A common garden of Halichondria sponges: taxonomic revision of Northeast Pacific Halichondriidae reveals many cryptic introduced species

Halichondria panicea MZ486738 New Hampshire Halichondria panicea MZ486927 New Hampshire Halichondria panicea MZ486929 Maine Halichondria panicea MZ487251 Maine Halichondria panicea MZ486931 Maine Halichondria panicea MZ487247 Maine Halichondria panicea MZ487246 Maine Halichondria panicea MZ487253 Maine Halichondria panicea MZ487259 New Hampshire Halichondria panicea MZ486745 New Hampshire Halichondria panicea MZ486928 Maine Halichondria panicea MZ487261 New Hampshire Halichondria panicea MZ487250 Maine Halichondria panicea MZ487249 Maine Halichondria panicea MZ486932 Maine Halichondria panicea MZ486935 Maine Halichondria panicea MZ487252 Maine Halichondria panicea MZ486742 New Hampshire Halichondria panicea MZ486933 Maine Halichondria panicea MZ486934 Maine Halichondria panicea MZ486930 Maine Halichondria panicea MZ487244 Maine Halichondria panicea MZ487258 New Hampshire Halichondria panicea MZ486925 Maine Halichondria panicea ESS17 Nova Scotia Halichondria panicea MZ486926 Maine Halichondria panicea MZ487245 Maine Halichondria panicea ESS74 Nova Scotia Halichondria panicea 180923 Sweeden Halichondria panicea ESS80 Nova Scotia Halichondria panicea Quoddy124 New Brunswick Halichondria panicea Quoddy113 New Brunswick Halichondria panicea 191026 Sweeden Halichondria panicea NIB0608 Ireland Halichondria panicea MK079945 Denmark Halichondria panicea MH756603 Iceland Halichondria panicea ESS11 Nova Scotia Halichondria panicea DUT0047 Alaska Halichondria panicea ESS18 Nova Scotia Halichondria panicea 160504 Sweeden Halichondria panicea 190424 Sweeden Halichondria panicea 160505 Sweeden Halichondria panicea 191116 Sweeden Halichondria panicea 190624 Sweeden Halichondria panicea 181204 Sweeden Halichondria panicea MZ486924 Maine Halichondria panicea MZ487248 Maine Halichondria panicea MZ487254 Maine Halichondria panicea ASGRD043-18 Bering Sea Halichondria panicea NLMAR577-20 North Sea Halichondria panicea KC869423 Ireland Halichondria panicea NLMAR567-20 North Sea Halichondria panicea NLMAR567-20 North Sea Halichondria panicea NLMAR577-20 North Sea Halichondria panicea 274026 Alaska Halichondria panicea BHAK03453 British Columbia Halichondria panicea TLT858 California Halichondria panicea TLT1371 California Halichondria panicea TLT1370 Washington Halichondria panicea BOIMB0056 Oregon Halichondria panicea OP535459 California Halichondria panicea BOIMB0189 Oregon Halichondria panicea TLT1430 Washington Halichondria panicea FH12x84 Washington Halichondria panicea FH12x38 Washington Halichondria panicea FH12x81 Washington Halichondria panicea FH12x39 Washington Halichondria panicea 274016 Alaska Halichondria panicea 274014 Alaska Halichondria panicea MG422903 British Columbia Halichondria panicea BHAK00536 British Columbia Halichondria panicea TLT1293 Washington Halichondria sitiens ESS130 Nova Scotia Halichondria sitiens ESS153 Nova Scotia Halichondria sitiens ESS44 Nova Scotia Halichondria sitiens ESS132 Nova Scotia Halichondria sitiens RBC11 British Columbia Halichondria sitiens RBC13 British Columbia Topsentia fernaldi TLT1134 California Topsentia fernaldi TLT1261 California Halichondria akesa ESS07 Nova Scotia Halichondria columbiana BHAK09466 British Columbia Halichondria columbiana BHAK09479 British Columbia Halichondria hygeia TLT645 California Halichondria sp MZ487266 Connecticut Halichondria californiana TLT256 California Halichondria hygeia FH12x114 Washington Halichondria hygeia WMS19x7 New York Halichondria hygeia TLT742 California Halichondria californiana TLT428 California Halichondria hygeia WMS19x6 New York Halichondria californiana TLT928 California Halichondria californiana TLT921 California Halichondria californiana TLT812 California Halichondria hygeia TLT1076 California Halichondria californiana TLT1010 California Halichondria sp MN448237 Washington Halichondria hygeia TLT706 California Halichondria hygeia TLT308 California Halichondria californiana TLT860 California Halichondria californiana TLT934 California Halichondria hygeia TLT940 California Halichondria californiana TLT417 California Halichondria californiana TLT937 California Halichondria californiana TLT160 California Halichondria sp KY492588 Portugal Halichondria hygeia BHAK03597 British Columbia Halichondria hygeia FH12x108 Washington Halichondria hygeia WMS21x10 New York Halichondria hygeia WMS21x9 New York Halichondria hygeia TLT724 California Halichondria hygeia TLT504 California Halichondria hygeia WMS19x5 New York Halichondria cascadia FH12x93 Washington Halichondria cascadia MZ580608 Alaska Halichondria cascadia FH12x92 Washington Halichondria akesa ROPOR090-18 Svalbard Halichondria akesa MH756604 Iceland Halichondria pinaza NIB0604 Ireland Halichondria pinaza TLT892 California Halichondria pinaza TLT1056 California Halichondria pinaza TLT873 California Halichondria sp BNSB100-21 Germany Halichondria pinaza TLT869 California Halichondria zabra TLT780 California Halichondria zabra TLT751 California Halichondria zabra TLT624 California Halichondria zabra TLT761 California Halichondria zabra TLT722 California Halichondria zabra TLT623 California Halichondria zabra TLT1317 California Halichondria pinaza NIB0603 Ireland Halichondria sp MZ487264 New Hampshire Halichondria zabra TLT978 California Halichondria zabra TLT1086 California Halichondria zabra TLT967 California Halichondria pinaza TLT1080 California Halichondria sp MZ486936 New York Halichondria sp MZ487241 New York Halichondria sp OQ323005 Virginia Halichondria sp OQ322884 Virginia Halichondria sp OQ323432 Virginia Halichondria sp OQ323360 Virginia Halichondria zabra TLT757 California Halichondria zabra TLT712 California Halichondria sp MZ487262 New Hampshire Halichondria zabra TLT760 California Halichondria sp MZ487265 New Hampshire Halichondria zabra TLT1077 California Halichondria zabra TLT815 California Halichondria zabra TLT759 California Halichondria zabra TLT979 California Halichondria zabra TLT1082 California Halichondria zabra TLT756 California Halichondria zabra TLT1064 California Halichondria zabra TLT612 California Halichondria zabra TLT737 California Halichondria zabra TLT874 California Halichondria sp MZ487260 New Hampshire Halichondria bowerbanki NLMAR086-20 Netherlands Halichondria bowerbanki NLMAR106-20 Netherlands Halichondria bowerbanki TLT490 California Halichondria bowerbanki TLT234 California Halichondria bowerbanki TLT415 California Halichondria bowerbanki TLT342 California Halichondria bowerbanki KY492591 Portugal Halichondria bowerbanki NIB0309 Ireland Halichondria bowerbanki MZ487221 Argentina Halichondria bowerbanki MZ487236 Argentina Halichondria bowerbanki MZ487239 Argentina Halichondria bowerbanki 190615 Portugal Halichondria bowerbanki 190616 Portugal Halichondria bowerbanki KY492592 Portugal Halichondria bowerbanki NIB0612 Ireland Halichondria bowerbanki NIB0614 Ireland Halichondria bowerbanki TLT1052 California Halichondria bowerbanki TLT1357 California Halichondria bowerbanki TLT1418 California Halichondria bowerbanki TLT337 California Halichondria bowerbanki TLT980 California Halichondria bowerbanki TLT738 California Halichondria bowerbanki TLT1358 California Halichondria bowerbanki TLT981 California Halichondria bowerbanki TLT1072 California Halichondria bowerbanki KY492589 Portugal Halichondria urca TLT766 California Halichondria urca TLT926 California Halichondria urca TLT747 California Halichondria urca TLT1063 California Halichondria urca TLT740 California Halichondria urca TLT534 California Halichondria urca TLT591 California Halichondria urca TLT960 California Halichondria urca TLT593 California Halichondria urca 15492 California Halichondria urca TLT733 California Halichondria urca TLT590 California Halichondria urca TLT1002 California Halichondria urca TLT951 California Halichondria urca TLT764 California Halichondria urca TLT1087 California Halichondria urca OQ322699 Virginia Halichondria urca OQ322807 Virginia Halichondria urca MZ487242 New York Halichondria urca OQ323255 Virginia Halichondria urca NIB0610 Ireland Halichondria coerulea MW059055 Hawaii Halichondria coerulea MT586735 Hawaii Halichondria loma TLT1225 California Halichondria loma TLT936 California Halichondria loma OP534737 California Halichondria loma TLT1024 California Halichondria loma TLT989 California Halichondria loma TLT976 California Halichondria loma TLT1130 California Halichondria loma OP534724 California Hymeniacidon actites TLT227 California Hymeniacidon actites TLT856 California Hymeniacidon actites TLT1158 California Hymeniacidon actites TLT847 California Halichondria dokdoensis MK105765 Korea Halichondria dokdoensis TLT386 California Halichondria dokdoensis NLMAR324-20 Netherlands Terpios hoshinota KJ008098 China Terpios hoshinota KP764915 Hymeniacidon kitchingi TLT412 California Hymeniacidon kitchingi TLT597 California Hymeniacidon kitchingi TLT491 California Hymeniacidon kitchingi TLT820 California Hymeniacidon kitchingi TLT977 California Hymeniacidon kitchingi TLT154 California Hymeniacidon kitchingi TLT941 California Hymeniacidon kitchingi TLT378 California Dictyonella funicularis MZ487627 Grand Cayman Hymeniacidon gracilis KY565326 Hawaii Hymeniacidon pierrei TLT599 California Aff Protosuberites KJ620398 Australia Homaxinella sp LN850194 Antarctica Hymeniacidon fusiformis TLT1194 California Semisuberites cribrosa BHAK09371 British Columbia Hymeniacidon sp Quoddy115 Bay of Fundy Canada Plicatellopsis fragilis LN850217 Antarctica Plicatellopsis antarctica LN850221 Antarctica Pseudosuberites hyalinus LN850222 Antarctica Aff Protosuberites KJ620404 Australia Protosuberites sp AY561979 Pacific Panama Terpios gelatinosus KX866738 Israel Protosuberites mereui KX601203 Sardinia Protosuberites sp MG825741 Ukraine Suberitidae sp MW059096 Hawaii Protosuberites sp MT226344 Vietnam Suberitidae sp MW144976 Hawaii Protosuberites incrustans KX601189 Norway Protosuberites sp KX601204 Thailand Protosuberites sp OR126051 California Halichondria galea EF217336 Korea Halichondria galea EF217337 Korea Halichondria galea EF217339 California Halichondria galea TLT496 California Halichondria galea EF217338 Korea Halichondria galea TLT495 California Halichondria galea TLT743 California Halichondria galea KX244759 Korea Halichondria okadai EF217341 Korea Halichondria okadai NC 037391 Korea Halichondria okadai MG267395 Halichondria sp MW059058 Hawaii Halichondria magniconulosa P19x171 Caribbean Panama Halichondria magniconulosa P19x173 Caribbean Panama Halichondria magniconulosa EF519616 Caribbean Halichondria melanodocia EF519617 Caribbean Halichondria melanodocia EF519618 Caribbean Halichondria magniconulosa EF519615 Caribbean Halichondriidae sp MK833923 Myanmar Halichondriidae sp MK833924 Myanmar Halichondria sp P19x40 Pacific Panama Halichondria sp P19x63 Pacific Panama Amorphinopsis sp MG029167 Hawaii Amorphinopsis maculosa JQ034560 Australia Amorphinopsis sp MW059039 Halichondriidae sp MK833925 Myanmar Halichondriidae sp MK833926 Myanmar Axinyssa sp KX894484 Vietnam Terpios sp MW059101 Hawaii Hymeniacidon chloris KY565323 Hawaii Amorphinopsis fenestrata JQ034540 Australia Amorphinopsis foetida JQ034541 Singapore Ciocalypta sp EF519604 Queensland Hymeniacidon sp MW059078 Hawaii Hymeniacidon sp MW143251 Hawaii Hymeniacidon perlevis ON062402 South Africa Hymeniacidon perlevis TLT129 California Hymeniacidon perlevis TLT79 California Hymeniacidon perlevis TLT651 California Hymeniacidon heliophila EU076812 Brazil Hymeniacidon perlevis KY492578 Portugal Hymeniacidon perlevis TLT247 California Hymeniacidon perlevis TLT736 California Hymeniacidon perlevis TLT1074 California Hymeniacidon perlevis KF192342 Korea Hymeniacidon perlevis NIB0615 Ireland Hymeniacidon perlevis NIB0616 Ireland Hymeniacidon perlevis NIB0611 Ireland Hymeniacidon flavia EF217333 Korea Hymeniacidon flavia EF217334 Korea Hymeniacidon flavia EF217335 Korea Protosuberites ectyoninus KX601188 Spain Halichondria sp JQ034561 Queensland Halichondria sp LC436885 Iran Halichondria sp LC436883 Iran Halichondria sp LC436882 Iran Halichondria sp LC436877 Iran Halichondria sp KX894504 Vietnam Homaxinella sp MT491669 South Africa Homaxinella sp MT491670 South Africa Pseudosuberites latke OR126050 California Pseudosuberites sp MN547324 Korea Pseudosuberites nudus LN850224 Antarctica Pseudosuberites sp MT491504 South Africa Protosuberites denhartogi KX601190 United Kingdom Suberites ficus OL877279 Korea Suberites ficus OL877281 Korea Suberites ficus OL877280 Korea Suberites latus OR126063 British Columbia Suberites latus MG423083 British Columbia Suberites latus MZ580937 Aleutian Islands Suberites latus MG423083 Suberites latus OR126065 British Columbia Suberites latus OR126067 British Columbia Suberites latus OR126066 British Columbia Suberites latus MG421261 British Columbia Suberites latus OR126068 British Columbia Suberites ficus AJ843891 Massachusetts Suberites dandelenae KY463455 South Africa Suberites dandelenae KY463456 South Africa Suberites sp MT483568 Suberites domuncula AM690374 Adriatic Suberites ficus HQ379429 Ireland Suberites pagurorum KC869422 Ireland Suberites latus OR126072 British Columbia Stylocordyla borealis MK372890 Stylocordyla chupachups LN850189 Antarctica Stylocordyla chupachups LN850243 Antarctica Suberites lambei OR126054 California Suberites lambei OR126055 California Suberites lambei OR126052 California Suberites lambei OR879308 Washington Suberites lambei OR879309 Washington Suberites lambei OR126056 California Suberites lambei AY561966 Suberites lambei OR126069 British Columbia Suberites lambei OR126070 Washington Suberites lambei OR126060 California Suberites lambei OR126057 California Suberites lambei OR126064 Oregon Suberites lambei OR879306 Washington Suberites lambei OR126071 Oregon Suberites lambei OR879307 Washington Suberites agaricus PP068272 California Suberites diversicolor FJ968448 Suberites diversicolor LC436878 Persian Gulf Suberites diversicolor KF568963 Indonesia Suberites diversicolor MT226353 Vietnam Suberites diversicolor MT226352 Vietnam Suberites aurantiacus EF519679 Florida Suberites aurantiacus EF519681 Florida Suberites aurantiacus EF519680 Florida Rhizaxinella sp AY561983 Rhizaxinella gadus OR126058 California Rhizaxinella gadus OR126059 California Rhizaxinella gadus OR126061 California Rhizaxinella gadus OR126062 California Rhizaxinella sp OP863334 Greece Rhizaxinella sp LR880962 Ireland Aaptos aaptos KX866769 Israel Hymeniacidon sp. JQ034565 Australia Aaptos suberitoides MH784603 Vietnam Aaptos papillata KY492599 Portugal Aaptos papillata KY492600 Portugal Aaptos nuda MT491602 Aaptos nuda MT491604 Homaxinella amphispicula OR126053 California Aaptos sp. MZ487069 Grand Cayman Polymastia littoralis KJ129611 Tethya actinia AY320033 69 99 93 72 91 24 70 100 29 100 13 95 100 100 51 96 29 96 65 48 100 59 68 28 82 22 91 62 96 93 73 1 20 100 100 79 34 78 99 92 23 100 14 3 3 100 100 100 77 20 89 7 99 21 48 46 100 100 97 1 58 87 98 73 70 37 93 100 100 79 100 42 57 30 57 75 100 56 66 22 83 3 100 48 76 5 100 3 35 96 85 31 100 47 8 98 92 96 50 67 4 35 100 90 89 63 10 96 55 36 90 32 60 100 100 100 81 100 97 38 40 100 96 61 100 9 100 81 100 90 4 100 88 29 82 1 71 100 84 16 4 6 38 95 58 20 15 1 90 10 59 1 5 100 1 7 83 98 20 14 63 76 88 59 97 86 2 2 91 99 91 39 42 94 50 73 12 81 100 100 100 100 98 99 87 11 8 90 69 2 17 86 89 94 2 2 90 2 56 51 43 87 42 100 90 46 7 67 55 41 4 93 14 100 99 4 100 14 63 12 47 2 85 87 97 37 1 44 73 100 2 86 59 4 49 100 59 50 84 63 12 96 73 17 100 100 83 65 100 42 95 19 100 98 2 92 72 100 100 82 86 49 100 69 92 37 90 16 9 98 17 70 2 77 61 42 65 13 100 5 63 11 64 54 100 100 77 5 84 100 100 100 52 64 52 100 65 78 31 100 99 100 2 99 47 3 31 98 2 7 99 4 96 14 97 2 99 4 47 7 94 99 99 100 100 29 100 4 96 100 60 97 28 98 52 5 38 23 86 99 84 100 100 47 100 Tree scale: 0.1