Two New Species of Encrusting Sponge (Porifera, Family Crellidae) from Eastern Canada

Home , Crella, Crellidae, Heteroscleromorpha

Canadian Journal of Zoology

Two new species of encrusting sponge (Porifera, Family Crellidae) from eastern Canada.

Journal: Canadian Journal of Zoology

Manuscript ID cjz-2021-0041.R1

Manuscript Type: Article

Date Submitted by the 31-Mar-2021 Author:

Complete List of Authors: Goodwin, Claire; Huntsman Marine Science Centre, Aquatic Biosciences; University of New Brunswick Saint John Dinn, Curtis; Fisheries and Oceans Canada Gulf Region, Nefedova, Ekaterina; Zoological Institute RAS Nijhof, Frauke;Draft Huntsman Marine Science Centre Murillo, Francisco; Fisheries and Oceans Canada Maritimes Region, Bedford Institute of Oceanography Nozères, Claude; Fisheries and Oceans Canada, Maurice Lamontagne Institute

Is your manuscript invited for consideration in a Special Not applicable (regular submission) Issue?:

PORIFERA < Taxon, Crella, Crellomima, Bay of Fundy, Keyword: Gulf of St. Lawrence, Scotian Shelf, new demosponges

1 Title: Two new species of encrusting sponge (Porifera, Family Crellidae) from eastern

2 Canada.

3 CLAIRE GOODWIN1,2, CURTIS DINN3, EKATERINA NEFEDOVA4, FRAUKE NIJHOF1,

4 FRANCISCO JAVIER MURILLO5, CLAUDE NOZÈRES6

5 1. Huntsman Marine Science Centre, St. Andrews, New Brunswick, Canada

6 2. University of New Brunswick, Saint John, New Brunswick, Canada

7 3. Fisheries and Oceans Canada, Gulf Fisheries Centre, Moncton, New Brunswick, Canada

8 4. Zoological Institute of the Russian Academy of Sciences, Saint Petersburg, Russia

9 5. Fisheries and Oceans Canada, Bedford Institute of Oceanography, Dartmouth, Nova Scotia, Canada

10 6. Fisheries and Oceans Canada, Maurice Lamontagne Institute, Mont-Joli, Quebec, Canada

11 * corresponding author: [email protected] 12 Draft 13 Competing interests: The authors declare there are no competing interests.

14 Contributors statement: CG, CD, and FJM co-ordinated collections, and managed specimen data 15 curation. All authors contributed to taxonomic investigations. CG and CD wrote the original draft of the 16 manuscript. All authors were involved in review and editing of the manuscript. 17 18 Data availability statement: Specimens and specimen collection data is publicly available through the 19 museum specified. 20 21 Funding statement: 22 The Brier Island and Gulf of St. Lawrence taxonomic work was funded by Fisheries and Oceans Canada 23 under Marine Conservation Targets programme supervised by Peter Lawton and J. Andrew Cooper (Brier 24 Island) as well as Denise Méthé and Geneviève Faille (Gulf of St Lawrence). The Gulf of St. Lawrence 25 surveys are part of regular activities conducted and funded by Fisheries and Oceans Canada. Sample 26 collection from the Scotian Shelf and taxonomic work on specimens was made possible by Fisheries and 27 Oceans Canada Strategic Program for Ecosystem-Based Research and Advice (SPERA) funding awarded 28 to Ellen Kenchington and Melisa Wong. SCUBA collection in the Bay of Fundy and taxonomic work on 29 these specimens was supported by an award to Huntsman Marine Science Centre from the New 30 Brunswick Government Total Development Fund.

1 © The Author(s) or their Institution(s) Canadian Journal of Zoology Page 2 of 39

32 Acknowledgements

33 Access to type material was facilitated by Helma Roggenbuck and Andreas Schmidt-Rhaesa (Zoological

34 Museum of the University of Hamburg); Emma Sherlock and Jasmin Perera (British Museum of Natural

35 History) and the Zoological Institute of the Russian Academy of Sciences. Thanks to L.A. Kartseva from

36 the Center for collective use of Komarov Botanical Institute of the Russian Academy of Sciences, Nhu

37 Trieu of the University of New Brunswick (Fredericton) and James M. Ehrman at the Digital Microscopy

38 Facility, Mount Allison University for assistance with SEM imaging. Thanks to J. Andrew Cooper, Peter

39 Lawton and Torben Brydges for their roles in organizing the Brier Island Survey. Thanks to Pat

40 Fitzgerald and Connie Bishop for their assistance with specimen collection in the Bay of Fundy. The

41 authors thank the CCGS Teleost, CGGS Alfred Needler, and RV Fundy Spray captains and crews.

42 Thanks to Mylène Roussel for microscope slideDraft preparation and Denise Méthé for sampling supervision.

43 We also thank Geneviève Côté, and Geneviève Faille from DFO Quebec for providing samples and

44 photos.

2 © The Author(s) or their Institution(s) Page 3 of 39 Canadian Journal of Zoology

47 Abstract

48 Two new species of Crellidae Dendy, 1922 from the east coast of Canada are described. The first is

49 Crella (Pytheas) cutis sp. nov., a massively encrusting species of Crella (Pytheas) collected from depths

50 of 84 to 249 m in the Gulf of St Lawrence and on the Scotian Shelf. The second is Crellomima

51 mehqisinpekonuta sp. nov., a thinly encrusting sponge found at diving depths in the Bay of Fundy. We

52 also report the first records of Crellomima derma Hentschel, 1929 from outside the type locality (Barents

53 Sea). All known species of Crellomima are reviewed based on type material.

55 Key words:

56 Porifera, Crella, Crellomima, Bay of Fundy, Gulf of St. Lawrence, Scotian Shelf, new demosponges,

57 Northwest Atlantic Ocean

58 Draft

59 Introduction

60 Sponges of the family Crellidae Dendy, 1922, a likely polyphyletic group (Redmond et al. 2013; Thacker

61 et al. 2013), consist of sponges with diverse growth forms that are often found in cold and deep water

62 (Van Soest 2002). The family Crellidae consists of five genera characterized by a tangential crust of

63 acanthostyles or acanthoxeas and areolated pore fields constructed of parallel spicules (Van Soest 2002).

64 Members of the family may have arcuate isochelae, occasionally anisochelae, and rarely sigmas as

65 microscleres (Van Soest 2002).

66 The genera Crella and Crellomima are distinguished from the other members of the family as

67 they do not have reduced anisochelae (Anisocrella, one species) or asterose acanthoxeas (Crellastrina,

68 Spirorhabdia, one and two species respectively). Crella species usually have arcuate chelae with rounded

69 alae, while Crellomima chelae are polydentate (Van Soest 2002). Crella is the most species rich genus

70 with 63 species (Van Soest et al. 2021) and contains four subgenera (Crella, Grayella, Pytheas, and

71 Yvesia), though these groups were erected for convenience and do not necessarily comprise phylogenetic

72 groupings (Van Soest 2002). Crellomima contains three named species (Van Soest et al. 2021). Only

3 © The Author(s) or their Institution(s) Canadian Journal of Zoology Page 4 of 39

73 three members of the family have been identified so far from eastern Canada: Crella (Yvesia) pyrula

74 (Carter 1876), Crella (Yvesia) guernei (Topsent 1890), and Crella (Yvesia) rosea (Topsent, 1892)

75 (Topsent 1892; Goodwin 2017; Dinn and Leys 2018).

76 Collections in the Gulf of St. Lawrence, Scotian Shelf, and the Bay of Fundy along Canada’s

77 eastern coast have revealed two new species of encrusting sponges in the family Crellidae. Here we

78 describe a new massively encrusting species of Crella (Pytheas) collected in deeper water and a new

79 species of a thin crust forming Crellomima found at diving depths in the Bay of Fundy. Other members of

80 Crellomima are also reviewed based on type material.

82 Materials and methods

83 Specimens were collected between 2017 and 2019 in three different regions of eastern Canada: the Gulf

84 of St. Lawrence, the Scotian Shelf and the bayDraft of Fundy (Figure 1). Gulf of St. Lawrence specimens were

85 collected during two surveys: the 2018 ecosystemic survey in the Estuary and northern Gulf of St.

86 Lawrence aboard the CCGS Teleost (August 4–September 1, 2018) using a Campelen 1800 trawl towed

87 for 15 minutes at 3 knots (Bourdages et al. 2020), and the 2019 multispecies survey in the southern Gulf

88 of St. Lawrence aboard the CCGS Teleost (September 5–October 2, 2019) using a Western IIA trawl

89 towed for 30 minutes at 3.5 knots (Hurlbut and Clay 1990) (Figure 1 A). Scotian Shelf specimens were

90 collected during the 2017 summer multispecies surveys aboard the CGGS Alfred Needler (June 28–

91 August 14, 2017) also collected by Western IIA trawl towed for 30 minutes at 3.5 knots (Murillo et al.

92 2018b) (Figure 1 A). On all cruises, sponge specimens were photographed on board with a scale for size

93 reference and then either preserved in 95% ethanol or frozen.

94 Sponges in the Bay of Fundy were collected using SCUBA as part of a study on Bay of Fundy

95 sponges (Goodwin 2017) and a survey surrounding Brier Island Nova Scotia (Cooper et al. 2019) (Figure

96 1 B-C). Sponges were photographed in situ using a Nikon D300 with 60 mm macro lens in a Subal

97 Housing with an Ikelite DS125 strobe. A small piece of each sponge was then taken by hand using a knife

98 and placed in a numbered sample bag. Specimens were preserved in 95% ethanol.

4 © The Author(s) or their Institution(s) Page 5 of 39 Canadian Journal of Zoology

99 Tissue sections were prepared by sectioning very thin portions of tissue dehydrated in ethanol.

100 Sections were clarified in clove oil for several minutes then mounted on a microscope slide in Canada

101 balsam. Tissue sections were imaged using a stereomicroscope or compound microscope. To isolate

102 spicules, pieces of sponge were placed in undiluted household bleach overnight to remove tissue, then

103 rinsed four times in distilled water and cleaned in two washes of 95% ethanol. Spicules were allowed to

104 settle for at least 15 minutes between rinses and then the upper layer of liquid was pipetted off, leaving

105 the spicules undisturbed. Cleaned spicules were dried on glass slides, mounted in DPX mounting medium

106 (Sigma-Aldrich, St. Louis, MO) or Canada Balsam and imaged using a compound microscope. For

107 scanning electron microscopy (SEM), cleaned spicules were placed on metal stubs, coated with gold and

108 viewed with a Hitachi SU3500 SEM (Mount Allison University, Sackville) or JEOL 6400 SEM (UNB,

109 Fredericton). Spicule measurements (n = 20–30, unless otherwise noted) were made with ImageJ 1.52 or

110 using an Olympus SC50 camera and OlympusDraft cellSens standard 1.16 software. Measurements are

111 reported as minimum(mean)maximum. Specimens were deposited at the Atlantic Reference Centre

112 (ARC), St. Andrews, New Brunswick. The World Porifera Database, which implements the classification

113 system for Demospongiae proposed by Morrow and Cárdenas (Morrow and Cárdenas 2015), was used as

114 the taxonomic authority (Van Soest et al. 2021). Type material was examined from the collections of

115 Zoological Institute of the Russian Academy of Sciences (ZIN); British Museum of Natural History

116 (BMNH) and Zoological Museum of the University of Hamburg (ZMH).

117 Total DNA was extracted using the DNeasy Blood and Tissue kit (Qiagen, Valencia, CA)

118 following the manufacturer’s instructions, using approximately 0.2 cm3 of sponge tissue to account for

119 spicule weight. The 5’ end region of COI was amplified using M13F-tailed dgLCO1490 and M13R-tailed

120 dgHCO2198 degenerate Folmer fragment primers (Folmer et al. 1994; Meyer 2003). PCR reactions used

121 Platinum Taq Polymerase (Invitrogen, Carlsbad, CA) and were run using a standard protocol of 94.0°C

122 for 5 min; (94.0°C for 30 s; 50.0°C for 30 s; 72.0°C for 45 s) × 30 cycles; 72.0°C for 2 min. Cleanup and

123 sequencing of PCR products were performed by Génome Québec (Montréal, QC) using an Applied

5 © The Author(s) or their Institution(s) Canadian Journal of Zoology Page 6 of 39

124 Biosystem's 3730xl DNA Analyzer. Consensus sequences were assembled in BioEdit version 7.2 and

125 manually checked for sequencing errors. Sequences were submitted to GenBank (Clark et al. 2016).

126

127 Ethics approval: Specimen collection was conducted in accordance with applicable laws, guidelines and

128 regulations. The majority of specimens were collected under the auspices of Fisheries and Oceans

129 Canada. Fieldwork conducted by Huntsman in the Bay of Fundy was covered by scientific collection

130 permits issued by Fisheries and Oceans Canada.

131

132 Results

133 Taxonomy

134 Phylum Porifera Grant, 1836

135 Class DemospongiaeDraft Sollas, 1885

136 Subclass Heteroscleromorpha Cárdenas, Pérez & Boury-Esnault, 2012

137 Order Poecilosclerida Topsent, 1928

138 Family Crellidae Dendy, 1922

139 Genus Crella Gray, 1867

140 Subgenus Crella (Pytheas) Topsent, 1890

141

142 Crella (Pytheas) cutis sp. nov.

143 (Figure 2, Table 1)

144

145 Zoobank no: urn:lsid:zoobank.org:act:4E29F07E-284F-480D-B124-54C03D1C6407

146

147 Diagnosis

148 Massively encrusting Crella with leathery skin, mostly smooth choanosomal acanthostyles, with chelae

149 and sigma microscleres.

6 © The Author(s) or their Institution(s) Page 7 of 39 Canadian Journal of Zoology

150 Material examined

151 Holotype:

152 ARC 81601 Northwest of St. Paul Island, Nova Scotia (47.2825 N, 60.3025 W), 205 m depth. DFO

153 CCGS Teleost southern Gulf of St. Lawrence multispecies survey, 1 October, 2019.

154 Paratypes:

155 ARC 81602 Northern Gulf of St. Lawrence, Esquiman Channel, (50.301 N, 58.704 W), 150 m depth.

156 DFO CCGS Teleost ecosystemic survey in the Estuary and northern Gulf of St. Lawrence, 22 August,

157 2018.

158 ARC 81603 Northern Gulf of St. Lawrence, Esquiman Channel (50.1685 N, 58.9112 W), 203 m depth.

159 DFO CCGS Teleost ecosystemic survey in the Estuary and northern Gulf of St. Lawrence, 22 August,

160 2018.

161 ARC 81500 Scotian Shelf (43.4485°N, 65.5166°W),Draft 84 m depth. DFO Maritimes CCGS Alfred Needler

162 multispecies survey, 6 July, 2017.

163

164 External appearance (Figure 2 A-B)

165 Massively encrusting species which has a distinctive leathery skin with many small conical papillae. The

166 underlying body is a light cream colour but darkens to grey or black after prolonged air exposure. The

167 sponge is often coated in thick, gelatinous mucous which persists after preservation in ethanol. The

168 ectosome of preserved specimens is detachable as a very thin, translucent skin. Specimens were not seen

169 in situ.

170

171 Spicules (Figure 2 C-O)

172 Choanosomal acanthostyles 427(478)539 x 13(17)19 μm. Large, mostly smooth, and have a mucronate

173 head, sometimes with multiple spines near the rounded head or scattered along the shaft.

174 Tylotes 360(389)413 x 8.9(10)12 μm. Fusiform and strongyle-like with minimal swelling at the ends.

175 Large ectosomal acanthostyles 294(331)384 x 11(12)14 μm.

7 © The Author(s) or their Institution(s) Canadian Journal of Zoology Page 8 of 39

176 Choanosomal echinating acanthostyles 99(132)192 x 7.0(9.6)11.5 μm. 177 Arcuate isochelae 21(24)27 μm (uncommon). Possess three or more pointed alae, often with a longer 178 centre ala. 179 Sigmas 32(36)39 μm. C-shaped, narrow abruptly near the pointed end. 180

181 Skeleton (Figure 2 P-Q)

182 The ectosomal skeleton consists of a multilayered and dense skin of long acanthostyles, with tangential

183 tylotes supporting the papillae. The skin can be several hundred microns thick. The chonaosomal skeleton

184 is formed by plumose bundles of mostly smooth acanthostyles with sigmas and isochelae scattered

185 throughout with no discernable pattern. The spicule bundles ascend towards the ectosomal skin, but

186 meander and converge. Echinating acanthostyles are present between the spicule bundles and are common

187 in the basal skeleton near the substrate, especially obvious near embedded sand grains. There is often a

188 wide subdermal space between the ectosomalDraft skin and choanosome, but this may be a result of damage

189 during collection.

190

191

192 Genetic data

193 COI mtDNA sequences were obtained for ARC 81603 (GenBank MW176031). Few Crella COI 194 sequences have been published; however, the obtained sequence does not appear similar to any other 195 Crellidae sequences submitted to Genbank. The sequence appears most similar to species of Tedania, 196 though not closely related. The megasclere composition of members of Tedania are similar to those of 197 Crella (Pytheas) cutis sp. nov. however, chelae and sigmas are not present in members of the Tedaniidae 198 and onychaete spicules are not usually present outside of the Tedaniidae (Hooper and Van Soest 2002). 199 Further genetic analysis can be performed when additional sequences of related taxa become available 200 from the North Atlantic, or from analyzing a different gene. 201

202 Distribution and ecology

203 Encountered often during trawl surveys in the Northern Gulf of St. Lawrence from 100–250 m depth.

204 Holotype from the southern edge of the Laurentian Channel, near St. Paul Island. Collected several times

8 © The Author(s) or their Institution(s) Page 9 of 39 Canadian Journal of Zoology

205 from the Esquiman Channel up to the shelf west of Newfoundland. Found encrusting pebbles and often

206 with many embedded sand grains. Also common on Scotian Shelf where it was collected from 84–225 m

207

208 Etymology

209 From the Latin cutis meaning skin, membrane, or leathery hide. The species has a very obvious

210 detachable skin that makes it distinct from other sponges in the region.

211

212 Remarks

213 The spiculation consisting of mostly smooth acanthostyles and common sigma microscleres is not

214 congruent with any other Crella specimens and is unique for the genus. Crellinspira Laubenfels, 1936;

215 Naniupi de Laubenfels, 1950; Ramosichela de Laubenfels, 1950 are considered synonyms of Crella

216 (Pytheas)(Van Soest et al. 2021). De LaubenfelsDraft separated the three sub-genera on the basis of the form

217 of their megascleres with Ramosichela having acanthostyles as both the primary and echinating spicules,

218 Naniupi with primary smooth styles and echinating acanthostyles and Crellinspira lacking echinating

219 acanthostyles. Crella (Pytheas) as currently defined includes all Crella with isochelae and basal or

220 echinating acanthostyles. A Phylogenetic revision of the genus might reveal if megasclere differences

221 have any significance.

222

223 Crella (Pytheas) cutis sp. nov. is easily identified in the field due to the presence of a distinctive and

224 easily removed skin, cream to black colouration, and copious mucous production. The ectosomal

225 acanthostyles of Crella (Pytheas) cutis sp. nov. are much longer than the echinating/basal acanthostyles, a

226 trait consistent only with C. (P.) affinis, C. (P.) alba, and C. (P.) altra, while other members of the

227 subgenus have echinating/basal acanthostyles which are longer than the ectosomal acanthostyles. C. (P.)

228 alba differs in lacking choanosomal acanthostyles and sigmas (Table 1). External morphology is similar

229 to C. (P.) atra, which is described as a black encrusting sponge that is soft with an easily detachable

230 smooth skin (Topsent 1892; Van Soest 2002), though the holotype of the species was said to be growing

9 © The Author(s) or their Institution(s) Canadian Journal of Zoology Page 10 of 39

231 on top of an arborescent coral (Topsent 1890, 1892). The echinating spicules of C. (P.) atra however are

232 the same size as its ectosomal acanthostyles but are partially smooth, and the species does not have

233 sigmas.

234 Sigma microscleres have only previously been reported in two species of Crella (Table 1). C.

235 (P.) sigmata (Topsent, 1925) is reported to have two sizes of sigma (Boury-Esnault and Lopes 1985),

236 which are generally larger than the sigmas in Crella (Pytheas) cutis sp. nov. Sigmas are also present in

237 Crella (Yvesia) topsenti (Babiç 1922) however that species has acanthoxeas rather than acanthostyles and

238 no echinating acanthostyles. Sigmas were also seen in C. (P.) donsi (Burton, 1931), but were considered

239 foreign (Van Soest 2002).

240

241 Genus Crellomima Rezvoi, 1925

242 Draft

243 Crellomima mehqisinpekonuta sp. nov.

244 (Figure 3, Table 2)

245

246 Zoobank no: urn:lsid:zoobank.org:act:8538E0A2-CB82-49D3-AC8B-6B0C0264CDB7

247

248 Diagnosis

249 Crellomima with polydentate chelae that have 5–8 alae and ectosomal spicules comprising solely of

250 acanthostyles which are comparatively long for the genus (120–188 µm).

251

252 Material examined

253 Holotype: ARC 80713, Can Cat Beach, Deer Island, New Brunswick, (44.92917°N, 66.98715°W), 6–12

254 m depth, SCUBA, Collected by Claire Goodwin, 6 November, 2016.

255 Paratypes: ARC 80709, ARC 80712 Can Cat Beach, Deer Island, New Brunswick, (44.92917°N,

256 66.98715°W), 6–12 m depth, SCUBA, Collected by Claire Goodwin, 6 November, 2016.

10 © The Author(s) or their Institution(s) Page 11 of 39 Canadian Journal of Zoology

257 Other specimens: ARC80705, Can Cat Beach, Deer Island, New Brunswick, (44.92917°N, 66.98715°W),

258 6–12 m depth, SCUBA, Collected by Claire Goodwin, 6 November, 2016.

259 ARC 81430 and ARC 81433 Peter’s Island, Brier Island, Nova Scotia (44.25860°N, 66.33667), 10 m

260 depth, SCUBA, collected by J. Andrew Cooper and Claire Goodwin, 27 August, 2017.

261 ARC 81520, Grand Passage, Brier Island, Nova Scotia, (44.27916°N, 66.34076°W), 17 m depth,

262 SCUBA, collected by Claire Goodwin and Torben Brydges, 27 September, 2017.

263 ARC 81528, ARC 81530 and ARC 81532 North end of Peter's Island, Nova Scotia, (44.26084°N,

264 66.33835°W), 13 m depth, collected by Andrew Cooper and Claire Goodwin, 27 September, 2017

265 ARC 81541 South-east shore Brier Island (near Brier 11), Nova Scotia, (44.24457°N, 66.35309°W), 21 m

266 depth, SCUBA, collected by Andrew Cooper and Claire Goodwin, 28 September, 2017.

267 ARC 81556 and ARC 81557 Brier Island SE shore near Brier 10, Nova Scotia, (44.24997°N,

268 66.34645°W), depth 20 m depth, collected ClaireDraft Goodwin and Torben Brydges, 28 September, 2017.

269

270 Comparative material examined: Syntype Crellomima imparidens Rezvoi, 1925 ZIN 7011. Holotype

271 Crellomima incrustans Hentschel, 1929 BMNH 1936.11.20.14; Holotype Crellomima derma Hentschel,

272 1929 ZMH S3259.

273

274 External appearance (Figure 3 A)

275 Bright orange thinly encrusting sponge. Irregularly spaced oscules are surrounded by stellate channels.

276 Between these the entire surface is covered with small punctate pore sieves (<0.5mm in diameter).

277

278 Skeleton (Figure 3 B)

279 A layer of erect basal acanthostyles, heads down, echinate the substrate. Ascending columns of 15–20

280 tornotes fan out towards the surface, supporting a dense ectosomal crust of tangential acanthostyles.

281 Chelae are scattered very sparsely throughout the tissue.

282

11 © The Author(s) or their Institution(s) Canadian Journal of Zoology Page 12 of 39

283 Spicules (Figure 3 C-I).

284 Measurements are from the holotype ARC 80713, measurements from additional specimens in Table 2.

285 Basal acanthostyles 152(166)192 x 8(14)18 µm, in one size category. Heads not tylote. Spined with large

286 spines, the smaller individuals are entirely spined, but the larger may be smooth or more sparsely spined

287 at the tip.

288 Ectosomal acanthostyles 120(138)188 x 5(5)6µm. Thin acanthostyles with an abrupt point. Entirely

289 spined with large spines.

290 Tornotes 185(204)218 x 4(6)7 µm. Fusiform tornotes with uneven ends.

291 Chelae 18(22)25µm with 5–8 alae, the alae continue down the shaft, diminishing in size.

292

293 Genetic data

294 Although sequencing of CO1 and the D2–D5Draft region of 28S was attempted, neither were successful.

295

296 Distribution and ecology

297 Currently known from the type locality Deer Island, New Brunswick in the Quoddy region of the Bay of

298 Fundy, and Brier Island, Nova Scotia on the outer edge of the Bay of Fundy. Specimens were collected

299 from between 6 and 21 m in depth. Forms thin (< 0.5 mm thick) crusts on bedrock. As the sponge fauna,

300 and particularly encrusting sponge fauna, of eastern Canada is poorly known, this species is likely to be

301 more widely distributed.

302

303 Etymology

304 From the Peskotomuhkati ‘Mehqi-sinpekonut’ meaning something reddish orange (animate) that gets

305 water squeezed out of it. The name was provided by the Peskotomuhkati language committee. The type

306 locality is in the traditional territory of the Peskotomuhkati nation.

307

308 Remarks

12 © The Author(s) or their Institution(s) Page 13 of 39 Canadian Journal of Zoology

309 Crellomima is defined as a Crellidae with anchorate chelae (Rezvoi 1925). An anchorate chela is defined

310 as ‘An isochela with three or more free alae (at each end) in the form of recurved processes shaped like

311 anchor claws or anchor blades; with two incipient lateral alae fused with the shaft over their entire length

312 and a gently curved, not abruptly arched shaft’ (Boury-Esnault and Rützler 1997). This differs from other

313 species in the family Crellidae which have arcuate chelae, defined as ‘An isochela with three free alae and

314 the shaft characteristically curved outward, often bow-shaped’ (Boury-Esnault and Rützler 1997).

315 There are currently three described species of Crellomima, all from the Arctic: Crellomima derma

316 Hetschel, 1929 (type locality Svalbard; Figure 5), Crellomima incrustans Hentshel, 1929, (type locality

317 Spitzbergen; Figure 6), and Crellomima imparidens Rezvoi, 1925 (type locality Russian Arctic; Figure 7)

318 (see Table 2). C. imparidens Rezvoi, 1925 is the type species of the genus. Re-examination of the type

319 specimens shows that all three do indeed have anchorate chelae, although of quite different forms. As

320 stated in Van Soest (2002), the possession ofDraft anchorate chelae is an exception in the Crellidae and the

321 value of chelae micromorphology for defining this group requires further investigation. Chelae

322 morphology can evolve independently in different poecilosclerid lineages so may be misleading as an

323 indicator of the phylogenetic history of the group (Vargas et al. 2011; Morrow and Cárdenas

324 2015). Molecular studies on this group may reveal better characters for delineation of genera.

325 Crellomima mehqisinpekonuta sp. nov. is distinct from the currently described species in the

326 genus. Crellomima imparidens can be distinguished as it has anchorate chelae with only 3–4 alae and

327 polytylote tornotes. C. incrustans has polydentate chelae but can be distinguished by its much shorter

328 dermal spicules (113–132 x 6-–7µm), longer tornotes (210–250 x 4.5–5 µm), and the presence of

329 ectosomal acanthostrongyles in addition to acanthostyles. The form of the chelae in C. incrustans is also

330 different as they have alae uniform in size and straight shafts. Crellomima derma has much shorter dermal

331 acanthostyles (96–168 x 5–7 µm) and smaller chelae (17–19 µm) which only have three to five alae at

332 each end.

333

334 Crellomima derma Hentschel, 1929

13 © The Author(s) or their Institution(s) Canadian Journal of Zoology Page 14 of 39

335 (Figure 4, Table 2)

336

337 Specimens:

338 ARC 80736; Joe's Point, St. Andrews, New Brunswick (45.07518°N, 67.08552°W), 8 m depth, SCUBA,

339 Collected by Claire Goodwin, 11 November, 2016.

340 ARC80747; McCann Head, Chamcook, New Brunswick 45.12430°N, 67.03565°W), 4–13 m depth,

341 SCUBA, Collected by Claire Goodwin, 7 December, 2016.

342

343 Comparative material examined: Syntype Crellomima imparidens Rezvoi, 1925 ZIN 7011. Holotype

344 Crellomima incrustans Hentschel, 1929 BMNH 1936.11.20.14; Holotype Crellomima derma Hentschel,

345 1929 ZMH S3259.

346 Draft

347 External appearance (Figure 4 A)

348 Very thinly encrusting orange sponge with a slightly hispid surface. No obvious oscules or channels

349 visible.

350

351 Skeleton (Figure 4 B)

352 A layer of erect basal acanthostyles, heads down, echinate the substrate. Ascending columns of 15–30

353 tornotes fan out towards the surface, supporting a dense ectosomal crust of tangential acanthostyles. Rare

354 chelae are present throughout the tissue.

355

356 Spicules (Figure 4 C-H). Measurements from ARC80736, see Table 2 for other specimens.

357 Basal acanthostyles 151(191)228 x 10(13)16 µm. Slightly tylote, densely spined head, spines become

358 progressively sparser towards the tip.

359 Ectosomal acanthostyles 134(149)165 x 3(5)7 µm. Often slightly curved. Densely spined. Abrupt point.

360 Tornotes 199(224)246 x 5(7)9 µm. Slightly fusiform. Asymmetrical.

14 © The Author(s) or their Institution(s) Page 15 of 39 Canadian Journal of Zoology

361 Chelae 16–18 µm, rare. Polydentate anchorate chelae with 3 teeth on the lower half and up to seven on

362 the upper.

363

364 Genetic data

365 Although sequencing of CO1 and the D2–D5 region of 28S was attempted, neither were successful.

366

367 Distribution and ecology

368 The type location of this species is the Barents Sea. Our records extend the range of this species to the

369 Bay of Fundy.

370

371 Remarks

372 The specimens from the Bay of Fundy do differDraft slightly in spicule size from the type specimen. They

373 have longer, thinner, basal acanthostyles and ectosomal acanthostyles (Table 2). The chelae are similar to

374 the type in possessing 3–5 teeth, but they seem to have a slightly more flattened shaft. However, the

375 overall form and size range of the spicules is similar, and therefore we assign these specimens to C.

376 derma. This species has so far only been recorded from the type locality in the Barents Sea. This disjunct

377 distribution is likely to be due to the under-recording of encrusting sponges. The Bay of Fundy sponge

378 fauna has many species that have a boreo-arctic distribution (Goodwin 2017), so the presence of this

379 boreal-arctic species in the Bay of Fundy is certainly possible from a biogeographic perspective.

380

381 Discussion

382

383 The sponge fauna of eastern Canada is rather poorly known compared to other regions in the North

384 Atlantic, with limited previous studies (Lambe 1896, 1900a, 1900b; Whiteaves 1901; Old 1941; Ginn et

385 al. 1998; Fuller 2011). Recent recognition of the ecological importance of sponges (Hogg et al. 2010;

386 Beazley et al. 2013; Maldonado et al. 2015) has prompted a resurgence of interest in the phylum across

15 © The Author(s) or their Institution(s) Canadian Journal of Zoology Page 16 of 39

387 the Atlantic, including eastern Atlantic Canada and the eastern Canadian Arctic (Kenchington et al. 2016;

388 Murillo et al. 2016b, 2018a; Bouchard Marmen et al. 2019). While studies have primarily focused on

389 habitat mapping and ecological associations, there has also been some effort into resolving taxonomic

390 issues (Fuller 2011; Tompkins et al. 2017; Baker et al. 2018; Bouchard Marmen et al. 2019). As a result

391 of these studies, new sponge species from eastern Canada have recently been described (e.g. Hestetun et

392 al. 2017; Dinn 2020), although more await description (Goodwin 2017). Further review may also help to

393 validate survey records that may have been misreported taxa, as has occurred for fan-like sponges (Dinn

394 et al. 2020). Despite this, it’s estimated that half of Canadian Porifera species remain unreported

395 (Mosquin et al. 1995). While several reported groups require review, sponges of family Crellidae do not

396 appear in historical lists for regions such as the Gulf of St. Lawrence (Brunel et al. 1998).

397

398 In this paper, we describe two speciesDraft new to science and report one new record for the region.

399 Eastern Canada has a cold temperate fauna (Archambault et al. 2010) and the majority of sponge species

400 found here have a wide, sometimes circumarctic distribution (Van Soest et al. 2021). However, many

401 sponges are known to have restricted dispersal potential (Maldonado 2006), and thus there may be several

402 endemic species in the region. Additionally, it is thought that worldwide only around half of the extant

403 sponge species have been described (Van Soest et al. 2012), so even those present in other areas may not

404 have scientific names. As in other regions such as the NE Atlantic (Picton and Goodwin 2007; Van Soest

405 et al. 2012), encrusting sponges are particularly poorly known. Small and encrusting sponges are not well

406 sampled in bottom trawl surveys (Wassenberg et al. 2002), although trawls remain a major source of

407 sponge distribution information in Canada (Kenchington et al. 2016; Murillo et al. 2016a). Dedicated

408 sampling of encrusting species and habitats through other means such as SCUBA, as done here for

409 Crellomima, or remotely-operated vehicles has the potential to enable the discovery of many species new

410 to science (Ginn et al. 1998; Vacelet and Perez 1998; Picton and Goodwin 2007; Goodwin et al. 2019).

411 Investing in additional collection and taxonomic study of sponges in Canadian waters will help to better

412 document the biodiversity of this ecologically important group.

16 © The Author(s) or their Institution(s) Page 17 of 39 Canadian Journal of Zoology

413

414 Figure Captions

415 Figure 1. Specimen collection locations. A, Eastern Canada showing sampling locations in the Quoddy

416 region of the Bay of Fundy (b) and Brier Island (c). B, St. Andrews and Deer Island SCUBA locations. C,

417 Brier Island SCUBA locations in southwest Nova Scotia. Circles, Crella (Pytheas) cutis sp. nov.

418 Triangles, Crellomima mehqisinpekonuta sp. nov. Squares, Crellomima derma. Figure was created using

419 ArcMap 10.7.1 and assembled from the following data sources: Collection data sourced from DFO and

420 CG. Base map from GSHHG (Wessel & Smith, 1996), and contour lines from NOAA National

421 Geophysical Data Center ETOPO1 (Amante & Eakins, 2009).

422 Figure 1. Crella (Pytheas) cutis sp. nov. A,B, collected specimens. C,D, choanaosomal acanthostyles.

423 E,F, tylotes. G,H, ectosomal acanthostyles. I,J, echinating/basal acanthostyles. K-M, isochelae. N,O,

424 sigmas. P,Q, skeleton. Draft

425 Figure 2. Crellomima mehqisinpekonuta sp. nov. A, Holotype in situ. B, skeleton. C,D basal acanthostyle

426 (two are illustrated to show variation in form but only one size class is present). E, ectosomal

427 acanthostyle. F,G, tornotes. H,I, isochelae.

428 Figure 3. Crellomima derma, Bay of Fundy Specimen. A, specimen in situ. B, skeleton. C, basal

429 acanthostyle. D, ectosomal acanthostyle. E,F, tornotes. G,H, isochelae.

430 Figure 4. Crellomima derma ZMH S3259. A, basal acanthostyle. B, ectosomal acanthostyle. C,D,

431 tornotes. E,F, isochelae.

432 Figure 5. Crellomima incrustans BMNH 1936.11.20.14. A,B, basal acanthostyle. C, ectosomal

433 acanthostyle. D,E, tornotes. F,G, isochelae.

434 Figure 6. Crellomima imparidens ZIN 7011. A, basal acanthostyle. B, ectosomal acanthostyle. C,D,

435 tornotes. E,F, isochelae.

436

17 © The Author(s) or their Institution(s) Canadian Journal of Zoology Page 18 of 39

437 References

438 Alander, H. 1937. Sponges from the Trondheimsfjord and AdjacentWaters. Part II. Det K. Nor. Vidensk.

439 Selsk. Forh. 10(19): 71-74.

440 Alander, H. 1942. Sponges from the Swedish west-coast and adjacent waters. . Ph.D. Thesis. (University

441 Lund, H. Struves Gøteborg). Available from

442 http://www.marinespecies.org/porifera/porifera.php?p=sourcedetails&id=7036.

443 Amante, C., and B.W. Eakins. 2009. ETOPO1 1 Arc-Minute Global Relief Model: Procedures, Data

444 Sources and Analysis. NOAA Technical Memorandum NESDIS NGDC-24. National Geophysical

445 Data Center, NOAA. doi:10.7289/V5C8276M [accessed 20 January 2021].

446 Archambault, P., Snelgrove, P.V.R., Fisher, J.A.D., Gagnon, J.M., Garbary, D.J., Harvey, M.,et al. . 2010.

447 From sea to sea: Canada’s three oceans of biodiversity. PLoS One 5(8): e12182.

448 doi:10.1371/journal.pone.0012182. Draft

449 Arndt, W. 1913. Zoologische Ergebnisse der ersten Lehr-Expedition der Dr. P. Schottländerschen

450 Jubiläums-Stiftung. Jahresbericht der Schlesischen gesellschaft für vaterländische Kult. 90(1): 110–

451 136.

452 Arnesen, E. 1903. Spongien von der norwegischen Küste. II. Monaxonida: Halichondrina. Bergen.

453 Museum Åarb.

454 Babiç, K. 1922. Monactinellida und Tetractinellida des Adriatischen Meeres. Zool. Jahrbücher. Abteilung

455 für Syst. Geogr. und Biol. der Tiere.

456 Baker, E., Odenthal, B., Tompkins-Macdonald, G., Walkusz, W., Siferd, T., and Kenchington, E.L.R.

457 2018. Sponges from the 2010-2014 Paamiut Multispecies Trawl Surveys, Eastern Arctic and

458 Subarctic: Class Demospongiae, Subclass Heteroscleromorpha, Order Poecilosclerida, Families

459 Crellidae and Myxillidae. Can. Tech. Rep. Fish. Aquat. Sci. 3253: 1–52.

460 Beazley, L.I., Kenchington, E.L., Murillo, F.J., and del Mar Sacau, M. 2013. Deep-sea sponge grounds

461 enhance diversity and abundance of epibenthic megafauna in the Northwest Atlantic. ICES J. Mar.

462 Sci. 70(7): 1471–1490. doi:10.1093/icesjms/fst124.

18 © The Author(s) or their Institution(s) Page 19 of 39 Canadian Journal of Zoology

463 Bouchard Marmen, M., Tompkins, G., Harrington, N., Savard-Drouin, A., Wells, M., Baker, E.,et al. .

464 2019. Sponges from the 2010-2014 Paamiut Multispecies Trawl Surveys, Eastern Arctic and

465 Subarctic: Class Demospongiae, Subclass Heteroscleromorpha, Order Poecilosclerida, Families

466 Microcionidae, Acarnidae and Esperiopsidae. Can. Tech. Rep. Fish. Aquat. Sci. 3349 3349: 1–76.

467 Bourdages, H., Brassard, C., Desgagnes, M., Galbraith, P., Gauthier, J., Nozères, C., Scallon-Chouinard,

468 P.-M., and Senay, C. 2020. Preliminary results from the ecosystemic survey in August 2019 in the

469 Estuary and northern Gulf of St. Lawrence. DFO Can. Sci. Advis. Sec. Res. Doc. 2020/009. iv + 93

470 p.

471 Boury-Esnault, N., and Lopes, M.T. 1985. Les Démosponges de la zone littorale de l’Archipel des

472 Açores. Ann. l’Institut océanographique 61(2): 149–225.

473 Boury-Esnault, N., and Rützler, K. 1997. Thesaurus of sponge morphology. Smithson. Contrib. Zool.

474 596: 1–55.. doi:10.5479/si.00810282.596.Draft

475 Brøndsted, H. V. 1932. Marine Spongia. The Zoology of the Faroes 1: 1–34.

476 Burton, M. 1931. The Folden Fiord. Report on the sponges collected by Mr. Soot-Ryven in the Folden

477 Fiord in the year 1923. Tromsø Museum Skr. 1(13): 1–8.

478 Carter, H.J. 1876. Descriptions and Figures of Deep-Sea Sponges and their Spicules, from the Atlantic

479 Ocean, dredged up on board H.M.S. Porcupine, chiefly in 1869 (concluded). Ann. Mag. Nat. Hist.

480 Clark, K., Karsch-Mizrachi, I., Liman, D., Ostell, J., and Sayers, E. 2016. GenBank. Nucleic

481 Acids Res. 44: 67–72.

482 Cooper, J.A., Goodwin, C., Lawton, P., Brydges, T., Hiltz, C., Armsworthy, S., and Mccurdy, Q. 2019.

483 Characterisation of the sublittoral habitats of the Brier Island/Digby Neck Ecologically and

484 Biologically Significant Area, Nova Scotia. Can. Tech. Rep. Fish. Aquat. Sci. 3349 3327: 1–163.

485 Dinn, C. 2020. A new species of Haliclona (Flagellia) Van Soest, 2017 (Porifera, Demospongiae,

486 Heteroscleromorpha) from the Gulf of St. Lawrence, Canada. Zootaxa 4778(2): 391–395.

487 doi:10.11646/zootaxa.4778.2.10.

488 Dinn, C., and Leys, S.P. 2018. Field Guide to Sponges of the Eastern Canadian Arctic. Univ. Alberta.

489 Educ. Res. Arch.: 1–102. doi:https://doi.org/10.7939/R3DF6KJ4G.

490 Dinn, C., Leys, S.P., Roussel, M., and Methe, D. 2020. Geographic range extensions of stalked,

491 flabelliform sponges (Porifera) from eastern Canada with a new combination of a species of

492 Plicatellopsis in the North Atlantic. Zootaxa 4755(2): 301–321. doi:10.11646/zootaxa.4755.2.6.

493 Folmer, O., Black, M., Hoeh, W., Lutz, R., and Vrijenhoek, R. 1994. DNA primers for amplification of

494 mitochondrial cytochrome c oxidase subunit I from diverse metazoan invertebrates. Mol. Mar. Biol.

495 Biotechnol. 3(5): 294–299. doi:10.1071/ZO9660275.

496 Fuller, S. 2011. Diversity of Marine Sponges in the Northwest Atlantic. Dalhousie University. Available

497 from papers3://publication/uuid/94E4A59B-4041-4287-BCD1-CACBBD196CB9.

498 Ginn, B.K., Logan, A., Thomas, M.L.H., and Van Soest, R.W.M. 1998. Hymedesmia canadensis

499 (Porifera: Poecilosclerida), a new speciesDraft among new geographical records from the Bay of Fundy,

500 New Brunswick, Canada. J. Mar. Biol. Assoc. United Kingdom 78: 1093–1100.

501 Goodwin, C.E. 2017. Field guide to sponges of the Bay of Fundy. Huntsman Marine Science Centre. St

502 Andrews, New Brunswick.

503 Goodwin, C.E., Berman, J., and Hendry, K.R. 2019. Demosponges from the sublittoral and shallow-

504 circalittoral (<24m depth) Antarctic Peninsula with a description of four new species and notes on in

505 situ identification characteristics. Zootaxa 4658(3): 461–508. doi:10.11646/zootaxa.4658.3.3.

506 Hentschel, E. 1929. Die Kiesel- und Hornschwämme des Nördlichen Eismeers. In Fauna Arctica. Eine

507 Zusammenstellung der arktischen Tierformen mit besonderer Berücksichtigung des Spitzbergen-

508 Gebietes auf Grund der Ergebnisse der Deutschen Expedition in das Nördliche Eismeer im Jahre

509 1898. Edited by F. Römer, F. Schaudinn, A. Brauer, and W. Arndt. G.Fischer, Jena. pp. 857–1042.

510 Hestetun, J.T., Tompkins-Macdonald, G., and Rapp, H.T. 2017. A review of carnivorous sponges

511 (Porifera: Cladorhizidae) from the Boreal North Atlantic and Arctic. Zool. J. Linn. Soc. 181: 1–69.

512 Hogg, M.M., Tendal, O.S., van Soest, R.W.M., Gutt, J., Krautter, M., and Roberts, J.M. 2010. Deep Sea

513 Sponge Grounds: Reservoirs of Biodiversity. UNEP-WCMC Biodiversity Series No. 32. UNEP-

514 WCMC, Cambridge.

515 Hooper, J.N.A., and Van Soest, R.W.M. 2002. Systema Porifera: A Guide to the Classification of

516 Sponges. Edited ByJ.N.A. Hooper and R.W.M. Van Soest. Springer US, Boston, MA.

517 doi:10.1007/978-1-4615-0747-5.

518 Hurlbut, T., and Clay, D. 1990. Protocols for research vessel cruises within the Gulf Region 540

519 (demersal fish) (1970-1987). Can. Manuscr. Rep. Fish. Aquat. Sci. 2082 143 p.

520 Kenchington, E.L.R., Beazley, L.I., Lirette, C., Murillo, F.J., Guijarro, J., Wareham, V., et al.2016.

521 Delineation of Coral and Sponge Significant Benthic Areas in Eastern Canada Using Kernel Density

522 Analyses and Species Distribution Models. DFO - Can. Sci. Advis. Secr. Res. Doc. (November):

523 178.

524 Koltun, V.M. 1959. [Siliceous horny sponges of the northern and fareastern seas of the U.S.S.R.] [In

525 Russian]. Opredeliteli po faune SSR, Izd.Draft Zool. muzeem Akad. Nauk 67: 1-236.

526 Lambe, L.M. 1896. Sponges from the Atlantic coast of Canada. Trans. R. Soc. Canada 2(2): 181–211.

527 Lambe, L.M. 1900a. Catalogue of the Recent Marine Sponges of Canada and Alaska. Ottawa Nat. 14(9):

528 153–172.

529 Lambe, L.M. 1900b. Sponges from the coasts of north-eastern Canada and Greenland. Trans. R. Soc.

530 Canada 6: 19–49.

531 Lévi, C. 1959. Résultats scientifiques des Campagnes de la ‘Calypso’’. Campagne de la “Calypso’’ dans

532 le Golfe de Guinée et aux îles Principe, São Tomé et Annobon. 5. Spongiaires.”’ Ann. l’Institut

533 océanographique.

534 Maldonado, M. 2006. The ecology of the sponge larva. Can. J. Zool. 84(2): 175–194. doi:10.1139/Z05-

535 177.

536 Maldonado, M., Aguilar, R., Bannister, R.J., Bell, J.J., Conway, K.W., Dayton, P.K., et al. 2015. Sponge

537 Grounds as Key Marine Habitats: A Synthetic Review of Types, Structure, Functional Roles, and

538 Conservation Concerns. In Marine Animal Forests. Springer International Publishing, Cham. pp. 1–

539 39. doi:10.1007/978-3-319-17001-5_24-1.

540 Meyer, C.P. 2003. Molecular systematics of cowries (Gastropoda: Cypraeidae) and diversification

541 patterns in the tropics. Biol. J. Linn. Soc. 79: 401–459.

542 Morrow, C.C., and Cárdenas, P. 2015. Proposal for a revised classification of the Demospongiae

543 (Porifera). Front. Zool. 12:7doi:https://doi.org/10.1186/s12983-015-0099-8.

544 Mosquin, T., Whiting, P.G., and McAllister, D.E. 1995. Canada’s biodiversity: The variety of life, its

545 status, economic benefits, conservation costs and unmet needs. Ottawa: Canadian Museum of

546 Nature. Canadian Museum of Nature., Ottowa.

547 Murillo, F., Kenchington, E., Tompkins, G., Beazley, L., Baker, E., Knudby, A., and Walkusz, W. 2018a.

548 Sponge assemblages and predicted archetypes in the eastern Canadian Arctic. Mar. Ecol. Prog. Ser.

549 597: 115–135. Available from https://www.int-res.com/abstracts/meps/v597/p115-135/.

550 Murillo, F.J., Kenchington, E., Beazley, L., Lirette, C., Knudby, A., Guijarro, J., Benoît, H., Bourdages,

551 H., and Sainte-Marie, B. 2016a. DistributionDraft Modelling of Sea Pens , Sponges , Stalked Tunicates

552 and Soft Corals from Research Vessel Survey Data in the Gulf of St . Lawrence for Use in the

553 Identification of Significant Benthic Areas. Can. Tech. Rep. Fish. Aquat. Sci. 3170: 1–132

554 Murillo, F.J., Kenchington, E., Lawson, J.M., Li, G., and Piper, D.J.W. 2016b. Ancient deep-sea sponge

555 grounds on the Flemish Cap and Grand Bank, northwest Atlantic. Mar. Biol. 163(3): 1–11.

556 doi:10.1007/s00227-016-2839-5.

557 Murillo, F.J., Kenchington, E.L., Clark, D., Emberley, J., Regnier-McKellar, C., Guijarro, J., Beazley,

558 L.I., and Wong, M.C. 2018b. Cruise Report for the CCGS Alfred Needler Maritimes Region

559 Research Vessel Summer Multispecies Survey, June 28 to August 14, 2017: Benthic Invertebrates.

560 Can. Tech. Rep. Fish. Aquat. Sci. 3262: 1–41.

561 Old, M.C. 1941. The Taxonomy and Distribution of the Boring Sponges (Clionidae) along the Atlantic

562 coast of North America. Chesap. Biol. Lab. Publ. 44: 1–30.

563 Picton, B.E., and Goodwin, C.E. 2007. Sponge biodiversity of Rathlin Island, Northern Ireland. J. Mar.

564 Biol. Assoc. UK 87(06): 1441–1458. doi:10.1017/S0025315407058122.

565 Pozdnyakov, I., and Karpov, S.A. 2016. Kinetid structure in Choanocytes of Sponges

566 (Heteroscleromorpha): Toward the ancestral Kinetid of Demospongiae: Kinetid Structure of

567 Heteroscleromorph Choanocytes. J. Morphol. 277(7): 925–34. doi:10.1002/jmor.20546.

568 Redmond, N.E., Morrow, C.C., Thacker, R.W., Diaz, M.C., Boury-Esnault, N., Cárdenas, P. et al.. 2013.

569 Phylogeny and systematics of demospongiae in light of new small-subunit ribosomal DNA (18S)

570 sequences. Integr. Comp. Biol. 53(3): 388–415. doi:10.1093/icb/ict078.

571 Rezvoi, P. 1925. Über neue Schwämme aus den Meeren der russischen Arctis. Zool. Anz. 62(9/10): 193–

572 201.

573 Sarà, M. 1969. Specie nuove di Demospongiae provenienti dal coralligeno pugliese. . Bolletino di Mus. e

574 Ist. dell’Università di Genova 37(255): 89-96.

575 Van Soest, R.W.M. 1984. Marine sponges from Curaçao and other Caribbean localities. Part III.

576 Poecilosclerida. In: Hummelinck, P.W. & Van der Steen, L.J. (Eds), Uitgaven van de

577 Natuurwetenschappelijke Studiekring Draftvoor Suriname en de Nederlandse Antillen. No. 112. Stud.

578 Fauna Curaçao other Caribb. Islands 66(199): 1–167.

579 Van Soest, R.W.M. 2002. Family Crellidae Dendy, 1922. In Systema Porifera. Edited by J.N.A. Hooper

580 and R.W.M. van Soest. Kluwer Academic/Plenum Publishers, New York. doi:10.1007/978-1-4615-

581 0747-5_59.

582 Van Soest, R.W.M., Boury-Esnault, N., Hooper, J.N.A., Rützler, K., de Voogd, N.J., Alvarez, B. et al..

583 2021. World Porifera Database (continually updated). doi:doi:10.14284/359.

584 Van Soest, R.W.M., Boury-Esnault, N., Vacelet, J., Dohrmann, M., Erpenbeck, D., de Voogd, N.J. et al..

585 2012. Global diversity of sponges (Porifera). PLoS One 7(4): e35105.

586 doi:10.1371/journal.pone.0035105.

587 Vargas, S., Erpenbeck, D., Göcke, C., Hall, K.A., Hooper, J.N.A., Janussen, D., Wörheide, G. 2013.

588 Molecular phylogeny of Abyssocladia (Cladorhizidae: Poecilosclerida) and Phellodermidae:

589 Poecilosclerida) suggests a diversificiation of chelae microscleres in clodorhizid sponges. Zoologica

590 scripta 42 (1):106–116. https://doi.org/10.1111/j.1463-6409.2012.00560.x

591 Thacker, R.W., Hill, A.L., Hill, M.S., Redmond, N.E., Collins, A.G., Morrow, C.C. et al.. 2013. Nearly

592 complete 28S rRNA gene sequences confirm new hypotheses of sponge evolution. Integr. Comp.

593 Biol. 53(3): 373–387. doi:10.1093/icb/ict071.

594 Tompkins, G., Baker, E., Anstey, L., Walkuzc, W., Siford, T., and Kenchington, E.L.R. 2017. Sponges

595 from the 2010-2014 Paamiut Multispecies Trawl Surveys, Eastern Arctic and Subarctic: Class

596 Demospongiae, Subclass Heteroscleromorpha, Order Poecilosclerida, Family Coelosphaeridae,

597 Genera Forcepia and Lissodendoryx. Can. Tech. Rep. Fish. Aquat. Sci. 3224: 1–129.

598 Topsent, E. 1890. Notice préliminaire sur les spongiaires recueillis durant les campagnes de l’Hirondelle.

599 Bull. la Société Zool. Fr. 15: 65-71.

600 Topsent, E. 1892. Contribution à l’étude des Spongiaires de l’Atlantique Nord (Golfe de Gascogne, Terre-

601 Neuve, Açores). Résultats des campagnes Sci. Accompl. par le Prince Albert I. Monaco. 2 : 1–165.

602 Topsent, E. 1925. Etude des Spongiaires du Golfe de Naples. Arch. Zool. expérimentale générale.

603 63 :623–725. Draft

604 Vacelet, J. 1961. Spongiaires (Démosponges) de la région de Bonifacio (Corse). Recl. des Trav. Stn. Mar.

605 d’Endoume 22:21–45.

606 Vacelet, J. 1969. Eponges de la Roche du Large et de l’étage bathyal de Méditerranée (Récoltes de la

607 soucoupe plongeante Cousteau et dragages). Mémoires du Muséum national d’Histoire naturelle.

608 Mémoires du Muséum Natl. d’Histoire Nat. (A, Zool.) 59:145–219.

609 Vacelet, J., and Perez, T. 1998. Two new genera and species of sponges (Porifera, Demospongiae)

610 without skeleton from a Mediterranean cave. Zoosystema 20: 5–22.

611 Wassenberg, T.J., Dews, G., and Cook, S.D. 2002. The impact of fish trawls on megabenthos

612 (sponges) on the north-west shelf of Australia. Fish. Res. 58(2): 141–151.

613 doi:10.1016/S0165-7836(01)00382-4.

614 Wessel, P., and W. H. F. Smith (1996), A global, self-consistent, hierarchical, high-resolution shoreline

615 database, J. Geophys. Res., 101(B4): 8741–8743, doi:10.1029/96JB00104.)

616 Whiteaves, J.F. 1901. Catalogue of the marine Invertebrata of eastern Canada. Rep. Geol. Surv. Canada

617 772: 1–272.

Draft

Tables Table 1. Comparison of spicule sizes of Atlantic and Mediterranean Crella species. Measurements are listed as min(mean)max.

Species/ Growth Acanthostyles Acanthostyles Acanthostyles Tylotes Isochelae Sigmas Distribution and notes source form choanosomal ectosomal echinating/ basal Crella Massively 427(478)539 x 294(331)384 x 99(132)192 x 360(389)413 x 21(24)27 32(36)39 Type locality Northwest of St. (Pytheas) encrusting 13(17)19 11(12)14 7(10)11 9(10)12 Paul Island, Nova Scotia. 205 cutis sp. nov. m. ARC 81601 Crella Massively 439(476)507 x 202(295)353 x 89(124)166 x 334(366)425 x 16(21)27 32(39)48 Northern Gulf of St. (Pytheas) encrusting 13(17)19 8(11)14 8(10)13 7(10)12 (n=19, very Lawrence, west of Daniels cutis sp. nov. rare) Harbour, Newfoundland and ARC 81602 Labrador. 150 m.

Crella Massively 377(485)512 x 260(319)384 x 92(155)234 x 305(370)422 x 18(20)22 28(32)42 Scotian shelf. 84 m. (Pytheas) encrusting 12(13)15 9(11)13 7(9)11 6(8)10 (n=7, very cutis sp. nov. Draft rare). ARC 81500 C. (P.) Encrusting - 83.3(96)104.6 73.6(86.1)110.5 185.3(201)229.5 16.9(18.7)21.7 43.4(50.6)56 Azores/Mediterranean. sigmata x 2.6(3.1)3.5 x 3.5(5)4.61 x 2.7(3.7)4.6 and Measurements from Boury- Topsent 1925 28.4(34.8)40.6 Esnault and Lopes 1985 C. (P.) donsi Encrusting - 69(84.1)98 x 153(178)210 x 171(204)243 x 18(20.1)24 present Norway. Burton 1931 3(4)5 5(5.8)7 2.5(3.2)4 Measurements from Van Soest (2002) C. (P.) Encrusting - 117(131.7)141 72(196.3)300 x 289(302.8)320 x 19(21.4)24 - Caribbean. chelifera x 2.5(3.0)3.5 3.5(5.16)7 2.5(3.2)4 Species has ectosomal Van Soest, (acanthoxea) (tornotes) acanthoxea instead of 1984 acanthostyles.

C. (P.) Encrusting 320–420 x 6–7 100–280 x – – 38 - Faroes. akraleitae 4–82 (Brøndsted, 1932) C. (P.) atra Encrusting - 375 365 375 30 - Azores.

1 Error in source, average basal acanthostyle thickness is larger than the range. 2 Acanthostyles are “larger at the base of the sponge” suggesting two size categories.

(Topsent Measurements from Topsent 1890) 18923 and Van Soest (2002) C. (P.) Massive - 250 x 7 630 x 8 560 x 7 25 - Norway. basispinosa Burton, 1931 C. (P.) Digitate - 90–130 x 5–7 130–150 x 5–7 260–300 x 2.5–5 17–22 - East Atlantic/ Mediterranean. digitifera Measurements from Vacelet (Lévi 1959) 1961 C. (P.) Massively - 120–140 x 250–350 x 325–375 x 5–7 35–40 - Norway. Jaegerskloldi encrusting 4–5 unknown 7–8 unknown if Alander, 1937 if ectosomal or ectosomal or basal basal C. (P.) alba Encrusting - 160–280 x 150–160 x 260–370 x 5 22–27 - Mediterranean. Vacelet, 1969 5–10 7.5–8

C. (P.) Encrusting - 70(96)110 x 91–135 x 10–13 220–265 x 3–5 17–21.5 - South European shelf. fusifera 3.5(7)7.5 Draft(tornotes) Species has ectosomal Sarà, 1969 (acanthoxea) acanthoxea instead of acanthostyles. C. (P.) plana Encrusting 105–165 x 2–8 120–175 x 2–8 185–230 x 4–8 310–370 x 5–7 15–18 - Northern Ireland. Includes a Picton and (acanthoxea) (tornotes) separate category of Goodwin, acanthoxea in addition to two 2007 sizes of acanthostyle.

C. (P.) Encrusting - 180(200)230 x 160(220)260 190(310)390 16(19)21 - Norway. Spicule widths not schottlaenderi 10 (tornotes) given. (Arndt 1913)

C. (Yvesia) Massive 200–500 x 6 80–160 x 4 - 300–400 x 4 24 - Norway. mammillata Has large choanosomal (Arnesen, spicules similar to C. (P.) 1903) cutis sp. nov., but lacks echinating acanthostyles and sigmas.

3 Listed as Crella (Pytheas) ater in Topsent 1892

Draft

Table 2 - Comparison of spicule sizes of Crellomima species. All spicule dimensions in µm. For type specimens, measurements in regular font are from type description, those in italics are from our measurements of type specimens. Measurements are listed as min(mean)max.

Species Basal Ectosomal Tornotes Chelae Distribution and Notes acanthostyles acanthostyles Crellomima imparidens 92–214 x 6–15 93–190 x 4–7 160–240 x 5–7 16–22 Type locality White Sea, 38 m. Also recorded from Rezvoi 1925. ZIN 7011. 138(160)184 x 106(117)147 x 185(198)226 x 21,22 Sweden (Alander 1942) and the Okhotsk Sea (Koltun 11(17)22 6(7)8.5 4(7)10 1959). Recent records from Kandalaksha bay, White Sea (Pozdnyakov and Karpov 2016,). Fine, dense, leather–like crusts on barnacles. Surface covered with small (0.5 mm) papillae spaced 1–1.5 mm apart. Colour bright rose. Draft Chelae with five, short, rounded alae on each end. Crellomima incrustans 182–224 x 9–11 113–132 x 6–7 210–250 x 4.5– 16–17 Type locality North and East Barents Sea (depth not Hentschel 1929 144(171)187 x 129(139)150 x 5 19(21)22 given). Encrusting on a delicate shell. Chelae very BMNH 1936.11.20.14. 13(15)18 7(10)13 239(252)273 x abundant. All chelae umbelliform with 7–9 alae at each 5(7)9 end and a straight shaft. Longer tornotes than other species and fatter ectosomal acanthostyles.

Crellomima 152(166)192 x 120(138)188 x 185(204)218 x 18(22)25 Type locality Deer Island, New Brunswick, also recorded mehqisinpekonuta sp. nov. 8(14)18 5(5)6 4(6)7 from Brier Island, Nova Scotia. Currently known from ARC 80713 depths of 6–21 m. Chelae very rare in most specimens, Crellomima 139(173)218 x 117(146)160 x 183(205)221 x 20,25 not found in ARC 80709. Chelae have 5-8 teeth, the alae mehqisinpekonuta sp. nov. 10(12)15 3(4)5 4(5)7 (rare) continue down the shaft, diminishing in size ARC 80705

Crellomima 123(156)199 x 112(136)148 x 154(192)246 x None mehqisinpekonuta sp. nov. 8(10)15 2(4)6 3(5)6 observed ARC 80709 Crellomima 139(189)209 x 113(146)167 x 191(207)222 x 14(19)24 mehqisinpekonuta sp. nov. 8(12)18 4(5)7 4(4)5 ARC 80712 Crellomima derma 126–210 96–168 200–232 17–19 Type locality North and East Barents Sea, 300 m. Thin Hentschel, 1929 124(165)233 x 109(119)133 x 188(225)250 x 14(19)21 crust on worm tube. Chelae have 3–5 alae and shaft with a ZMH S3259 12(17)21 7(9)10 4(6)8 pronounced curve. Crellomima derma 151(191)228 x 134(149)165 x 199(224)246 x 16,18 Thin, orange, hispid, thinly encrusting sponge. Chelae with ARC 80736 10(13)16 3(5)7 Draft5(7)9 (rare) 3–5 alae at each end, fairly flat, wide shaft. Crellomima derma 165(209)236 x 103(131)154 x 213(228)252 x 16(19)21 ARC 80747 12(16)20 4(5)7 5(7)9

Draft

Figure 1. Specimen collection locations. A, Eastern Canada showing sampling locations in the Quoddy region of the Bay of Fundy (b) and Brier Island (c). B, St. Andrews and Deer Island SCUBA locations. C, Brier Island SCUBA locations in southwest Nova Scotia. Circles, Crella (Pytheas) cutis sp. nov. Triangles, Crellomima mehqisinpekonuta sp. nov. Squares, Crellomima derma. Figure was created using ArcMap 10.7.1 and assembled from the following data sources: Collection data sourced from DFO and CG. Base map from GSHHG (Wessel & Smith, 1996), and contour lines from NOAA National Geophysical Data Center ETOPO1 (Amante & Eakins, 2009).

210x152mm (600 x 600 DPI)

Draft

Figure 2. Crella (Pytheas) cutis sp. nov. A,B, collected specimens. C,D, choanaosomal acanthostyles. E,F, tylotes. G,H, ectosomal acanthostyles. I,J, echinating/basal acanthostyles. K-M, isochelae. N,O, sigmas. P,Q, skeleton.

170x199mm (300 x 300 DPI)

Draft

Figure 3. Crellomima mehqisinpekonuta sp. nov. A, Holotype in situ. B, skeleton. C,D basal acanthostyle. E, ectosomal acanthostyle. F,G, tornotes. H,I, isochelae.

170x187mm (300 x 300 DPI)

Draft

Figure 4. Crellomima derma, Bay of Fundy Specimen. A, specimen in situ. B, skeleton. C basal acanthostyle. D, ectosomal acanthostyle. E,F, tornotes. G,H, isochelae.

170x177mm (300 x 300 DPI)

Draft

Figure 5. Crellomima derma ZMH S3259. A, basal acanthostyle. B, ectosomal acanthostyle. C,D, tornotes. E,F, isochelae.

170x118mm (300 x 300 DPI)

Draft Figure 6. Crellomima incrustans BMNH 1936.11.20.14. A,B, basal acanthostyle. C, ectosomal acanthostyle. D,E, tornotes. F,G, isochelae.

170x111mm (300 x 300 DPI)

Draft Figure 7. Crellomima imparidens ZISP ZIN7011. A, basal acanthostyle. B, ectosomal acanthostyle. C,D, tornotes. E,F, isochelae.

170x110mm (300 x 300 DPI)