Addition to Sweden’s freshwater sponge fauna and a phylogeographic study of Spongilla lacustris (Spongillida, Porifera) in southern Sweden Author Robert, Chloé 384E6BC8-C928-4D4E-B7DB-BC285556ABCB Dept. Organismal Biology, Uppsala University, Norbyvägen 18 D, SE- 752 36 Uppsala, Sweden. & Dept. Marine Sciences, University of Gothenburg, Box 461, SE- 405 30 Göteborg, Sweden. chloe.robert@gu.se Author Pereira, Raquel 2E0077C7-7540-4E45-B3B0-32A4A4F1D50B Dept. Organismal Biology, Uppsala University, Norbyvägen 18 D, SE- 752 36 Uppsala, Sweden. raquel.pereira@ebc.uu.se Author Thollesson, Mikael C1EF10A0-3D59-4C66-A2A6-2BD48F154721 Dept. Organismal Biology, Uppsala University, Norbyvägen 18 D, SE- 752 36 Uppsala, Sweden. mikael.thollesson@ebc.uu.se text European Journal of Taxonomy 2022 2022-07-13 828 1 138 167 http://dx.doi.org/10.5852/ejt.2022.828.1861 journal article 97042 10.5852/ejt.2022.828.1861 db902fb6-703b-4457-a4e8-37be2a702d74 2118-9773 6839811 92E8908C-818C-49CD-8E84-A95479F8E455 Spongilla lacustris ( Linnaeus, 1759 ) Fig. 2 Spongia lacustris Linnaeus, 1759: 1348 . Material examined ( 109 specimens , Table 1 ) SWEDEN Dalsland 3 specs ; Lake Vänern , Mellerud , Sunannå harbour ; 58.7092° N , 12.5072° E ; 11 Oct.2017 ; Chloé Robert and Raquel Pereira leg.; UPSZMC 188195 , 188197 , 188199 4 specs ; Åklång , Dalslands kanal , Håverud ; 58.8214° N , 12.4061° E ; 11 Oct. 2017 ; Chloé Robert and Raquel Pereira leg.; UPSZMC 188201 , 188203 , 188205 , 188207 . – Bohuslän 1 spec. ; Bohuslän , Lake S Bullaresjön , Naverstad , Sundshult ; 58.7403° N , 11.5743° E ; 8 Oct. 2017 ; Chloé Robert leg.; UPSZMC 188185 3 specs ; Lake S Bullaresjön , Östad ; 58.7719° N , 11.5693° E ; 8 Oct. 2017 ; Chloé Robert and Raquel Pereira leg.; UPSZMC 188189 , 188191 , 188193 . Södermanland 5 specs ; Lake Likstammen , Öster Malma ; 58.9468° N , 17.1708° E ; 29 Jun. 2013 ; Mikael Thollesson leg.; UPSZMC 188012 , 188015 to 188018 9 specs ; Lake Naten , Stjärnhof bath ; 59.0774° N , 17.0129° E ; 30 Jun. 2013 ; Mikael Thollesson leg.; UPSZMC 188023 , 188029 to 188036 5 specs ; Lake Kyrksjön , bath ; 59.0923° N , 16.9951° E ; 30 Jun. 2013 ; Mikael Thollesson leg.; UPSZMC 188024 to 188028 5 specs ; Klämmingen , Solbacken ; 59.1307° N , 17.2454° E ; 29 Jun. 2013 ; Mikael Thollesson leg.; UPSZMC 188014 , 188019 to 188022 .– Uppland 1 spec. ; Krägga herrgård ; 59.603° N , 17.394° E ; 2 Aug. 2017 ; Chloé Robert and Raquel Pereira leg.; UPSZMC 188114 2 specs ; Viks bath ; 59.7346° N , 17.4639° E ; 2 Aug. 2017 ; Chloé Robert and Raquel Pereira leg.; UPSZMC 188110 , 188112 3 specs ; Hammarskogs bath ; 59.7636° N , 17.5762° E ; 2 Aug. 2017 ; Chloé Robert and Raquel Pereira leg.; UPSZMC 188122 to 188124 4 specs ; Flottsund ; 59.7875° N , 17.6626° E ; 2Aug.2017 ; Chloé Robert and Raquel Pereira leg.; UPSZMC 188107 , 188115 , 188120 , 188121 1 spec. ; Lake Erken , Norr Malma , Erken freshwater laboratory ; 59.8353° N , 18.6331° E ; 30 May 2016 ; Raquel Pereira leg.; UPSZMC 188213 4 specs ; Lake Siggeforasjön ; 59.9759° N , 17.157° E ; 26 May 2014 ; Tove Fällman leg.; UPSZMC 188100 , 188102 , 188104 , 188106 2 specs ; Gimo damm ; 60.1786° N , 18.1718° E ; 3 Aug. 2017 ; Chloé Robert and Raquel Pereira leg.; UPSZMC 188137 , 188140 5 specs ; Simbadet , Österbybruk ; 60.195° N , 17.9143° E ; 3 Aug. 2017 ; Chloé Robert and Raquel Pereira leg.; UPSZMC 188126 , 188129 , 188131 , 188133 , 188135 .– Dalarna 4 specs ; Dalälven , Grådö ; 60.2471° N , 16.027° E ; 28 Aug. 2013 ; Mikael Thollesson leg.; UPSZMC 188037 , 188056 , 188059 , 188061 1 spec. ; Lake Viksjön , Strands boat club ; 60.4841° N , 15.7447° E ; 28 Aug. 2013 ; Mikael Thollesson leg.; UPSZMC 188064 4 specs ; Lake Flosjön , harbour , Dala-Floda ; 60.513° N , 14.8089° E ; 28 Aug. 2013 ; Mikael Thollesson leg.; UPSZMC 188048 , 188050 , 188054 , 188057 6 specs ; Lake S Mojesjön , bath ; 60.6003° N , 15.0885° E ; 28 Aug. 2013 ; Mikael Thollesson leg.; UPSZMC 188039 , 188041 to 188043 , 188045 , 188046 2 specs ; Malungs camping , outlet into Västerdalälven ; 60.6847° N , 13.7023° E ; 29 Aug. 2013 ; Mikael Thollesson leg.; UPSZMC 188072 , 188075 5 specs ; Lake Orsasjön , Strandrestaurangen ; 61.1186° N , 14.5985° E ; 30 Aug. 2013 ; Mikael Thollesson leg.; UPSZMC 188082 to 188084 , 188086 , 188088 2 specs ; Lake Skattungen , Oresjöns boat club ; 61.1649° N , 15.1327° E ; 30 Aug. 2013 ; Mikael Thollesson leg.; UPSZMC 188090 , 188091 2 specs ; Lake Hällsjön , northern rest stop ; 62.0681° N , 12.6744° E ; 28 Aug. 2003 ; Mikael Thollesson leg.; UPSZMC 188000 , 188003 . – Hälsingland 3 specs ; Lake Ängratörn , Vikstenstorpet ; 61.9159° N , 15.3872° E ; 31 Aug. 2013 ; Mikael Thollesson leg.; UPSZMC 188092 , 188094 , 188096 . – Härjedalen 12 specs ; Hedeviken ; 62.4083° N , 13.6697° E ; 30 Aug. 2017 ; Chloé Robert and Raquel Pereira leg.; UPSZMC 188165 , 188166 , 188170 to 188177 , 188179 , 188181 . – Medelpad 2 specs ; Lake Gissjön ; 62.5233° N , 16.179° E ; 29 Aug. 2017 ; Chloé Robert and Raquel Pereira leg.; UPSZMC 188147 , 188148 . – Jämtland 2 specs ; Lake Storsjön , Galhammar ; 62.7784° N , 14.4321° E ; 30 Aug. 2017 ; Chloé Robert and Raquel Pereira leg.; UPSZMC 188161 , 188183 3 specs ; Lake Sundsjön ; 62.9513° N , 15.1623° E ; 29 Aug. 2017 ; Chloé Robert and Raquel Pereira leg.; UPSZMC 188155 , 188157 , 188159 4 specs ; Lake Storsjön , Slandrom ; 63.1347° N , 14.6478° E ; 29 Aug. 2017 ; Chloé Robert and Raquel Pereira leg.; UPSZMC 188209 to 188212 . Table 1 (continued). Material examined in the present study. Abbreviations of Swedish provinces: Bhl = Bohuslän; Dlr = Dalarna; Dls = Dalsland; Hls = Hälsingland; Hrj = Härjedalen; Jmt = Jämtland; Mpd = Medelpad; Sdm = Södermanland; Sth = Stockholm; Upp = Uppland.
Collection site GenBank accession numbers
Species Specimen ID Designation Coordinates Catchment coxI 28S i56 i56 Voucher
area variety number(s)
Spongilla P059-170830-1 Storsjön, Galhammar, Jmt 62°46.7′ N Indalsälven UPSZMC 188161,
lacustris P059-170830-9 14°25.93′ E OL979196 188162 UPSZMC 188183,
P059-190702-1 Storsjön, Slandrom, Jmt 63°8.08′ N 14°38.87′ E 188184 UPSZMC 188209
P059-190702-2 UPSZMC 188210
P059-190702-3 UPSZMC 188211
P059-190702-4 UPSZMC 188212
P059-170829-1 Gissjön, Mpd 62°31.4′ N Ljungan OL979206 UPSZMC 188147
P059-170829-2 16°10.74′ E UPSZMC 188148
P059-170829-6 Sundsjön, Jmt 62°57.08′ N OM105905 w-377G UPSZMC 188155,
P059-170829-7 15°9.74′ E OM105907 w-377G 188156 UPSZMC 188157, 188158
P059-170829-8 OM105933 w-377G UPSZMC 188159,
P059-130831-1 Ängratörn, 61°54.95′ N Ljusnan 188160 UPSZMC 188092,
P059-130831-2 by Vikstenstorpet, His 15°23.23′ E OL979184 OM105936 y-290A 188093 UPSZMC 188094,
P059-130831-3 OL979198 OL985646 188095 UPSZMC 188096, 188097
P059-170830-11 Hedeviken, Hrj 62°24.5′ N 13°40.18′ E UPSZMC 188165
P059-170830-12 OM105912 y-290G UPSZMC 188166,
P059-170830-14 OM105904 y-290G 188167 UPSZMC 188170
P059-170830-15 UPSZMC 188171
P059-170830-16 OM105919 y-290G UPSZMC 188172
P059-170830-17 UPSZMC 188173
P059-170830-18 OM105901 y-290G UPSZMC 188174
Table 1 (continued on next six pages). Material examined in the present study.
Collection site GenBank accession numbers
Species Specimen ID Designation Coordinates Catchment coxI 28S i56 i56 Voucher
area variety number (s)
Spongilla P059-170830-19 OM 105903 y-290G UPSZMC 188175
lacustris P059-170830-20 OM 105932 y-290A UPSZMC 188176
P059-170830-21 UPSZMC 188177,
P059-170830-22 188178 UPSZMC 188179, 188180
P059-170830-23 OM 105924 y-290G UPSZMC 188181, 188182
P059-130828-1 Dalälven, Grådö, Dlr 60°14.83′ N Dalälven OL985645 UPSZMC 188037,
P059-130828-2 16°1.62′ E 188038 UPSZMC 188056
P059-130828-3 OM 105921 z-242 T UPSZMC 188059,
145 P059-130828-4 188060 UPSZMC 188061
P059-130828-6 Strands boat club, Dlr 60°29.05′ N 15°44.68′ E UPSZMC 188064, 188065
P059-130828-16 Flosjön, boat site, Dlr 60°30.78′ N 14°48.53′ E UPSZMC 188048, 188049
P059-130828-17 UPSZMC 188050,
P059-130828-19 188051 UPSZMC 188054,
P059-130828-20 OM 105913 z-242 T 188055 UPSZMC 188057, 188058
P059-130828-10 S Mojesjön, bathing place, Dlr 60°36.02′ N 15°5.31′ E UPSZMC 188039, 188040
P059-130828-11 UPSZMC 188041
P059-130828-12 UPSZMC 188042
P059-130828-13 UPSZMC 188043,
P059-130828-14 188044 UPSZMC 188045
P059-130828-15 UPSZMC 188046, 188047
Table 1 (continued). Material examined in the present study.
Collection site GenBank accession numbers
Species Specimen ID Designation Coordinates Catchment coxI 28S i56 i56 Voucher
area variety number (s)
Spongilla P059-130829-1 Malungs camping, mouth of 60°41.08′ N UPSZMC 188072,
lacustris P059-130829-2 the river in Västerdalälven Dlr 13°42.14′ E 188073 UPSZMC 188074
P059-130830-1 Orsasjön, at the beach restaurant, Dlr 61°7.12′ N 14°35.91′ E UPSZMC 188082
P059-130830-2 UPSZMC 188083
P059-130830-3 OM 105910 y-290G UPSZMC 188084, 188085
P059-130830-4 UPSZMC 188086,
P059-130830-5 OM 105899 y-290G 188087 UPSZMC 188088,
P059-130830-6 Skattungen, Oresjöns boat club, Upp 61°9.89′ N 15°7.96′ E OM 105915 z-242G 188089 UPSZMC 188090
P059-130830-7 OM 105925 z-242 T UPSZMC 188091
P052-030828-1 Hällsjön, northern resting place, Dlr 62°4.09′ N 12°40.46′ E OL979188 UPSZMC 188000
P052-030830-1 OL979195 UPSZMC 188003
P059-170803-6 Gimo dam, Upp 60°10.72′ N Olandsån UPSZMC 188137,
P059-170803-7 18°10.31′ E OM 105931 y-290G 188138 UPSZMC 188140,
UP-16-2-1 Dock at Erken-lab, Upp 59°50.12′ N 18°37.99′ E Norrtäljeån OM 105917 w-377G 188141 UPSZMC 188213, 188214
P059-170802-13 Krägga mansion, Upp 59°36.18′ N 17°23.64′ E Norrström UPSZMC 188114
P059-170802-11 Viks badplats, Upp 59°44.08′ N UPSZMC 188110,
P059-170802-12 17°27.83′ E OM 105902 w-377G 188111 UPSZMC 188112,
P059-170802-7 Hammarskogs badplats, Upp 59°45.82′ N 17°34.57′ E OL985663 OM 105920 z-242 T 188113 UPSZMC 188122
P059-170802-8 OL979192 UPSZMC 188123
P059-170802-9 UPSZMC 188124, 188125
Table 1 (continued). Material examined in the present study.
Collection site GenBank accession numbers
Species Specimen ID Designation Coordinates Catchment coxI 28S i56 i56 Voucher
Spongilla P059-170802-1 Flottsund, Upp 59°47.25′ N area OL979179 variety OM105923 y-290G number(s) UPSZMC 188107
lacustris P059-170802-2 17°39.76′ E UPSZMC 188115
P059-170802-5 UPSZMC 188120
P059-170802-6 UPSZMC 188121
P059-140526-1 Siggefora Lake, Upp 59°58.55′ N 17°9.42′ E UPSZMC 188100, 188101
P059-140526-2 UPSZMC 188102,
P059-140526-3 188103 UPSZMC 188104,
P059-140526-4 188105 UPSZMC 188106
P059-170803-1 Simbadet, Österbybruk, Upp 60°11.7′ N 17°54.86′ E UPSZMC 188126, 188127
P059-170803-2 UPSZMC 188129, 188130
P059-170803-3 OM105928 w-377G UPSZMC 188131,
P059-170803-4 188132 UPSZMC 188133,
P059-170803-5 188134 UPSZMC 188135, 188136
P059-130629-1 Likstammen,east Malma,Sdm 58°56.81′ N 17°10.25′ E Svärtaån OM105930 y-290G UPSZMC 188012, 188013
P059-130629-2 UPSZMC 188015
P059-130629-3 OM105897 y-290G UPSZMC 188016
P059-130629-4 OL985649 OM105922 z-242T UPSZMC 188017
P059-130629-5 OM105896 y-290G UPSZMC 188018
P059-130630-1 Naten, Stjärnhof bathing 59°4.64′ N OL979207 OL985655 OM105927 y-290G UPSZMC 188023
P059-130630-2 place, Sdm 17°0.77′ E OL985644 OM105908 y-290G UPSZMC 188029
P059-130630-3 OL979201 OM105911 w-377G UPSZMC 188030
Table 1 (continued). Material examined in the present study.
Collection site GenBank accession numbers
Species Specimen ID Designation Coordinates Catchment coxI 28S i56 i56 Voucher
area variety number (s)
Spongilla P059-130630-4 OM 105900 z-242 T UPSZMC 188031
lacustris P059-130630-5 OL979189 OL985652 UPSZMC 188032
P059-130630-6 OL985667 UPSZMC 188033
P059-130630-7 OL985658 UPSZMC 188034
P059-130630-8 OL985660 OM 105934 z-242 T UPSZMC 188035
P059-130630-9 OL985659 OM 105914 w-377G UPSZMC 188036
P059-130630-10 Kyrksjön, bathing place, Sdm 59°5.54′ N UPSZMC 188024
P059-130630-11 16°59.71′ E UPSZMC 188025
P059-130630-12 UPSZMC 188026
P059-130630-13 UPSZMC 188027
P059-130630-14 UPSZMC 188028
P059-130629-10 Klämmingen, Solbacken, Sdm 59°7.84′ N 17°14.72′ E OL979203 OL985664 OM 105926 w-377G UPSZMC 188014
P059-130629-6 OL979187 OL985656 UPSZMC 188019
P059-130629-7 OL979182 OL985670 UPSZMC 188020
P059-130629-8 OL985648 OM 105916 y-24G UPSZMC 188021
P059-130629-9 OL979191 OM 105909 y-290G UPSZMC 188022
P059-171011-1 Vänern, Sunannå Hamn, Mellerud, Dls 58°42.55′ N 12°30.43′ E Göta Älv UPSZMC 188195, 188196
P059-171011-2 OM 105935 w-377G UPSZMC 188197,
P059-171011-4 OM 105898 w-377G 188198 UPSZMC 188199,
P059-171011-6 Åklång, Dalslands kanal, Håverud, Dls 58°49.28′ N 12°24.37′ E 188200 UPSZMC 188201, 188202
P059-171011-7 UPSZMC 188203, 188204
P059-171011-8 OM 105906 w-377 T UPSZMC 188205, 188206
Table 1 (continued). Material examined in the present study.
Collection site GenBank accession numbers
Species Specimen ID Designation Coordinates Catchment coxI 28S i56 i56 Voucher
area variety number (s)
Spongilla P059-171011-9 UPSZMC 188207,
lacustris 188208
P059-171008-1 Bullaresjön, Sundshult, 58°44.42′ N Enningdalsälven OM 105929 y-290G UPSZMC 188185,
P059-171008-3 Naverstad, Bhl S Bullaresjön, Östad, Bhl 11°34.46′ E 58°46.31′ N 188186 UPSZMC 188189,
P059-171008-4 11°34.16′ E 188190 UPSZMC 188191,
188192
P059-171008-5 OL979199 UPSZMC 188193,
188194
Eunapius P059-170830-10 Storsjön, Galhammar, Jmt 62°46.7′ N Indalsälven OL979181 UPSZMC 188163,
fragilis P059-170830-2 14°25.93′ E OL979200 188164 UPSZMC 188215,
188216
P059-170830-3 OL979183 UPSZMC 188221,
188222
P059-170830-4 OL979180 OL985661 UPSZMC 188217,
P059-170830-5 OL979202 OL985653 188218 UPSZMC 188223,
P059-170830-6 OL979209 188224 UPSZMC 188225,
188226
P059-170830-7 OL979185 UPSZMC 188227,
188228
P059-170830-8 OL979210 OL985651 UPSZMC 188219,
P059-170802-10 Hammarskogs badplats, Upp 59°45.82′ N Norrström OL979193 OL985650 188220 UPSZMC 188108,
P059-170802-3 Flottsund, Upp 17°34.57′ E 59°47.25′ N OL979205 188109 UPSZMC 188116,
17°39.76′ E 188117
P059-170802-4 OL979197 UPSZMC 188118,
188119
Ephydatia P059-170829-3 Gissjön, Mpd 62°31.4′ N Ljungan OL979204 OL985642 UPSZMC 188149,
muelleri P059-170829-4 16°10.74′ E OL985669 188150 UPSZMC 188151,
188152
P059-170829-5 OL979190 OL985668 UPSZMC 188153,
188154
Table 1 (continued). Material examined in the present study.
Collection site GenBank accession numbers
Species Specimen ID Designation Coordinates Catchment coxI 28S i56 i56 Voucher
Ephydatia P059-170828-1 Delsbo, Södra Dellensjön, Hls 61°48.58′ N area Delångersån variety number(s) UPSZMC 188143,
muelleri P059-170828-2 16°34.24′ E OL979186 OL985654 188144 UPSZMC 188145,
188146
P059-170830-13 Hedeviken, Hrj 62°24.5′ N Ljusnan OL979194 OL985657 UPSZMC 188168,
13°40.18′ E 188169
P059-130828-5 Dalälven, Grådö, Dlr 60°14.83′ N Dalälven OL985662 UPSZMC 188062,
P059-130828-18 Flosjön, boat site, Dlr 16°1.62′ E 60°30.78′ N 188063 UPSZMC 188052,
P059-130828-7 14°48.53′ E S Mojesjön, bathing place, Dlr 60°36.02′ N OL985647 188053 UPSZMC 188066,
15°5.31′ E 188067
P059-130828-8 OL985671 UPSZMC 188068,
188069
P059-130828-9 OL985666 UPSZMC 188070
P059-130829-3 Malungs camping, mouth of 60°41.08′ N OL985665 UPSZMC 188076,
P059-130829-4 the river in Västerdalälven, Dlr 13°42.14′ E 188077 UPSZMC 188078,
P059-130829-5 188079 UPSZMC 188080,
188081
UP-16-2-2 Dock at Erken-lab, Upp 59°50.12′ N Norrtäljeån UPSZMC 188244,
UP-16-1-2 Badplatsen Svanberga: Erken 18°37.99′ E 59°50.22′ N 188244 UPSZMC 188242,
- S bathing place drainage 18°39.35′ E 188243
P059-171011-3 pipe, Upp Vänern, Sunannå Hamn, 58°42.55′ N Göta Älv UPSZMC 188237,
P059-171011-5 Mellerud, Dls 12°30.43′ E 188238 UPSZMC 188239,
Ephydatia P059-170902-1 Helgarsviken, Sth 59°17.34′ N Östersjön OL979208 OL985643 188240 UPSZMC 188229,
fluviatilis 18°42.19′ E (the Baltic) 188230
P059-170902-2 UPSZMC 188231,
P059-170902-3 188232 UPSZMC 188233,
P059-211231-1 188234 PENDING
Fig. 2. Spongilla lacustris ( Linnaeus, 1759 ) . A–B . Habitus. A . Specimen growing under a pontoon (P059-171011-6). B . Specimen growing on anchor chain, with finger–like projections (P059-170802- 48). C–D . Spiculae. C . Megasclere (P059-171008-4). D . Microsclere (P059-171008-1). E–F . Estimated spicula size distribution within and between specimens, P059-130831-1 (solid line), P059-170802- 8 (dashed) and P059-171008-4 (dotted). Marks on the x-axis represent spiculae measured. Brackets correspond to ranges in literature (red = Tendal 1967b ; blue = Penney & Racek 1968 ; black = Evans & Montagnes 2019 ). E . Megascleres. F . Microscleres. Table 2. Co-occurrence of freshwater sponges at different sites in the present study. On the diagonal are the number of sites where the species was found, lower triangle gives the number of co-occurrences, and upper diagonal shows Sørensen-Dice indices of species association (ranging from 0 – never found together to 1 – always found together). At two sites the three species Spongilla lacustris ( Linnaeus, 1759 ) , Ephydatia muelleri (Lieberkühn, 1856) and Eunapius fragilis ( Leidy, 1851 ) were found together; at no site did all four species co-occur.
Spongilla lacustris Ephydatia fluviatilis Ephydatia muelleri Eunapius fragilis
Spongilla lacustris 30 0 0.40 0.18
Ephydatia fluviatilis 0 1 0 0
Ephydatia muelleri 8 0 10 0
Eunapius fragilis 3 0 0 3
Description HABITUS. Specimens usually bright green ( Fig. 2A ), often with finger-like projections ( Fig. 2B ). Surface hispid due to protruding spicules. SPICULAE. Skeleton with two classes of oxeas: smooth megascleres ( Fig. 2C ) and spined microscleres ( Fig. 2D ). Megascleres fusiform, microscleres acerate ( Boury-Esnault & Rützler 1997 ). Gemmuloscleres were not observed. Megasclere length 265 μm (190–375 μm), width 11 μm (2.5–20 μm) ( Fig. 2E ). Microsclere length 76 μm (50–117.5 μm), width 4.8 μm (2.5–7.5 μm) ( Fig. 2F ). Distribution and habitat Widely distributed, entire sampling area; hard substrates in lakes and rivers. Remarks The finger-like projections distinguish this species from the other Swedish freshwater sponges, as does the presence of microscleres.