.1 1 NHMUK Rakia Ayari Tunisia 36.7505 Wadi Bezirk 6 10.5317 9 955 NHMUK 2020.1 1 A66832 NMNH Rakia Ayari & Prof Tunisia Wadi Tinja 9 955 1 Z18077 NIWA Piombino Italy Stn 42.9495 Leon Gaston Seurat 6 10.5439 University 9 955 NIWA 134508 1 Toscana Z18078 2016-07-26 NIWA Italy 44.8435 Stn 6 12.2898 9 955 NIWA 134512 1 Z18079 2016-09-23 NIWA Italy 44.819 Stn 6 12.3468 9 955 NIWA 134513 1 Z18071 NIWA New Zealand Bay of Islands -35.3124 Stn 6 174.121 Opua 9 955 NIWA 98801 1 [677,832,1495,1520] New Zealand Gordon 9 955 1 Z18614 NIWA Lyttelton New Zealand -43.6108 Mawatari 6 172.7048 Stn 9 955 NIWA 98802 1 [975,1134,1530,1555] New Zealand Gordon 9 955 1 Z18081 NIWA New Zealand Whanganui Inlet -40.5569 Mawatari 6 172.6315 Stn 9 955 NIWA 98946 1 [935,1028,1599,1624] New Zealand Marston 9 955 1 OPX20201 2002-11-14 2015-02-18 2002-11-14 MITS New Zealand Whangarei 2 10 956 -35.3143 Whangarei 6 174.1223 Whangarei 9 955 MITS 1863, 2, MITS 1925, MITS 66480, MITS 66498, MITS 66516, MITS 66550, MITS 66578, MITS 66588, MITS 71244 1