taxonID	type	description	language	source
03BCE061A162FFD3B790FE5A5F69FA05.taxon	type_taxon	Type species. Cribrilina (Juxtacribrilina) flavomaris Yang, Seo, Min, Grischenko & Gordon, 2018, by original designation.	en	Dick, Matthew H., Grischenko, Andrei V., Gordon, Dennis P., Ostrovsky, Andrew N. (2021): The “ Cribrilina annulata " problem and new species of Juxtacribrilina (Bryozoa Cheilostomata: Cribrilinidae) from the North Pacific. Zootaxa 5016 (3): 333-364, DOI: 10.11646/zootaxa.5016.3.2
03BCE061A162FFD3B790FE5A5F69FA05.taxon	diagnosis	Diagnosis. Colony encrusting, multiserial, usually unilaminar except for scattered reduced or dwarf ovicellate zooids overlying basal zooidal layer (limited self-overgrowth by non-ovicellate zooids can occur in J. mutabilis). Zooids with convex frontal shield of pinnate costae, with intercostal lacunae between adjacent costae; each costa typically with lumen pseudopore at or near tip. Gymnocyst negligible laterally; exposed in proximal corners of zooid or narrowly along proximal margin, or as extended proximal " caudal " zone; frontal pore chambers present or absent on proximal gymnocyst. Non-ovicellate zooids with approximately semicircular secondary orifice and two or more articulated oral spines. Avicularia lacking. Ovicellate zooids occurring in basal layer along with non-ovicellate zooids; as large (including ooecium) as or smaller than latter. Highly reduced dwarf ovicellate zooids occurring frontally, budded from basal or frontal pore chambers; sometimes present in basal layer, budded basally. Ovicell immersed, brood cavity intrazooidal, occupying distal part of ovicellate zooid. Ooecial complex in ovicellate zooids comprising ooecium and pair of modified, non-articulated, latero-oral spines arching over proximal part of ooecium and meeting or fusing at midline. Ooecium reduced to greater or lesser extent, hood-like, cap-like; kenozooidal, budded distally from maternal zooid or from roof of distal pore chamber, with one or more pseudopores; less commonly, ooecium vestigial, narrow, non-kenozooidal, produced as spine-like outgrowth from roof of distal basal pore chamber. Modified latero-oral spines either cylindrical, overlying proximal part of ooecium but not obscuring ooecial proximal edge, or wider and flatter, obscuring ooecial proximal edge; each spine bearing single lumen pseudopore at tip. Ancestrula cribriform, with oral spines, similar to but smaller than subsequent autozooids.	en	Dick, Matthew H., Grischenko, Andrei V., Gordon, Dennis P., Ostrovsky, Andrew N. (2021): The “ Cribrilina annulata " problem and new species of Juxtacribrilina (Bryozoa Cheilostomata: Cribrilinidae) from the North Pacific. Zootaxa 5016 (3): 333-364, DOI: 10.11646/zootaxa.5016.3.2
03BCE061A162FFD3B790FE5A5F69FA05.taxon	discussion	Remarks. Yang et al. (2018) erected Juxtacribrilina as a subgenus in Cribrilina to accommodate four species (C. annulata, C. corbula, C. mutabilis, and C. flavomaris) that differed from other species in Cribrilina in having the ooecium reduced in size, lacking avicularia, having modified latero-oral spines associated with the proximal ooecial margin, having dwarf ovicellate zooids, and having only a cribriform ancestrula. The discovery of additional species in Juxtacribrilina, as described herein, shows the existence of a sizeable clade that appears to have radiated independently from Cribrilina, justifying the change in the rank of Juxtacribrilina from subgenus to genus (López- Gappa et al. 2021).	en	Dick, Matthew H., Grischenko, Andrei V., Gordon, Dennis P., Ostrovsky, Andrew N. (2021): The “ Cribrilina annulata " problem and new species of Juxtacribrilina (Bryozoa Cheilostomata: Cribrilinidae) from the North Pacific. Zootaxa 5016 (3): 333-364, DOI: 10.11646/zootaxa.5016.3.2
03BCE061A162FFD8B790F9925CD3FD81.taxon	description	(Figs 2, 3; Table 1)	en	Dick, Matthew H., Grischenko, Andrei V., Gordon, Dennis P., Ostrovsky, Andrew N. (2021): The “ Cribrilina annulata " problem and new species of Juxtacribrilina (Bryozoa Cheilostomata: Cribrilinidae) from the North Pacific. Zootaxa 5016 (3): 333-364, DOI: 10.11646/zootaxa.5016.3.2
03BCE061A162FFD8B790F9925CD3FD81.taxon	materials_examined	Material examined. Specimen from NW Pacific region: ZIRAS 32 / 50735, western Kamchatka Shelf, Sea of Okhotsk, KamchatNIRO Collection, RV Professor Probatov, Station 6 (57.46722 ° N, 156.43417 ° E), one colony on interior surface of bivalve shell fragment, collected by T. B. Morozov, 16 June 2013, Sigsbee trawl, gravel bottom, 40 m depth. Other material examined: ZIRAS 33 / 50736, White Sea, Kandalaksha Bay, Chupa Inlet, near Matrenin Island, close to the Belomorskaia Educational and Research Station, Saint Petersburg State University (66.30917 ° N, 33.62861 ° E), six colonies on algae, dried and mounted on SEM stubs, collected by O. N. Kotenko, 20 September 2020, dredge and SCUBA diving, 5 – 10 m depth.	en	Dick, Matthew H., Grischenko, Andrei V., Gordon, Dennis P., Ostrovsky, Andrew N. (2021): The “ Cribrilina annulata " problem and new species of Juxtacribrilina (Bryozoa Cheilostomata: Cribrilinidae) from the North Pacific. Zootaxa 5016 (3): 333-364, DOI: 10.11646/zootaxa.5016.3.2
03BCE061A162FFD8B790F9925CD3FD81.taxon	description	Measurements. See Table 1. Description (Kamchatka specimen). Colonies encrusting, unilaminar (Fig. 2 A) except for reduced ovicellate zooids overgrowing zooids of basal layer (Fig. 2 E, F). Zooids oval or spindle-shaped in outline, delineated by deep groove, with little space between them (Fig. 2 B); gymnocyst negligible, sometimes exposed at proximal angles or on narrowing proximal part of zooid (Fig. 2 C, D); frontal pore chambers lacking. Frontal shield highly convex, comprising 13 – 17 costae (one abnormally large zooid with 20 treated as outlier, not included in measurements or average). Costae oval or flattened in cross section, with 8 – 10 intercostal lacunae in complete transverse series between them; in non-ovicellate zooids (Fig. 2 B), suboral pair of costae swollen, not wider than rest and often narrower, their tips sometimes slightly raised to form median projection. Each costa with single lumen pseudopore, at tip or somewhat proximal (Fig. 2 C) to tip. Non-ovicellate zooids with secondary orifice roughly semicircular, corners rounded; 4 (rarely, 3) blunt, articulated oral spines, lateral pair thicker than distal pair. Ovicellate zooids present in basal layer (Fig. 2 C, D) toward periphery or at margin; approximately as large (including ooecium) as non-ovicellate zooids, with 9 – 12 costae, suboral pair somewhat swollen, as wide as or somewhat wider than adjacent pair, often bi-lobed at ends, with more-distal lobes sometimes turned distally to form slight median projection. Ooecium (Fig. 2 C – F) hood-like, terminal; kenozooidal, budded from maternal zooid, occasionally bearing 1 or 2 basal pore chambers distally or disto-laterally; with thickened, non-articulated pair of modified latero-oral spines, each with single lumen pore, extending across proximal part of ooecium, usually meeting at midline, forming smooth or chevron-shaped arc, but not obscuring proximal edge of ooecium. Ooecium with 5 – 10 pseudopores, 1 – 4 in center and rest around peripheral margin. Median suture usually evident in proximal ooecial margin, sometimes extending distally to center of ooecium. Secondary orifice somewhat lozenge-shaped, bilabiate due to thickened costae proximally and modified spines distally. Ovicellate zooids present frontally (Fig. 2 E, F); similar to but somewhat shorter than (Table 1) basal ovicellate zooids; ooecial complex (Fig. 2 E, F) as in basal ovicellate zooids; sometimes with 1 ‒ 2 basal pore chambers distolaterally (Fig. 2 E). Frontally positioned ovicellate zooids appear associated with reparative growth or disruptions to zooidal packing. No true dwarf zooids observed; one possible developing dwarf (arrowhead, Fig. 2 D) evident as bud from basal pore chamber of zooid in basal layer.	en	Dick, Matthew H., Grischenko, Andrei V., Gordon, Dennis P., Ostrovsky, Andrew N. (2021): The “ Cribrilina annulata " problem and new species of Juxtacribrilina (Bryozoa Cheilostomata: Cribrilinidae) from the North Pacific. Zootaxa 5016 (3): 333-364, DOI: 10.11646/zootaxa.5016.3.2
03BCE061A162FFD8B790F9925CD3FD81.taxon	discussion	Remarks. Powell's (1967) description of J. annulata, based primarily on material from Vancouver Island in the boreal eastern Pacific and Prince Edward Island in the boreal western Atlantic, is problematic, because these populations are distant from one another and are not conspecific, as is evident from Powell's illustrations (see also below). The most detailed description of Atlantic J. annulata is that of Bishop (1994), based primarily on material from the North Atlantic and the Atlantic sector of the Arctic, and including SEM images. Both of these studies assumed that J. annulata is a single species with a circumpolar, Arctic-boreal distribution and included at least some information from Pacific specimens. Juxtacribrilina annulata was originally described from Greenland (Fabricius 1780), with a minimal description and no illustrations. Although Bishop (1994) did examine two specimens from Greenland, his description was based largely on material from Hardanger Fjord, Norway. Assuming that Bishop had illustrated type material, Yang et al. (2018) inadvertently referred to Hardanger Fjord as the type locality, but to our knowledge material from Greenland has not been redescribed, nor has a type specimen been designated, and J. annulata remains poorly defined taxonomically. Nonetheless, specimens from the boreal North Atlantic and European Arctic share zooidal characters that likely apply to J. annulata and that differ in key aspects from those in boreal Pacific populations. For the purposes of our study, and until the identity of J. annulata can be fixed, we consider Bishop's (1994) description of J. annulata to be representative of that species. The Kamchatka (Fig. 2), Hardanger Fjord (Bishop 1994, figs 25 – 32), and White Sea (Fig. 3) populations of J. annulata show some differences among one another, such as in zooid size, the predominant number of oral spines, and the number of costae and costal lacunae (Table 1). In addition, while no ancestrula was observed in the Kamchatka specimen, ancestrulae from the White Sea have only two oral spines (Fig. 3 F) (see also Nikulina 2002, fig. 18; Yagunova & Ostrovsky 2008, top photo and fig. 4), whereas the ancestrula from Hardanger Fjord illustrated by Bishop (1994, fig. 32) has four. These differences are minor compared to several major features shared by the three J. annulata populations, as follows. 1) Zooids in the basal layer have the marginal gymnocyst typically limited to a slight exposure along the proximal margin or in the proximal corners, and frontal pore chambers are lacking (Figs 2 B, 3 B; Bishop 1994, figs 25 – 30). 2) Ovicellate zooids occur in the basal layer, ranging (including the ooecium) from somewhat longer than to somewhat shorter than non-ovicellate zooids, but having fewer costae (Figs 2 A, C, D, 3 C; Bishop 1994, figs 25, 28, 29). 3) The ooecium is moderately well developed, hood-like, with a pair of modified latero-oral spines extending over the proximal surface and meeting at the midline, but leaving the ooecial proximal margin visible (e. g., Figs 2 C, 3 D; Bishop 1994, fig. 28). The ooecium has a faint median suture showing on the proximal margin and sometimes extending as far as the center; there are 5 – 10 pseudopores (occasionally fewer), typically arranged with one to four near the center and the rest distributed around the distal curvature (Figs 2 F, 3 D; Bishop 1994, fig. 28), but sometimes appearing haphazardly arranged. Some ooecia in the basal layer possess one or two basal pore chambers distally or disto-laterally (Fig. 3 C; Bishop 1994, fig. 29) and occasionally bud a distal zooid from these chambers; Fig. 2 C shows one such instance. 4) Dwarf ovicellate zooids are relatively uncommon. We did not observe any fully dwarfed zooids in the Kamchatka specimen, nor were any evident in representative material that Bishop (1994) illustrated from Hardanger Fjord. Likewise, we found none among hundreds of small colonies on algae collected in 2020 from the White Sea, although they do occur in that population (Fig. 3 E; Ostrovsky 1998, fig. 13 A; Nekliudova et al. 2019, fig. 5 B – D), where their presence or absence is related to life history. In a study of J. annulata inhabiting algal substrates in the White Sea, Nekliudova et al. (2019) found that colonies produce no dwarf zooids in their first season of sexual maturity, but after overwintering produce frontal dwarf ovicellate zooids near the colony margin. In the White Sea population, frontal dwarfs are thus entirely absent from many colonies, as in our specimens from 2020. By examining histological sections of White Sea specimens, Ostrovsky (1998) showed that frontally positioned dwarfs are not budded frontally, due to the lack of frontal pore chambers, but instead arise from basal pore chambers. The basally budded dwarfs are narrow as they extend interzooidally to the colony surface and then widen to form the frontal shield and ooecium (Fig. 3 E, arrowhead; Ostrovsky 1998, fig. 14). Once a colony has formed, basal (ovicellate and non-ovicellate) zooids in the central region of the colony are unable to bud frontal dwarfs from basal pore chambers, because the latter are prevented from budding by the tight packing of the autozooids, and in most cases are already being used for interzooidal communication. This explains the general paucity of dwarfs in the interior of J. annulata colonies. Occurrence. The presumed range of J. annulata is circum-Arctic (Powell 1968; Kluge 1975), southward to Nantucket Island in the western Atlantic (Osburn 1912), the Isle of Man in the eastern Atlantic (Hayward & Ryland 1998), and western Kamchatka, Sea of Okhotsk (one record, this study) in the western Pacific. This nominal species, however, has not been adequately described from its type locality (Greenland) or most of the rest of its range and potentially represents a complex of cryptic species.	en	Dick, Matthew H., Grischenko, Andrei V., Gordon, Dennis P., Ostrovsky, Andrew N. (2021): The “ Cribrilina annulata " problem and new species of Juxtacribrilina (Bryozoa Cheilostomata: Cribrilinidae) from the North Pacific. Zootaxa 5016 (3): 333-364, DOI: 10.11646/zootaxa.5016.3.2
03BCE061A169FFDFB790FD175B97FAB6.taxon	description	(Figs 4 – 8; Table 2) urn: lsid: zoobank. org: act: 312 B 8 C 48 - E 297 - 4377 - B 55 B- 11 B 8 EA 330130	en	Dick, Matthew H., Grischenko, Andrei V., Gordon, Dennis P., Ostrovsky, Andrew N. (2021): The “ Cribrilina annulata " problem and new species of Juxtacribrilina (Bryozoa Cheilostomata: Cribrilinidae) from the North Pacific. Zootaxa 5016 (3): 333-364, DOI: 10.11646/zootaxa.5016.3.2
03BCE061A169FFDFB790FD175B97FAB6.taxon	diagnosis	Diagnosis. Non-ovicellate zooids with 12 – 18 costae (colony averages 12.8 – 15.2), 3 or 4 oral spines (predominantly 3); intercostal lacunae medium-sized, 5 – 9 in transverse series (colony averages 6.9 – 7.5). Most zooids with 1 or 2 frontal pore chambers on proximal gymnocyst. Ovicellate zooids present in basal layer in subtidal specimens, rare in basal layer in intertidal specimens; roughly same size (including ooecium) as non-ovicellate zooids. Reduced and dwarf ovicellate zooids occur in basal layer near or at colony margin. Frontally budded dwarf ovicellate zooids typically present, often tightly packed in colony center, sometimes few or lacking; frontal shield usually with 3 or 4 costae. Ooecium reduced, cap-like, kenozooidal, bearing several irregular pseudopores in ovicellate zooids in basal layer, typically two transversely arranged pseudopores in dwarf zooids. Ancestrula cribriform, with 4 oral spines.	en	Dick, Matthew H., Grischenko, Andrei V., Gordon, Dennis P., Ostrovsky, Andrew N. (2021): The “ Cribrilina annulata " problem and new species of Juxtacribrilina (Bryozoa Cheilostomata: Cribrilinidae) from the North Pacific. Zootaxa 5016 (3): 333-364, DOI: 10.11646/zootaxa.5016.3.2
03BCE061A169FFDFB790FD175B97FAB6.taxon	etymology	Etymology. The specific name derives from Ezo, the old Japanese name for Hokkaido Island.	en	Dick, Matthew H., Grischenko, Andrei V., Gordon, Dennis P., Ostrovsky, Andrew N. (2021): The “ Cribrilina annulata " problem and new species of Juxtacribrilina (Bryozoa Cheilostomata: Cribrilinidae) from the North Pacific. Zootaxa 5016 (3): 333-364, DOI: 10.11646/zootaxa.5016.3.2
03BCE061A169FFDFB790FD175B97FAB6.taxon	materials_examined	Material examined. Holotype: YPM-IZ- 106549, Akkeshi Narrows, Akkeshi Bay, Hokkaido, Japan (43.048073 ° N, 144.85021 ° E), collected by M. H. Dick, July 2014, dredge, depth 6 – 8 m, encrusting dead bivalve shell; on SEM stub. Paratypes: YPM-IZ- 106550, YPM-IZ- 106551, YPM-IZ- 106552, same collection information as for holotype, on SEM stubs; YPM-IZ- 106553 (CaAkk- 1), Kakijima Island, Akkeshi-ko (= Akkeshi estuary), Akkeshi, Japan (43.04145 ° N, 144.86061 ° E), collected by A. V. Grischenko and M. H. Dick, 4 June 2004, rocky intertidal; YPM-IZ- 106554, YPM-IZ- 106555, NHM 2006.2.27.31, Aininkappu, Akkeshi Bay, Japan (42.99267 ° N, 144.85500 ° E), collected by A. V. Grischenko and M. H. Dick, 5 June 2004, rocky intertidal, on SEM stubs; NHM 2006.2.27.50, NHM 2006.2.27.51, Daikokujima I., Akkeshi Bay, Japan (42.95433 ° N, 144.87567 ° E), collected by A. V. Grischenko and M. H. Dick, 7 June 2004, rocky intertidal, on SEM stubs; NHM 2006.2.27.52, Barasan Cape, Akkeshi Bay, Japan (43.02417 ° N, 144.83683 ° E), collected by A. V. Grischenko and M. H. Dick, 3 July 2004, rocky intertidal, on SEM stub. Other material: YPM-IZ- 106556, East Tongass Narrows, Ketchikan, Alaska (55.31657 ° N, 131.51990 ° W), collected by M. H. Dick, 11 – 12 September 2003, rocky intertidal; ZIRAS 01 / 50734, Bering Island (Commander Islands), Station 10, Cape Gaupta (Kitovyy) (55.24583 ° N, 165.92056 ° E), one colony encrusting barnacle fragment, collected by A. V. Grischenko, 1 August 1991, lower horizon of rocky-boulder intertidal, Laminaria dentigera zone, rock face.	en	Dick, Matthew H., Grischenko, Andrei V., Gordon, Dennis P., Ostrovsky, Andrew N. (2021): The “ Cribrilina annulata " problem and new species of Juxtacribrilina (Bryozoa Cheilostomata: Cribrilinidae) from the North Pacific. Zootaxa 5016 (3): 333-364, DOI: 10.11646/zootaxa.5016.3.2
03BCE061A169FFDFB790FD175B97FAB6.taxon	description	Measurements. See Table 2. Description. Colonies encrusting, multiserial, unilaminar except for frontally positioned dwarf zooids (Figs 4 B, C, 6 A, B, 7 A, 8 A). Non-ovicellate zooids (Figs 4 A, B, 5 B, 6 B, 7 B, 8 B) oval or sometimes spindle-shaped, widest in middle, closely packed; gymnocyst negligible, sometimes wider in proximal corners or extending proximally as long caudal zone. Most ovicellate and non-ovicellate zooids in basal layer bearing 1 or 2 (occasionally 3) oval, triangular, or quadrangular frontal pore chambers on proximal gymnocyst (Figs 4 A, B, 7 B, 8 B); frontal pore chambers multiporous (Figs 4 E, 5 E). Frontal shield highly convex, sometimes slightly keeled; 12 – 18 costae (average values, 12.8 – 15.2) in non-ovicellate zooids; oval or somewhat flattened in cross section; suboral pair not appreciably wider than rest but sometimes somewhat swollen, often with small, fused tips extending fronto-distally (Figs 4 B, E, 5 B, 8 B); each costa with minute lumen pseudopore at or near tip. Intercostal lacunae moderate in size; circular, oval, subcircular, or heart-shaped (end closest to midline tapering); 5 – 9 in transverse series (average per colony, 6.9 – 7.5), most zooids with 6 – 8. Secondary orifice of non-ovicellate zooids roughly semicircular, wider than long, with corners widely rounded. Oral spines most commonly 3 (sometimes 4; rarely, 2 or 5), lateral pair often longer and heavier than spine (s) in between; single median spine usually flattened, widest basally, tapering distally, paired median spines usually cylindrical, occasionally fused. Ovicellate zooids lacking in basal layer (except at margin) in intertidal colonies but usually present in basal layer in subtidal colonies, either scattered individually (Fig. 6 B) or in circumferential bands (Figs 5 A – C, 6 A). Ovicellate zooids (including ooecium) in colony interior as large as non-ovicellate zooids (Figs 5 A, 6 B), with 11 – 17 costae; suboral pair often markedly wider than rest, with part of fused ends projecting fronto-distally to form mucro proximal to and overhanging orifice. Mucro sharp, with double or single tip, or wide, blunt, flattened. Reduced zooids (with as few as ~ 8 costae) and dwarf zooids sometimes occurring in basal layer at or near colony margin (Figs 4 F, 6 A, 7 B, 8 A, B, D). Narrow, concentric zone of somewhat to markedly reduced ovicellate zooids seen toward margin of one colony (Fig. 6 A). Ovicellate zooids in basal layer with distal pore chamber, sometimes nonbudding and evident as rounded opening below ooecium (Fig. 5 B, arrow), but usually budding distal zooid (Fig. 5 B, arrowheads); basal ovicellate zooids thus commonly occurring in columnar series with other zooids (Figs 5 A, 6 A). Ooecium in basal zooids (Figs 5 B – D, 6 A) kenozooidal, arising from roof of distal pore chamber of maternal zooid (Fig. 6 D), connecting with pore chamber via two or more pores (Fig. 6 D, F); reduced, cap-shaped, abutting arching, modified pair of wide, flattened, non-articulated, latero-oral spines fusing at midline; modified spines tapering and twisted toward midline, forming distal convexity of secondary orifice; ends often flattened and raised to become blunt median projection. Secondary orifice shaped like smiling mouth flanked by lips (widened suboral costae and modified latero-oral spines). Ooecium of zooids in basal layer with 2 – 6 small to moderately large, scattered, circular to irregular pseudopores; proximal margin of cap not visible inside secondary orifice. Dwarf zooids often abundant in center of colony (Figs 4 C, 6 B, 7 A, 8 A), budded from frontal pore chambers of basal zooids (Fig. 4 E); costal shield (Figs 4 D, 6 C, 8 C, F) reduced to 3 or 4 costae (occasionally 2 or 5) including suboral pair; suboral pair often curved in distal direction, forming sharp mucro or evenly tapering triangular process. Ooecium appears to be kenozooidal, budded distally from maternal zooid. Ooecial complex (Figs 4 D, 6 C, 7 C, 8 C, E, F) in dwarf zooids similar to that in ovicellate zooids in basal layer; modified latero-oral spines not twisted as much and usually not forming median process; cap-like ooecium similar but narrower, usually with 2 circular or oval, transversely arranged pseudopores (Figs 4 D, 6 C, 7 C, 8 C), these sometimes partly merged. Margin of ooecial cap meeting modified spines; sometimes slightly thickened and bearing 1 or 2 tiny additional pseudopores. Dwarf zooids sometimes with 1 – 2 basal pore chambers evident in distal (in ooecial kenozooid), disto-lateral, lateral, or proximo-lateral positions (Figs 6 C, 8 C, E, F). Marginal, basally budded dwarf zooids (Figs 4 F, 8 D) more common in intertidal than in subtidal colonies. Ancestrula (Fig. 5 F) cribriform, similar to but smaller than subsequent zooids, with 4 articulated oral spines.	en	Dick, Matthew H., Grischenko, Andrei V., Gordon, Dennis P., Ostrovsky, Andrew N. (2021): The “ Cribrilina annulata " problem and new species of Juxtacribrilina (Bryozoa Cheilostomata: Cribrilinidae) from the North Pacific. Zootaxa 5016 (3): 333-364, DOI: 10.11646/zootaxa.5016.3.2
03BCE061A169FFDFB790FD175B97FAB6.taxon	discussion	Remarks. Grischenko et al. (2007) described this species in detail (as Cribrilina annulata) from the intertidal zone at Akkeshi Bay, Japan. Juxtacribrilina ezoensis n. sp. differs from J. annulata in several clear-cut ways. In J. ezoensis n. sp., most zooids in the basal layer have one or two frontal pore chambers on the proximal gymnocyst, whereas in J. annulata, frontal pore chambers occur only rarely and are usually entirely absent. Consequently, J. ezoensis n. sp. can readily bud frontal dwarfs at any time during colony astogeny, and in any part of the colony. In contrast, as far as is known, J. annulata produces frontal dwarfs only from basal pore chambers, which once a colony has formed are largely unavailable for budding in the center of the colony. Frontal dwarfs are therefore much more common in J. ezoensis n. sp. and often occur abundantly in the central region of the colony, whereas in J. annulata they are uncommon and usually occur at or near the colony margin. The two species also differ markedly in the size of the ooecium and the form and structure of the ooecial complex. In J. annulata, the ooecium is hood-like and only moderately reduced from the full-sized ooecium seen in most cheilostomes (Ostrovsky 1998, 2008, 2013, 2020; Martha et al. 2020); the modified latero-oral spines arching over the proximal part of the ooecium retain a spine-like appearance and do not obscure the proximal ooecial margin (Figs 2 D – F, 3 C, D). The ooecium is kenozooidal, budded from the distal area of the maternal zooid occupied by the septulum of the forming basal pore chamber; in both basal and dwarf zooids, the kenozooidal ooecium may itself give rise to one or two distal pore chambers, but only rarely (Fig. 2 C) has been observed to produce another autozooid distally. Ovicellate zooids in the basal layer thus do not usually bud a distal autozooid, and usually do not lie in columnar series with following zooids (Fig. 2 D). In contrast, the ooecium in J. ezoensis n. sp. is reduced to a broad, low cap and is more highly integrated with the modified spines, which are flatter and broader, forming the transverse bar that comprises the proximal margin of the ooecial complex and whose concave proximal edge contributes to the shape of the secondary orifice (Fig. 5 B, D). The ooecium in basal zooids is kenozooidal, but arises from the roof of the distal pore chamber (Fig. 6 D), to which it is connected via one or more communication pores (Fig. 6 F), thus leaving the pore chamber free to produce another zooid distally (Fig. 6 E); basal ovicellate zooids are thus typically arranged in columnar series (Figs 5 A, B, 6 A, B). In the infrequent cases where a basal ovicellate zooid is not in columnar series, the distal pore chamber remains evident as an oval opening below the ooecium (Fig. 5 B, arrow). Most measurements for zooids in the basal layer overlap considerably between J. annulata (Table 1) and J. ezoensis n. sp. (Table 2), especially considering variation among the populations within each of these species. However, in White Sea J. annulata, dwarfs are larger for all measurements and have more costae (range 4 – 7, average 5.7) than dwarfs in J. ezoensis n. sp. (overall range 2 – 5; range of averages 3.2 – 3.7). Ooecia in J. annulata have more pseudopores, both in basal ovicellate zooids (Kamchatka, 5 – 10; White Sea, 5 – 7) and in frontal dwarfs (White Sea, 3 – 5), than J. ezoensis n. sp. (basal ooecia, 2 – 6; dwarf ooecia, usually 2). At Akkeshi, subtidal colonies of J. ezoensis n. sp. have " full-sized " ovicellate zooids in the basal layer, whereas intertidal colonies lack them (Grischenko et al. 2007). Upon examination of additional intertidal material from Akkeshi, we again saw no " full-sized " ovicellate zooids in the basal layer, but did find dwarfs to occur in the basal layer at the colony margin (Figs 4 C, F) — as was also the case for intertidal specimens of J. ezoensis n. sp. from the Commander Islands (Fig. 7 B) and Ketchikan (Fig. 8 A, D). While the differences in the intra-colony complement of ovicellate zooids between intertidal and subtidal material might seem to indicate two distinct species, we believe that the two forms are conspecific. The measurements for non-ovicellate and dwarf zooids (Table 2) are remarkably similar between intertidal and subtidal specimens from Akkeshi, as are the number of intercostal lacunae in nonovicellate zooids, the form of the ooecium, and the form of the costal shield in dwarf zooids. Previous studies have found the distribution of various types of ovicellate zooids (frontal or basal; full-sized or dwarf) in J. annulata colonies (Nekliudova et al. 2019), and of zooids having different types of frontal shield (Ito et al. 2015) in J. mutabilis, to be related to life history. In the case of J. ezoensis n. sp. at Akkeshi, we speculate that the paucity of ovicellate zooids in the basal layer in intertidal colonies is related to the less stable, more stressful intertidal environment compared to the subtidal environment. Subtidal colonies of J. ezoensis n. sp. at Akkeshi typically reach larger sizes than intertidal colonies and almost certainly overwinter. It may be that intertidal colonies, on the other hand, settle each season from subtidally produced larvae and do not overwinter, foregoing the production of larger ovicellate zooids in the basal layer in favor of frontal and marginal dwarfs. Frontal dwarfs typically appear first toward the colony center and are added centrifugally as the colony grows, presumably as conditions are favorable, allowing flexibility in the timing of reproduction and the resources devoted to it. Marginal dwarfs likely appear late in the season, as colony growth begins to wind down. Among the Pacific Juxtacribrilina species treated in the subsequent descriptions, J. ezoensis n. sp. is most similar to J. pushkini n. sp. described next; a comparison of the two is deferred to the Remarks section for the latter. Occurrence. J. ezoensis n. sp. has a trans-Pacific distribution; it was common intertidally and subtidally at Akkeshi Bay, Japan, and occurred intertidally in the Commander Islands (one colony) and at Ketchikan, Alaska (one colony).	en	Dick, Matthew H., Grischenko, Andrei V., Gordon, Dennis P., Ostrovsky, Andrew N. (2021): The “ Cribrilina annulata " problem and new species of Juxtacribrilina (Bryozoa Cheilostomata: Cribrilinidae) from the North Pacific. Zootaxa 5016 (3): 333-364, DOI: 10.11646/zootaxa.5016.3.2
03BCE061A16EFFC3B790FA035A59FD81.taxon	description	(Figs 9, 10; Table 3) urn: lsid: zoobank. org: act: CDED 1 A 27 - C 2 BC- 4867 - 9 F 62 - E 395036 D 70 F 5	en	Dick, Matthew H., Grischenko, Andrei V., Gordon, Dennis P., Ostrovsky, Andrew N. (2021): The “ Cribrilina annulata " problem and new species of Juxtacribrilina (Bryozoa Cheilostomata: Cribrilinidae) from the North Pacific. Zootaxa 5016 (3): 333-364, DOI: 10.11646/zootaxa.5016.3.2
03BCE061A16EFFC3B790FA035A59FD81.taxon	diagnosis	Diagnosis. Non-ovicellate zooids with 14 – 19 costae, 3 – 5 oral spines (usually 3 or 4); intercostal lacunae small, 6 – 10 in transverse series. One or two frontal pore chambers on proximal gymnocyst of most zooids. Ovicellate zooids (including ooecium) roughly same size as non-ovicellate zooids occurring but uncommon in basal layer. Reduced or dwarf marginal ovicellate zooids occurring; frontal dwarfs often abundant in colony center. Frontal shield of dwarf zooids as large as or larger than ooecial complex, with 5 or 6 costae. Ooecium in zooids in basal layer moderately reduced, cap-like, with several irregular pseudopores; ooecium in dwarf zooids typically with 2 transversely arranged, oval or irregular pseudopores, with additional pair of smaller pseudopores sometimes present. Ancestrula cribriform, with 4 or 5 oral spines.	en	Dick, Matthew H., Grischenko, Andrei V., Gordon, Dennis P., Ostrovsky, Andrew N. (2021): The “ Cribrilina annulata " problem and new species of Juxtacribrilina (Bryozoa Cheilostomata: Cribrilinidae) from the North Pacific. Zootaxa 5016 (3): 333-364, DOI: 10.11646/zootaxa.5016.3.2
03BCE061A16EFFC3B790FA035A59FD81.taxon	etymology	Etymology. Named for Dr. Alexander Fedorovitch Pushkin (1938 – 2019), research associate at the Zoological Institute, Russian Academy of Sciences, zoologist and hydrobiologist, working mostly on Arctic and Antarctic benthic invertebrates and a leading world expert on Pycnogonida.	en	Dick, Matthew H., Grischenko, Andrei V., Gordon, Dennis P., Ostrovsky, Andrew N. (2021): The “ Cribrilina annulata " problem and new species of Juxtacribrilina (Bryozoa Cheilostomata: Cribrilinidae) from the North Pacific. Zootaxa 5016 (3): 333-364, DOI: 10.11646/zootaxa.5016.3.2
03BCE061A16EFFC3B790FA035A59FD81.taxon	materials_examined	Material examined. Holotype: YPM-IZ- 100361, vicinity of Ketchikan, Alaska, 9 – 12 September 2003, lower rocky intertidal, - 0.46 to - 0.49 m relative to mean low water level (MLWL); on rock, mounted on SEM stub. Paratypes: YPM-IZ- 106544, YPM-IZ- 106545, YPM-IZ- 106546, East Tongass Narrows, Ketchikan, Alaska (55.31657 ° N, 131.51990 ° W), 11 – 12 September 2003, lower rocky intertidal; YPM-IZ- 106547, collection information as for holotype, on SEM stub; YPM-IZ- 100481, Settlers Cove, Ketchikan vicinity, Alaska (55.51161 ° N, 131.72517 ° W), 9 September 2003, lower rocky intertidal, - 0.46 m from MLWL; all on SEM stubs. Other material: YPM-IZ- 106548, collection information as for holotype; on SEM stub. All specimens collected by M. H. Dick.	en	Dick, Matthew H., Grischenko, Andrei V., Gordon, Dennis P., Ostrovsky, Andrew N. (2021): The “ Cribrilina annulata " problem and new species of Juxtacribrilina (Bryozoa Cheilostomata: Cribrilinidae) from the North Pacific. Zootaxa 5016 (3): 333-364, DOI: 10.11646/zootaxa.5016.3.2
03BCE061A16EFFC3B790FA035A59FD81.taxon	description	Measurements. See Table 3. Description. Colonies encrusting, multiserial, unilaminar except for frontal dwarf zooids (Fig. 9 D). Nonovicellate zooids (Figs 9 A, 10 C, D) typically oval in outline, widest in middle, closely packed; gymnocyst negligible, sometimes wider in proximal corners or tapering proximally as long caudal zone. Ovicellate and non-ovicellate zooids in basal layer each bearing 1 or 2 (exceptionally 3) oval, triangular, or quadrangular frontal pore chambers on proximal gymnocyst (Fig. 9 A – C); frontal pore chambers multiporous. Frontal shield convex, tumid, sometimes with slight keel. Costae oval or somewhat flattened in cross section, 14 – 19 in number (average per colony 15.1 – 17.1) in non-ovicellate zooids, suboral pair slightly swollen but not wider than rest, sometimes with small, fused tips extending fronto-distally (Fig. 10 C) to form slight mucro; each costa with minute lumen pseudopore at or near tip. Intercostal lacunae small; circular, oval, or subcircular; 6 – 10 (usually 8 or 9) in transverse series (average per colony 8.2 – 9.0). Secondary orifice (Figs 9 A, 10 C) roughly semicircular, markedly wider than long; oral spines 3 or 4 (rarely 5), lateral pair often longer and heavier than those in between; median spine (s) flattened, basally wide, tapering, single or paired, in latter case sometimes fused. Ovicellate zooids uncommonly present in basal layer (Fig. 9 B, C), scattered singly and in small groups. Ovicellate zooids in colony interior as large (including ooecium) as non-ovicellate zooids, with 11 – 15 costae; suboral pair of costae wider, thicker than rest, with part of fused tips often extending fronto-distally to form sharp or blunt, often flattened mucro (Fig. 9 B) proximal to and overhanging orifice. Ooecial complex (Fig. 9 B, C) in zooids in basal layer comprising ooecium and pair of widened, flattened, nonarticulated, modified latero-oral spines meeting at midline to form proximal margin; spines tapering and twisted in distal half to two-thirds to form concave proximal margin of ooecial complex, and sometimes raised and thickened in midline to form low mucro limited by conspicuous " fold " near middle of modified spine on each side (small arrowheads, Fig. 9 B); each spine bearing lumen pseudopore near truncate tip at midline. Proximal margin of ooecial complex and proximal margin of orifice both labiate in appearance. Ooecium moderately reduced, cap-like, roughly twice as wide as long, with 2 – 3 moderately large, circular to irregular pseudopores; ooecial cap abutting rather than passing underneath pair of modified spines forming proximal margin of complex; cap sometimes with thickened or folded proximal margin (abutting modified pair of spines) bearing additional 2 or 3 small pseudopores (Fig. 9 C). Ooecium likely kenozooidal, arising from roof of distal pore chamber, which buds next-distal zooid (Fig. 9 B, arrow) or fails to bud, remaining as circular opening (Fig. 9 B, arrowheads); basal ovicellate zooids thus often but not always in columnar series with other zooids. Dwarf ovicellate zooids typically present in center of colony (Fig. 9 D), often densely packed, budded from proximal frontal pore chambers of zooids in basal layer; frontal shield (Fig. 10 A, B) conspicuous, occupying half of zooid length; costae usually 5 or 6 (rarely, 7 or 8); suboral pair thicker than others, similar to suboral pair in basal ovicellate zooids, often forming median mucro; other costae centripetally directed toward midline, forming tight shield, with small, circular intercostal lacunae. Ooecial complex as described above for basal ovicellate zooids; ooecium small, cap-like (Fig. 10 B), but kenozooidal, terminal, budded from maternal zooid; typically bearing two moderately large, circular or oval, transversely arranged pseudopores; proximal margin of ooecium (abutting modified pair of latero-oral spines) sometimes thickened and bearing additional smaller pseudopores (Fig. 10 B, arrowheads). Dwarf zooids sometimes with up to three basal pore chambers in distal (ooecial), disto-lateral, or lateral positions (Fig. 10 A). Dwarf zooids occasionally present marginally in basal layer (Fig. 10 C). Ancestrula (Fig. 10 D) cribriform, similar to but smaller than subsequent zooids, with 10 – 13 costae, surrounded by 6 periancestrular zooids (1 distal, 2 disto-lateral, 2 proximolateral, 1 proximal); 4 (n = 2) or 5 (n = 5) oral spines; if 5, distalmost, median spine thinner and shorter than rest.	en	Dick, Matthew H., Grischenko, Andrei V., Gordon, Dennis P., Ostrovsky, Andrew N. (2021): The “ Cribrilina annulata " problem and new species of Juxtacribrilina (Bryozoa Cheilostomata: Cribrilinidae) from the North Pacific. Zootaxa 5016 (3): 333-364, DOI: 10.11646/zootaxa.5016.3.2
03BCE061A16EFFC3B790FA035A59FD81.taxon	discussion	Remarks. Dick et al. (2005) described this species (as Cribrilina annulata) from the same locality, Ketchikan, Alaska. The zooidal measurements differ somewhat between that study and this, possibly due to different methods of measurement (light microscopy using an ocular micrometer in the previous study; measurements from SEM images here). The costal count for non-ovicellate zooids also differs (12 – 17 previously; 14 – 19 here), with the lower previous numbers possibly due to inclusion of some zooids in the zone of astogenetic change and a lower sample size. The number of oral spines in the ancestrula was previously given as four, but ancestrulae more often have five spines. Juxtacribrilina pushkini n. sp. differs from J. annulata in the same ways as does J. ezoensis n. sp. (see above). Juxtacribrilina pushkini n. sp. and J. ezoensis n. sp. are quite similar in most aspects, including zooidal dimensions, the number of oral spines, and the morphology of ovicellate zooids in the basal layer. However, non-ovicellate zooids in J. pushkini n. sp. tend to have more costae (14 – 19, overall average 16.2) than in J. ezoensis n. sp. (12 – 18, overall average 14.8); the intercostal lacunae in basal zooids are smaller and more numerous (8 or 9 in transverse series) than in J. ezoensis n. sp. (usually 6 – 8); the costal shield in dwarf zooids is larger, tighter, and more convex, bearing 5 or 6 costae, compared to J. ezoensis n. sp. (smaller and flatter, typically bearing 3 or 4 costae); and there is a conspicuous angular " fold " (Fig. 9 B, small arrowheads) near the middle of each of the modified latero-oral spines of ovicellate zooids in the basal layer, whereas this fold is lacking in J. ezoensis n. sp.. The ancestrula in J. pushkini n. sp. has four or five oral spines but more commonly five; in J. ezoensis n. sp., only four oral spines were observed. The two species are most readily distinguished by the size and number of the intercostal lacunae in zooids in the basal layer, and by the number of costae and size of the costal shield in dwarf zooids. Occurrence. At present, J. pushkini n. sp. is known only from Ketchikan, Alaska.	en	Dick, Matthew H., Grischenko, Andrei V., Gordon, Dennis P., Ostrovsky, Andrew N. (2021): The “ Cribrilina annulata " problem and new species of Juxtacribrilina (Bryozoa Cheilostomata: Cribrilinidae) from the North Pacific. Zootaxa 5016 (3): 333-364, DOI: 10.11646/zootaxa.5016.3.2
03BCE061A172FFC6B790FD145B73FD49.taxon	description	(Fig. 11; Table 4) urn: lsid: zoobank. org: act: C 88 EE 938 - BEEE- 4 FD 3 - B 3 B 4 - EBA 40 EEE 38 EA	en	Dick, Matthew H., Grischenko, Andrei V., Gordon, Dennis P., Ostrovsky, Andrew N. (2021): The “ Cribrilina annulata " problem and new species of Juxtacribrilina (Bryozoa Cheilostomata: Cribrilinidae) from the North Pacific. Zootaxa 5016 (3): 333-364, DOI: 10.11646/zootaxa.5016.3.2
03BCE061A172FFC6B790FD145B73FD49.taxon	diagnosis	Diagnosis. Non-ovicellate zooids with 12 – 16 costae, 3 or 4 oral spines (predominantly 3); intercostal lacunae 6 – 9 in transverse series. Most zooids in basal layer with 1 or 2 frontal pore chambers on proximal gymnocyst. Ovicellate zooids occurring in basal layer, roughly same size (including ooecium) as non-ovicellate zooids; lacuna evident in midline at base of projection formed by suboral costae. Frontally budded dwarf ovicellate zooids occurring profusely in colony center; 4 – 7 costae (average 4.9); dwarf costal shield variable; tight, with wide, tapering costae and small intercostal lacunae, or reticulate, with narrow costae and very wide intercostal lacunae. Dwarf ooecium moderately reduced, cap-like, with a few irregular pseudopores; typically bearing single slit-like, oval, or ramifying pseudopore in midline.	en	Dick, Matthew H., Grischenko, Andrei V., Gordon, Dennis P., Ostrovsky, Andrew N. (2021): The “ Cribrilina annulata " problem and new species of Juxtacribrilina (Bryozoa Cheilostomata: Cribrilinidae) from the North Pacific. Zootaxa 5016 (3): 333-364, DOI: 10.11646/zootaxa.5016.3.2
03BCE061A172FFC6B790FD145B73FD49.taxon	etymology	Etymology. Named for Dr. Andrej Alexandrovitch Dobrovolskij (1939 – 2019), Head of the Department of Invertebrate Zoology at Saint Petersburg State University, Russia, zoologist, protistologist, and parasitologist, specializing on Digenea.	en	Dick, Matthew H., Grischenko, Andrei V., Gordon, Dennis P., Ostrovsky, Andrew N. (2021): The “ Cribrilina annulata " problem and new species of Juxtacribrilina (Bryozoa Cheilostomata: Cribrilinidae) from the North Pacific. Zootaxa 5016 (3): 333-364, DOI: 10.11646/zootaxa.5016.3.2
03BCE061A172FFC6B790FD145B73FD49.taxon	materials_examined	Material examined. Holotype and paratype: ZIRAS 01 / 50733 - A and - B, single pebble with one mature colony (holotype, A) and one ancestrular colony (paratype, B), Krabovaya Bay, Cape Severnyy Vkhodnoy, Shikotan Island, Lesser Kuril Chain (43.87722 ° N, 146.82000 ° E), collected by A. P. Tsurpalo, 28 August 1996; mid-intertidal, rocky shore; prepared for SEM examination.	en	Dick, Matthew H., Grischenko, Andrei V., Gordon, Dennis P., Ostrovsky, Andrew N. (2021): The “ Cribrilina annulata " problem and new species of Juxtacribrilina (Bryozoa Cheilostomata: Cribrilinidae) from the North Pacific. Zootaxa 5016 (3): 333-364, DOI: 10.11646/zootaxa.5016.3.2
03BCE061A172FFC6B790FD145B73FD49.taxon	description	Measurements. See Table 4. Description. Colony (Fig. 11 A) encrusting, multiserial, unilaminar except for frontally located dwarf zooids. Zooids (Fig. 11 B – D) oval or spindle shaped, sometimes extended proximally; gymnocyst irregular, present as intermittent narrow band, widest at proximolateral corners (Fig. 11 B, C), bearing 1 or 2 small, oval frontal pore chambers in most zooids. Frontal shield highly convex, sometimes slightly keeled; non-ovicellate zooids with 12 – 16 costae (average 14.5); intercostal lacunae circular, oval, irregular, or heart-shaped, 6 – 9 (average 7.5) in transverse series, each costa with minute lumen pseudopore at or near tip. In non-ovicellate zooids, suboral costae similar to or somewhat wider than others in width; ends of suboral costae often biramous, with more-distal pair of rami extending frontally or fronto-distally, meeting at midline to form projection proximal to orifice. Secondary orifice roughly semicircular, corners relatively sharp; 3 – 4 (average 3.1) articulated oral spines present; median one or two spines thinner than lateral spines, flattened, wider basally and tapering distally if single, cylindrical if paired. Ovicellate zooids abundant in basal layer (Fig. 11 A, C), occurring in broad band five or six zooid generations wide between central region and margin, with ovicellate zooids again giving rise to non-ovicellate zooids at margin. Ovicellate zooids in basal layer with suboral costae markedly wider than rest (Fig. 11 C), each with terminal lumen pseudopore; bifurcate at end, with more-distal rami meeting at midline to form broad, labiate flange directed fronto-distally and overhanging orifice; median lacuna at base of flange. Ooecial complex (Fig. 11 C) with broad, thickened, non-articulated, modified latero-oral spines, each with near-terminal lumen pseudopore, arching toward and meeting at midline, twisted and narrower distally, often flattened and elevated, giving rise to wide, blunt, frontodistally directed flange. Secondary orifice of ovicellate zooids wider than long, with appearance of smiling mouth, flanked by proximal (suboral costae) and distal (latero-oral spines) labiate flanges. Ooecium likely kenozooidal, arising from roof of distal pore chamber; cap-shaped, broader than long, proximal edge overlain by modified spines but not visible in frontal view; with one to several irregular or ramifying pseudopores. Basal ovicellate zooids often in columnar series with other zooids. No. oral spines 3.1 (3 – 4) No. costae 14.5 (12 – 16) No. intercostal lacunae 1 7.5 (6 – 9) Ovicellate zooids, basal n = 7, 1 Zooid length 2 552 (495 – 685) Zooid width 294 (258 – 353) Orifice length 77 (47 – 99) Orifice width 180 (163 – 199) Ooecial complex length 146 (116 – 172) Ooecial complex width 234 (174 – 280) Ooecium length 62 (33 – 89) Ooecium width 172 (151 – 194) No. costae 12.0 (11 – 13) Frontal dwarf zooids n = 13, 1 Zooid length 2 317 (225 – 577) Zooid width 227 (178 – 357) Orifice length 80 (52 – 130) Orifice width 151 (117 – 231) Ooecial complex length 143 (95 – 231) Ooecial complex width 200 (138 – 312) Ooecium length 71 (52 – 111) Ooecium width 142 (112 – 215) No. costae 4.9 (4 – 7) 1 Number of lacunae in transverse series between second and third pairs of costae from orifice. 2 Includes the ooecium. Frontally budded dwarf zooids (11 D, E) scattered abundantly in colony center; ooecial complex similar to that of larger, basal ovicellate zooids but with flanges proximal and distal to orifice less prominent; suboral flange comprising blunt or sharp " double " mucro formed from cylindrical, more-distal rami of suboral costae; distal, elevated flange on proximal margin of ooecial complex often lacking. Ooecium of dwarf zooids cap-like, kenozooidal, terminal, variable in size and shape, as wide as or wider than long; with one or two circular or elongate pseudopores in midline, or single ramifying pseudopore. Dwarf zooids with 4 – 7 costae (including suboral pair; average 4.9); costal shield either nearly flat, with wide, tapering costae and small intercostal lacunae, or markedly convex, inflated, with narrow costae and very wide intercostal lacunae forming a reticulate shield (both types evident in Fig. 11 E). Ooecium sometimes with basal pore chamber distally (Fig. 11 E). Only one reduced-size ovicellate zooid observed in basal layer — a dwarf zooid closer to colony center than to margin (not illustrated). One newly settled ancestrula observed (Fig. 11 F); cribriform; tight shield composed of 14 costae; with 5 oral spines, median spine shorter and thinner than rest.	en	Dick, Matthew H., Grischenko, Andrei V., Gordon, Dennis P., Ostrovsky, Andrew N. (2021): The “ Cribrilina annulata " problem and new species of Juxtacribrilina (Bryozoa Cheilostomata: Cribrilinidae) from the North Pacific. Zootaxa 5016 (3): 333-364, DOI: 10.11646/zootaxa.5016.3.2
03BCE061A172FFC6B790FD145B73FD49.taxon	discussion	Remarks. Juxtacribrilina dobrovolskii n. sp. differs from J. annulata in the same ways as do J. pushkini n. sp. and J. ezoensis n. sp.: most basal zooids bear one or two frontal pore chambers on the proximal gymnocyst; frontal dwarf zooids arise from these chambers and are often abundant and closely packed in the colony center; and the ooecium is cap-like and closely integrated with the modified latero-oral spines forming its proximal boundary. Juxtacribrilina dobrovolskii n. sp. differs most conspicuously from J. pushkini n. sp. and J. ezoensis n. sp. in features of the dwarf zooids. In J. dobrovolskii n. sp., the dwarf ooecium usually has a single irregular (slit-like, elliptical, or ramifying) pseudopore in the midline, and the costal shield bears 4 – 7 costae and is variable (Fig. 11 E), either flatter with fewer costae, or more tumid, with the costae forming a reticulate shield. In the latter two species, the dwarf ooecium usually has two transversely arranged pseudopores, and the costal shield is either somewhat tumid, usually with 5 or 6 costae (J. pushkini n. sp.; Fig. 10 A, B), or flatter, usually with 3 or 4 costae (J. ezoensi s n. sp.; Figs 4 D, 6 C). Basal zooids in both J. dobrovolskii n. sp. (Fig. 11 C) and J. ezoensis n. sp. (Fig. 4 A) have larger intercostal lacunae relative to costal width than J. pushkini n. sp. (Fig. 9 A), and this is reflected in lower average numbers of intercostal lacunae in transverse series: 7.5, 6.9 – 7.5, and 8.2 – 9.0, respectively (Tables 2 – 4). Likewise, basal ovicellate zooids in both J. dobrovolskii n. sp. (Fig. 11 C) and J. ezoensis n. sp. (Fig. 5 C, D) tend to lack the distinct fold, seen in J. pushkini n. sp. (Fig. 9 B), in each of the modified latero-oral spines contributing to the ooecial complex. Finally, the conspicuous median lacuna at the base of the flange formed by the suboral costae in basal ovicellate zooids in J. dobrovolskii n. sp. (Fig. 11 B, C) is lacking in J. pushkini n. sp. (Fig 9 B) and J. ezoensis n. sp. (Fig. 5 B – D). Occurrence. Shikotan Island in the Lesser Kuril Chain is the only known locality for J. dobrovolskii n. sp.	en	Dick, Matthew H., Grischenko, Andrei V., Gordon, Dennis P., Ostrovsky, Andrew N. (2021): The “ Cribrilina annulata " problem and new species of Juxtacribrilina (Bryozoa Cheilostomata: Cribrilinidae) from the North Pacific. Zootaxa 5016 (3): 333-364, DOI: 10.11646/zootaxa.5016.3.2
03BCE061A177FFC9B790FD5F5B8CFE85.taxon	description	(Fig. 12; Table 5) urn: lsid: zoobank. org: act: CEF 34 BD 6 - BE 69 - 40 A 7 - 99 E 0 - CE 1 B 69 F 860 D 2	en	Dick, Matthew H., Grischenko, Andrei V., Gordon, Dennis P., Ostrovsky, Andrew N. (2021): The “ Cribrilina annulata " problem and new species of Juxtacribrilina (Bryozoa Cheilostomata: Cribrilinidae) from the North Pacific. Zootaxa 5016 (3): 333-364, DOI: 10.11646/zootaxa.5016.3.2
03BCE061A177FFC9B790FD5F5B8CFE85.taxon	diagnosis	Diagnosis. Non-ovicellate zooids with 7 – 14 costae, 2 or 3 oral spines (predominantly 3), lateral pair long, stout, with enlarged bases. Costae inflated, elliptical in transverse section, separated by relatively deep intercostal grooves; suboral pair often thicker than rest; intercostal lacunae moderately large, 5 – 7 in transverse series. Most zooids in basal layer with 1 or 2 frontal pore chambers on proximal gymnocyst. Ovicellate zooids not observed in basal layer. Frontally budded dwarf ovicellate zooids with 4 costae. Ooecium in dwarf zooids cap-like, kenozooidal, bearing single large, circular pseudopore in midline.	en	Dick, Matthew H., Grischenko, Andrei V., Gordon, Dennis P., Ostrovsky, Andrew N. (2021): The “ Cribrilina annulata " problem and new species of Juxtacribrilina (Bryozoa Cheilostomata: Cribrilinidae) from the North Pacific. Zootaxa 5016 (3): 333-364, DOI: 10.11646/zootaxa.5016.3.2
03BCE061A177FFC9B790FD5F5B8CFE85.taxon	etymology	Etymology. The specific name derives from the Latin adjective tumidus (swollen), referring to the swollen appearance of the costae.	en	Dick, Matthew H., Grischenko, Andrei V., Gordon, Dennis P., Ostrovsky, Andrew N. (2021): The “ Cribrilina annulata " problem and new species of Juxtacribrilina (Bryozoa Cheilostomata: Cribrilinidae) from the North Pacific. Zootaxa 5016 (3): 333-364, DOI: 10.11646/zootaxa.5016.3.2
03BCE061A177FFC9B790FD5F5B8CFE85.taxon	materials_examined	Material examined. Holotype: YPM-IZ- 106557, on SEM stub, Eider Island, Narrow Strait, Kodiak, Alaska (57.88560 ° N, 152.40095 ° W), collected by M. H. Dick, December 1982, lower rocky intertidal. Similar material of uncertain identity: ZIRAS 01 / 50737, Ptichy Island, 11 km northwest of Cape Raduzhnyy, western Kamchatka Peninsula, Sea of Okhotsk (57.16361 ° N, 156.56194 ° E), collected by A. V. Grischenko, 6 September 1992, colony detached from inner surface of Mya truncata shell, Balanus zone, middle to lower rocky intertidal, on SEM stub. NHM 2006.2.27.51, Akkeshi Bay, Hokkaido, Japan, small, non-ovicellate colony, collected by A. V. Grischenko and M. H. Dick, rocky intertidal, June – July 2004. YPM-IZ- 106558, on SEM stub, Higgins Point, Ketchikan, Alaska (55.455183 ° N, 131.834305 ° W), collected by M. H. Dick, 9 September 2003, rocky intertidal.	en	Dick, Matthew H., Grischenko, Andrei V., Gordon, Dennis P., Ostrovsky, Andrew N. (2021): The “ Cribrilina annulata " problem and new species of Juxtacribrilina (Bryozoa Cheilostomata: Cribrilinidae) from the North Pacific. Zootaxa 5016 (3): 333-364, DOI: 10.11646/zootaxa.5016.3.2
03BCE061A177FFC9B790FD5F5B8CFE85.taxon	description	Measurements. See Table 5. Description (holotype). Colony (Fig. 12 A) 5 x 2 mm in size; encrusting, multiserial, unilaminar except for frontally budded dwarf zooids. Zooids (Fig. 12 B) closely appressed, delineated by deep groove; oval, long-oval, or spindle-shaped in outline. Gymnocyst negligible except as narrow zone proximally, bearing one or two frontal pore chambers on most zooids; frontal pore chambers occasionally also occurring on lateral gymnocyst. Frontal shield highly convex, with 7 – 14 relatively thick, tumid costae; intercostal lacunae moderately large, circular or irregular, 5 – 7 in transverse series; each costa with lumen pseudopore at or close to tip. Suboral pair of costae usually stouter than rest, sometimes elevated in midline or curving fronto-distally to form raised suboral projection. Secondary orifice approximately semicircular, broader than long, with rounded corners. Non-ovicellate zooids with 2 – 3 articulated oral spines; disto-lateral pair tall, thick, cylindrical, arising from enlarged bases (Fig. 12 B), with or without smaller, single, tapering spine between them. No ovicellate zooids observed in basal layer. Dwarf zooids (Fig. 12 C, D) with 4 costae, including thickened suboral pair. Proximal border of ooecial complex concave, formed by pair of thickened, tapering, modified latero-oral spines meeting at midline, each with small lumen pseudopore near tip. Ooecium reduced, cap-like, kenozooidal, terminal, with single, large pseudopore in center (Fig. 12 C, D). No marginal dwarf zooids observed. No ancestrula observed.	en	Dick, Matthew H., Grischenko, Andrei V., Gordon, Dennis P., Ostrovsky, Andrew N. (2021): The “ Cribrilina annulata " problem and new species of Juxtacribrilina (Bryozoa Cheilostomata: Cribrilinidae) from the North Pacific. Zootaxa 5016 (3): 333-364, DOI: 10.11646/zootaxa.5016.3.2
03BCE061A177FFC9B790FD5F5B8CFE85.taxon	discussion	Remarks. Juxtacribrilina tumida n. sp. differs from J. annulata in the same ways as do J. ezoensis n. sp., J. pushkini n. sp., and J. dobrovolskii n. sp. (see above). It differs from the latter three species in having fewer costae (average, 10.1, Table 5; range of averages for the latter three, 12.8 – 17.1, Tables 2 – 4); fewer intercostal lacunae in transverse series (average, 6.3; range of averages for the other three, 7.1 – 8.6); the suboral pair of costae markedly thicker than the rest (Fig. 12 B); and the disto-lateral pair of oral spines thick, with large bases evident in bleached material (Fig. 12 B). The dwarf zooidal ooecium in J. tumida n. sp. has a single, relatively large, circular pseudopore in the center (Fig. 12 C, D), whereas those in J. ezoensis n. sp. (Fig. 6 C) and J. pushkini n. sp. (Fig. 10 A) typically have two smaller, transversely arranged pseudopores, and that in J. dobrovolskii n. sp. (Fig. 11 D) has a single slitlike, oval, or ramifying pseudopore. A single intertidal specimen from Akkeshi Bay, northern Japan (Fig. 13 A; Table 5) shows non-ovicellate zooids very similar to J. tumida n. sp. and could be this species, although the identification is tentative because no ovicellate zooids were present. A single intertidal specimen from Ptichy Island, Sea of Okhotsk (Fig. 13 B; Table 5), likewise showed non-ovicellate zooids similar to those in J. tumida n. sp., with relatively few costae and intercostal lacunae, a similar complement of oral spines, and enlarged bases of the disto-lateral oral spines, but differed in having flatter, less tumid costae and markedly larger zooid size. As the Ptichy specimen lacked ovicellate zooids, its identity likewise remained ambiguous. Specimen YPM-IZ- 106558 from Ketchikan (Fig. 13 C – F) is unusual in showing a band two generations wide of zooids (both ovicellate and non-ovicellate) similar in morphology to J. tumida n. sp. in having fewer costae and fewer, larger intercostal lacunae (Table 5), and in the somewhat enlarged bases of the disto-lateral oral spines. This zone, however, gives rise to a marginal zone of zooids having the more numerous costae and smaller, more numerous intercostal lacunae characteristic of J. pushkini n. sp. (Fig. 13 D). The costae in the J. tumida - like zooids in specimen YPM-IZ- 106558 are not as inflated as in J. tumida n. sp. In addition, the ooecia in frontal dwarfs are intermediate between the single-pseudopore condition in J. tumida n. sp. and that in J. pushkini n. sp.; in Fig. 13 F, one ooecium has a single large pseudopore, three have two smaller pseudopores transversely arranged but closer together than is typical for J. pushkini n. sp., and two have a dumbbell-shaped opening resulting from the incomplete separation of two close pseudopores. The costal shields in dwarf zooids in YPM-IZ- 106558 are flatter and composed of four costae, as in J. tumida n. sp.; those in J. pushkini n. sp. typically have five or six costae. The nature of the J. tumida - like band in the Ketchikan specimen is not clear. This specimen is not clearly either J. tumida n. sp. or J. pushkini n. sp.; none of the many other specimens from Ketchikan identified as J. pushkini n. sp. showed the mixed zooid types. It is noteworthy that another Juxtacribrilina species, J. mutabilis, shows distinct, alternative zooid types, with individual colonies containing one to three of the types (Ito et al. 2015; Dick et al. 2020), intermingled or occurring in separate zones. One explanation for Ketchikan specimen YPM-IZ- 106558 is that it represents a distinct species showing alternative zooid types. Another possibility is that hybridization has occurred between J. tumida n. sp. and J. pushkini n. sp. at Ketchikan, although no J. tumida n. sp. colonies were detected there. Without more material from across the North Pacific, we cannot resolve this issue. Occurrence. We found J. tumida n. sp. only at Kodiak, although specimens of ambiguous identity from other localities (Ptichy Island, Akkeshi, Ketchikan), if confirmed, would indicate a broader, amphi-Pacific distribution.	en	Dick, Matthew H., Grischenko, Andrei V., Gordon, Dennis P., Ostrovsky, Andrew N. (2021): The “ Cribrilina annulata " problem and new species of Juxtacribrilina (Bryozoa Cheilostomata: Cribrilinidae) from the North Pacific. Zootaxa 5016 (3): 333-364, DOI: 10.11646/zootaxa.5016.3.2
