Josephella marenzelleri Caullery & Mesnil, 1896

Bastida-Zavala, J. Rolando, McCANN, Linda D., Keppel, Erica & Ruiz, Gregory M., 2017, The fouling serpulids (Polychaeta: Serpulidae) from United States coastal waters: an overview, European Journal of Taxonomy 344, pp. 1-76 : 7-12

Key to shallow-water serpulids from the United States

1. With operculum ................................................................................................................................ 2

– Without operculum ......................................................................................................................... 45

2. Opercular peduncle thin, with pinnules (operculum on one radiole) ............................................... 3

– Opercular peduncle thick and smooth .............................................................................................. 4

3. Five thoracic chaetigers; capillary ( Fig. 9L ) and sharply-limbate collar chaetae ( Fig. 9M ) .................... .................................................................................... Josephella marenzelleri Caullery & Mesnil, 1896

– Six to 12 thoracic chaetigers; collar with fin-and-blade chaetae, with basal denticulate expansion and flattened distal blade ( Fig. 9N ) ................................................... Filogranula calyculata ( Costa, 1861) 1

4. Collar with chaetae ...................................................................................................................................5

– Collar without chaetae .................................................................................................................... 43

5. Operculum funnel-shaped, without chitinous plates ( Figs 2 A–D, 4A–J, 9A–B) ...................................6

– Operculum with chitinous ( Fig. 7A ) or calcareous plates ( Fig. 9C, G ), sometimes fused, sometimes with spines like “deer horns”, never funnel-shaped ....................................................................... 28

– Operculum otherwise ...................................................................................................................... 40

6. Operculum as a simple funnel, calcified ........................................ Pyrgopolon ctenactis ( Mörch, 1863)

– Operculum as a simple funnel, fleshy ( Fig. 2 A–D) .......................................................................... 7

– Operculum as a fleshy basal funnel and a distal chitinous verticil ( Fig. 4 A–J) .................................. ......................................................................................................... Hydroides Gunnerus, 1768 …12

7. Operculum with basal digitate or rounded processes ( Fig. 2 A–D) ......... Crucigera Benedict, 1887 …8

– Operculum with no basal processes ( Fig. 9 A–B) ................................ Serpula Linnaeus, 1758 …9 2

8. Operculum with one basal rounded process; inner surface of funnel smooth ....................................................................................................... Crucigera irregularis Bush, 1905

– Operculum with three basal rounded processes, one dorsal and two laterals ( Fig. 2 C–D), sometimes the dorsal rounded process split in two; inner surface of funnel smooth ........................................... ............................................................................................................. C. zygophora ( Johnson, 1901)

– Operculum peduncle with four basal digitate processes ( Fig. 2B ); inner surface of funnel with tubercles ( Fig. 2B ) ................................................................................... C. websteri Benedict, 1887

9. Opercular funnel with less than 20 radii; shallow constriction between peduncle and operculum .......................................................................... Serpula sp. A in ten Hove & Wolf 1984 3

– Opercular funnel with more than 20 radii; deep constriction between peduncle and operculum .. 10

10. Opercular funnel short, with 46–160 radii ( Fig. 9 A–B), pointed tips, inner surface with numerous tiny conical tubercles ......................................................................... S. columbiana Johnson, 1901 4

– Opercular funnel elongate; with 21–35 radii, rounded tips, inner surface smooth ..........................11

11. Tube with five longitudinal ridges; collar with bayonet chaetae (similar to Fig. 9 H–J) with five teeth ............................................................................................................... S. watsoni Willey, 1905

– Tube with 6–8 longitudinal ridges; collar with bayonet chaetae with two teeth ................................ ................................................................................................... S. vossae Bastida-Zavala, 2012 2

12. Verticil spines without lateral spinules, with expanded tips, without wings ( Fig. 4D ) .................. 13

– Verticil spines with lateral spinules, without wings ( Fig. 4A, 4E, 4 H–I) ....................................... 14

– Verticil spines without lateral spinules ( Fig. 4 B–C, 4 G), without expanded tips, sometimes with wings ( Fig. 4F, 4J ) .......................................................................................................................... 21

13. Verticil spines with rounded tips; bayonet chaetae with two teeth and proximal rasp ( Fig. 9K ) ......................................................................................... Hydroides microtis Mörch, 1863

– Verticil spines with flat almost T - shaped tips ( Fig. 4D ); bayonet chaetae with two teeth, without proximal rasp ( Fig. 9 H–I) ......................................................................... H. dirampha Mörch, 1863

14. Verticil spines with only one pair of lateral spinules ( Fig. 4A, 4I ) ................................................. 15

– Verticil spines with more than one pair of lateral spinules ( Fig. 4E, 4H ) ....................................... 18

15. Funnel without radii, with 3–6 blunt processes only; verticil with straight spines ............................ ............................................................................................................ H. cf. mucronatus Rioja, 1958

– Funnel formed by radii; verticil spines curving inwards ................................................................ 16

16. Funnel radii with blunt tip ( Fig. 4A ) ........................................................... H. bispinosa Bush, 1910

– Funnel radii with pointed tip ( Fig. 4I ) ............................................................................................ 17

17. Verticil spines abruptly curving inwards, with a pronounced distal knob .......................................... ................................................................................................................. H. parva ( Treadwell, 1902)

– Verticil spines curving inwards rather smoothly, without distal knob ...... H. crucigera Mörch, 1863

18. Verticil spines complex, with many lateral and external spinules; dark-brown basally and hyaline distally; internal spinules enlarged ...................................................... H. mongeslopezi Rioja, 1958

– Verticil spines simple, with few lateral and without external spinules; uniform colour pattern; internal spinules short .................................................................................................................................. 19

19. Verticil spines with 2–4 pairs of lateral spinules; central spine short ( Fig. 4E ) or without this spine ................................................................................................................................................ 20 – Verticil spines with 5–8 pairs of lateral spinules; central spine long ( Fig. 4H ) .................................. ......................................................................................................... H. longispinosa Imajima, 1976b

20. Verticil spines straight or curving outwards ( Fig. 4E ) ............................ H. elegans ( Haswell, 1883) – Verticil spines curving inwards ...................................... Hydroides sp. A in ten Hove & Wolf 1984 3

21. Funnel radii with laterally expanded tips, T-shaped ........................... H. gairacensis Augener, 1934 – Funnel radii with pointed tips ( Fig. 4 B–C, 4 F–G, 4 J) .................................................................... 22

22. Verticil spines straight or curving outwards; bayonet chaetae with two teeth and proximal rasp ( Fig. 9K ) ............................................................................................. H. protulicola Benedict, 1887

– Verticil with at least dorsal –if not all– spines curving inwards; bayonet chaetae with two teeth without proximal rasp ( Fig. 9 H–I) .................................................................................................. 23

23. Dorsal spines curving inwards, ventral spines curving outwards ( Fig. 4C, 4J ) .............................. 24 – All spines curving inwards ( Fig. 4B, 4 F–G) ................................................................................... 25

24. Verticil spines without external spinule and wings ( Fig. 4C ) .................. H. dianthus ( Verrill, 1873) – Verticil spines with external spinule (at least ventral ones) and wings ( Fig. 4J ) ................................ ......................................................................................... H. sanctaecrucis (Krøyer in Mörch, 1863)

25. Verticil with spines long and slender, gradually curving inwards, without wings, all spines similar in shape and size ( Fig. 4G ) .............................................................................. H. gracilis ( Bush, 1905)

– Verticil with spines short and thick, strongly curving inwards, without wings, 1–5 dorsal spines larger than the others ( Fig. 4B ) ....................................................................................................... 26

– Verticil with spines long and slender, gradually curving inwards, with wings, all spines similar in shape and size ( Fig. 4F ) .................................................................................................................. 27

26. Only one dorsal spine, remaining verticil spines equally sized, with subdistal knobs ....................... .......................................................................................................... H. brachyacantha Rioja, 1941a

– Only one dorsal spine, remaining verticil spines equally sized, without subdistal knobs ( Fig. 4B ) .. ..................................................................................................... H. cf. brachyacantha Rioja, 1941a

– 3–5 dorsal bigger spines, with external tubercles on smaller spines .................................................. ............................ H. cf. amri Sun, Wong, ten Hove, Hutchings, Williamson & Kupriyanova, 2015 5

27. Branchial crown with more than 29 pairs of radioles; funnel with 45–61 radii ................................. ............................................................................................................. H. spongicola Benedict, 1887

– Branchial crown with less than 24 pairs of radioles; funnel with 27–38 radii ( Fig. 4F ) .................... ................................................................................................................... H. floridana ( Bush, 1910)

28. Operculum with 1–7 distal calcareous plates ( Fig. 9C, 9G ), sometimes with spines like “deer horns” ....................................................................................... Spirobranchus Blainville, 1818 …29

– Operculum with a column of chitinous concentric plates ( Fig. 7A ), sometimes with a distal stalk stellate ............................................................................. Pomatostegus stellatus ( Abildgaard, 1789)

– Operculum with 1–14 distal chitinous plates fused ( Fig. 7G ), sometimes with one distal branched spine or a hook ................................................................................................................................ 34

29. Operculum with 1–7 successive calcareous plates, distal spines absent ................................................... ......................................................................................... Spirobranchus latiscapus (Marenzeller, 1885)

– Operculum with one calcareous plate, with spines similar to deer horns, or when the plate is smooth has an ovoid or conical shape ......................................................................................................... 30

30. Operculum lacks spines ( Fig. 9C, 9G ); with a few limbate collar chaetae ( Fig. 9M ), sometimes none .........................................................................................................................................................31

– Operculum with spines; collar chaetae always present .........................................................................33

31. Tube with two longitudinal ridges, with spine or flap projection over entrance, lacks alveoli (holes) ( Fig. 9D ); collar without chaetae ................................................. Spirobranchus kraussii ( Baird, 1865)

– Tube with one or three longitudinal ridges, never two ridges, lacks projection over entrance, with alveoli ( Fig. 9F ); collar with few chaetae ....................................................................................... 32

32. Operculum ovoid, with a projection; tube with one longitudinal ridge and one line of small alveoli in older sections ..................................................................................................... S. americanus Day, 1973

– Operculum conical ( Fig. 9G ) or bilobed, with 1–2 small projections; tube with three longitudinal ridges and two lines of large alveoli along tube ( Fig. 9F ) ........................... S. minutus Rioja, 1941b

33. Operculum with two very large (equal to or larger than opercular plate) and thin horns and one short horn, with several short spines ............................................................... S. giganteus (Pallas, 1766) 6

– Operculum with two large (smaller than opercular plate) and thick horns, sometimes with a third large horn, with few large spines ..................................................... S. corniculatus (Grube, 1862) 6, 7

– Operculum with five short and wide horns, with several large spines ....... S. spinosus Moore, 1923

34. Thoracic membrane extends up to 3–6 chaetigers; thoracic uncini with anterior blunt tooth ...................................................................................... Vermiliopsis Saint-Joseph, 1894 8 …35

– Thoracic membrane extends up to second chaetiger; thoracic uncini with anterior bifurcate tooth ................................................................................................................................................ 36

35. Operculum with 1–7 fused discs ...................................... Vermiliopsis annulata ( Schmarda, 1861) 8

– Operculum with 10–14 fused discs ................................................. V. multiannulata Gravier, 1906 8

36. Insertion of opercular peduncle is at second radiole; thoracic uncini saw to rasp-shaped ................. ......................................................................... Semivermilia pomatostegoides ( Zibrowius, 1969b) 9

– Insertion of opercular peduncle just below and between first and second radioles; thoracic uncini saw-shaped ...................................................................................... Pseudovermilia Bush, 1907 …37

37. Operculum black or dark brown ( Fig. 7 G ) ..................................................................................... 38

– Operculum yellow or colorless ....................................................................................................... 39

38. Tube with longitudinal ridges and alveoli rows, lacks peristomes and transverse ridges; operculum with tree-shaped spines ............................................... Pseudovermilia multispinosa ( Monro, 1933)

– Tube with longitudinal and transverse ridges, sometimes with peristomes, lacks alveoli; operculum with a simple curved spine, with some minor spines or spines absent ( Fig. 7 G ) ............................... ......................................................................................................... P. occidentalis ( McIntosh, 1885)

39. Tube with several denticulate ridges and alveoli rows, with transverse brown bands ....................... ............................................................................................................. P. fuscostriata ten Hove, 1975

– Tube smooth, with a longitudinal ridge, without alveoli rows, entirely white ................................... ................................................................................................................. P. conchata ten Hove, 1975

40. Tube with a longitudinal keel, without peristomes; operculum with a smooth convex to conical chitinous cap ( Fig. 7 F ); collar with fin-and-blade chaetae, with basal denticulate expansion and flattened distal blade ( Fig. 9 N ) ............................................................... Pseudochitinopoma occidentalis ( Bush, 1905)

– Tube without a longitudinal keel, with peristomes ( Fig. 2 E, 2 I ); operculum with spines ( Fig. 2 G, 2 N ) or smooth ( Fig. 2 J–K); collar chaetae coarsely serrated, without basal expansion ( Fig. 9 O ) ...................................................................................... Ficopomatus Southern, 1921…41

41. Thoracic membranes dorsally fused ( Fig. 2 M, 2 O ); opercular spines curved outwards ( Fig. 2 N ); tropical species ....................................................................... Ficopomatus uschakovi ( Pillai, 1960)

– Thoracic membranes free ( Fig. 2 F ); opercular spines curved inwards ( Fig. 2 G ) or absent ( Fig. 2 J– K) .................................................................................................................................................... 42

42. Operculum concave with black incurving spines ( Fig. 2 G ), not black or lacking in juveniles; subtropical species .............................................................................. F. enigmaticus ( Fauvel, 1923)

– Operculum bulbous ( Fig. 2 J ), slightly convex ( Fig. 2 K ), without spines; tropical species ................ .......................................................................................................... F. miamensis ( Treadwell, 1934)

43. Opaque tube without dorsal ridge, sometimes with brood chambers; thorax with sickle chaetae or “ Apomatus ” chaetae (distally curved) ............................................. Rhodopsis pusilla Bush, 1905 10

– Hyaline tube with dorsal ridge, without brood chambers; thorax without sickle chaetae .................. .......................................................................................................... Placostegus Philippi, 1844 …44

44. Tube without transverse ridges; operculum bulbous proximally, concave and zygomorphic dorsally .............................................................................. Placostegus californicus Hartman, 1969

– Tube with transverse ridges in posterior half section; operculum unknown ...................................... ................................................................................................................... P. incomptus Ehlers, 1887

45. Collar chaetae limbate ( Fig. 9 M ) or capillaries ( Fig. 9 L ), without basal denticulate expansion; uncini with long main tooth ....................................................................................................................... 46

– Collar with fin-and-blade chaetae, with basal denticulate expansion and flattened distal blade ( Fig. 9 N ), or bayonet chaetae with several teeth ( Figs 7 I , 9 J ); uncini with short main tooth ........ 49

46. Seven to nine thoracic chaetigers ............................................................... Protula Risso, 1826 … 47

– 11–14 thoracic chaetigers .............................................. Filogranella sp. in Fosså & Nielsen 1996 11

47. Nine thoracic chaetigers; radioles with a single row of pinnules ....... Protula setosa ( Bush, 1910) 12

– Seven thoracic chaetigers; radioles with a double row of pinnules ............................................. 48 13

48. Tube with longitudinal ridges; branchial crown lacks basal round processes; thoracic membrane narrow, reaches fourth chaetiger ........................................................................... P. longiseta Schmarda, 1861 14

– Tube with rough texture, without longitudinal ridges ( Fig. 7 B ); branchial crown with basal round process on each radiole ( Fig. 7D ); thoracic membrane wide, reaches seventh chaetiger ( Fig. 7 E ) ...................................................................................................... P. balboensis Monro, 1933 14

– Tube smooth, sometimes with shallow transverse ridges; branchial crown with 31–51 radioles; thoracic membrane narrow, reaches seventh chaetiger; uncini smooth .............................. P. atypha Bush, 1905

49. Tube with internal longitudinal ridges ..................................... Spiraserpula Regenhardt, 1961 …50

– Tube without internal longitudinal ridges ....................................... Salmacina Claparède, 1870 …51

50. Tube with dorsal internal ridge serrated, ventral ones with Y-shape .................................................. ................................................................................. Spiraserpula ypsilon Pillai & ten Hove, 1994 15

– Tube with dorsal internal ridge not serrated, ventral ones serrated .................................................... .............................................................................................. S. caribensis Pillai & ten Hove, 1994 15

51. Collar chaetae with many small teeth ............................................................................................. 52

– Collar chaetae with 4–8 large teeth ( Fig. 7 I ) .................................................................................. 53

52. Three, sometimes four, pairs of radioles ( Fig. 7 J ); thoracic uncini with 8–9 rows of teeth and 3–4 teeth per row ................................................................. Salmacina tribranchiata ( Moore, 1923) 16, 17

– Four pairs of radioles; thoracic uncini with seven rows of teeth and 2–3 teeth per row .................... .................................................................................................................. S. dysteri ( Huxley, 1855) 16

– Five pairs of radioles; thoracic uncini with more than 10 rows of teeth ............................................ ................................................................................ Salmacina sp. A in ten Hove & Wolf 1984 3, 16, 17

53. Without prostomial eyes; teeth of collar chaetae distally increasing regularly in size ( Fig. 7 I ); 20 or fewer abdominal segments ( Fig. 7 H ) ....................................................... S. huxleyi ( Ehlers, 1887) 16

– With prostomial eyes; collar chaetae with 4–6 large teeth of similar size; up to 40 abdominal segments ......................................................................................... S. incrustans Claparède, 1870 16

Figures

Fig. 9. Serpulids from United States fouling plates. Serpula columbiana. A. Operculum, from Ketchikan Bay, Alaska (SERC-81981). B. Operculum, adult from San Diego, California (LACMNH-Poly4928);. – Spirobranchus kraussii. C-D. Operculum and tube fragment, from Oahu, Hawaii (SERC-114603). E. Colony, from Natal, South Africa (LACMNH-N5217). – S. minutus. F. Tube, from Santa Catarina, Brazil (USNM-43239). G. Operculum, From Tampa Bay, Florida (SERC-93759). Types of collar chaetae. H–J. Bayonet. K. Bayonet with proximal rasp. L. Capillaries. M. Limbate. N. Fin-andblade chaetae. O. Coarsely serrated. – Hydroides sanctaecrucis (H, L). – H. cf. brachyacantha (I). – Spiraserpula ypsilon (J). – H. elegans (K). – H. ochotereana (M).– Pseudochitinopoma occidentalis (N).– Ficopomatus uschakovi (O, modified from Arteaga-Flórez et al. 2014).

Fig. 2. Serpulids from United States fouling plates. Crucigera websteri. A. Operculum, juvenile from Humboldt Bay, California. B. Operculum, adult from San Pedro, California (LACMNH-N8819). – C. zygophora. C. Operculum, juvenile from Alaska (SERC). D. Operculum, adult from Canoe Bay, Alaska (LACMNH-N2128). – Ficopomatus enigmaticus. E. Tubes from Lake Merritt, California (LACMNH-N5141). F. Body, from Chesapeake Bay, Virginia (SERC-59327). G. Operculum, from Chesapeake Bay, Virginia (SERC-60530R). H. Colonies in Long Beach, California (photo by Bruno Pernet).– F. miamiensis. I. Tubes, from Chetumal Bay, Mexican Caribbean (ECOSUR). J–K. Operculum, from Galveston Bay, Texas (SERC-88344RF). – F. uschakovi. L. Tube, from Corpus Christi, Texas (SERC-88883). M. Thorax and operculum, from Galveston Bay, Texas (SERC-86995). N. Operculum, from La Encrucijada, Chiapas (UMAR-Poly 113). O. Thorax and operculum, from Corpus Christi, Texas (SERC-88883).

Fig. 7. Serpulids from United States fouling plates. Pomatostegus stellatus. A. Operculum, from Biscayne Bay, Florida (SERC-118825). – Protula balboensis. B–D. Tube, body and branchial crown, white arrow shows detail of rounded processes, Biscayne Bay, Florida (SERC-118825). E. Body of live adult from Smithsonian Institution station in Bocas del Toro, Panama (photo by Betel Martínez- Guerrero). – Pseudochitinopoma occidentalis. F. Operculum, Puget Sound, Washington (SERC- 33385R). – Pseudovermilia occidentalis. G. Operculum, Biscayne Bay, Florida (SERC-118450). – Salmacina huxleyi. H–I. Tubes, body and collar chaetae, Biscayne Bay, Florida (SERC-118449). – S. tribranchiata. J. Body, Galapagos Island (SERC-233687).

Fig. 4. Serpulids from United States fouling plates. Hydroides bispinosa. A. Operculum, from Biscayne Bay, Florida (SERC-118756). – H. cf. brachyacantha. B. Operculum, Pensacola Bay, Florida (SERC- 92415). – H. dianthus. C. Operculum, Corpus Christi, Texas (SERC-87878RB). – H. dirampha. D. Operculum, Biscayne Bay, Florida (SERC-118709). – H. elegans. E. Operculum, San Diego, California (SERC-04075R). – H. floridana. F. Operculum, Corpus Christi, Texas (SERC-87968). – H. gracilis. G. Operculum, San Diego, California (SERC-34585R). – H. longispinosa. H. Operculum, Oahu, Hawaii (SERC-19654). – H. parva. I. Operculum, Biscayne Bay, Florida (SERC-118709). – H. sanctaecrucis. J. Operculum, Tampa Bay, Florida (SERC-66920R).