First inventory of sea anemones (Cnidaria: Actiniaria) from La Paz Bay, southern Gulf of California (Mexico) Author Barragán, Yamaly Author Sánchez, Carlos Author Rodríguez, Estefanía text Zootaxa 2019 2019-02-21 4559 3 501 549 journal article 27440 10.11646/zootaxa.4559.3.4 73a6e2e6-fbe3-4f6f-b946-0636fe6fa234 1175-5326 2627177 D3C170C0-D928-4564-8BD9-447D84E141A0 Phymactis papillosa ( Lesson, 1830 ) ( FIgS. 2 , 18–20 , TAble 8 ) Actinia papillosa Lesson, 1830 Actinia clematis Drayton in Dana, 1846 Bunodes papillosa Verrill, 1869 Bunodes papillosus Lesson, 1830 : Andres 1883 ( 1884 ) Bunodactis papillosa (Verrill, 1849) : Carlgren 1949 Cereus papillosus: Milne-Edwards, 1857 Phymactis clematis Milne-Edwards & Haime, 1851 : Verrill, 1869 Phymactis florida: Milne-Edwards 1857 Phymactis papillosa Lesson, 1830 : Stephenson, 1922 Phymactis papillosa var. cyanea : Häussermann, 2004 Phymactis papillosa var. fusca : Häussermann, 2004 Phymactis papillosa var. rubra : Häussermann, 2004 Phymactis papillosa var. rubra-cyanea : Häussermann, 2004 Phymactis papillosa var. rubra-fusca : Häussermann, 2004 Phymactis papillosa var. rubra-viridis : Häussermann, 2004 Phymactis papillosa var. viridis : Häussermann, 2004 Phymactis papillosa var. viridis-fusca : Häussermann, 2004 Non Phymactis clematis: Milne-Edwards 1857 ;? Stephenson 1918 ;? Orlando Muñoz et al. 1976 ; Zamponi 1977 , 1981 , 1989 , 1993 , 2000; Pollero 1983 ; Patronelli et al. 1987 ; Excoffon & Zamponi 1991 , 1997 , 2000 ; Acuña & Zamponi 1995 ; Genzano et al. 1996 ; Acuña & Zamponi 1996; Zamponi & Perez 1996; Acuña et al . 1996; Zamponi et al . 1997 (1998); Gomes et al . 1998 ; Excoffon et al . 1999 ; Zamponi et al . 1998 Rivetia papillosa Lesson, 1830 : Pax 1912 ? Phymactis capensis: Pax 1912 ? Phlyctenactis tuberculosa ( Quoy & Gaimard, 1833 ) : Zamponi & Excoffon 1992 (1995) Material eXamined. (See TAble 2 ). Description. EXTeRNAl ANATOmY ( FIg. 18 ): Well deVelOped pedAl dISC, TO 20 mm dIAmeTeR IN pReSeRVed SpeCImeNS. COlUmN CYlINdRICAl, TO 12 mm heIghT ANd TO 19 mm dIAmeTeR IN pReSeRVed SpeCImeNS, deNSelY COVeRed WITh SImple OR COmpOUNd VeSICleS fROm dISTAl mARgIN TO pedAl dISC. EACh mARgINAl pROjeCTION WITh AN ACRORhAgUS WITh bASITRIChS ANd hOlOTRIChS. Deep fOSSe. FUllY eXpANded ORAl dISC ANd TeNTACleS TO 19 mm dIAmeTeR, ORAl dISC TO 14 mm dIAmeTeR IN pReSeRVed SpeCImeNS. COUNTed Up TO 180 TeNTACleS, heXAmeROUSlY ARRANged IN SIX CYCleS, TO fIVe mm leNgTh IN pReSeRVed SpeCImeNS, SImple, ShORT, STICkY, SmOOTh, All Of SImIlAR leNgTh. MeSeNTeRIAl INSeRTIONS VISIble IN ORAl dISC. INTeRNAl ANATOmY ANd mICROANATOmY ( FIg. 19 ): MeSeNTeRIeS heXAmeROUSlY ARRANged IN Up TO SeVeN CYCleS: fIRST TO fOURTh CYCleS peRfeCT; OTheRS ImpeRfeCT ANd INCOmpleTe. EXAmINed SpeCImeNS STeRIle. ThRee SIphONOglYphS ATTAChed TO NON-dIReCTIVe meSeNTeRIeS. SAme NUmbeR Of meSeNTeRIeS dISTAllY ANd pROXImAllY. ReTRACTOR mUSCleS STRONg ANd dIffUSe, OCCUpYINg OVeR hAlf-leNgTh Of meSeNTeRIeS. PARIeTObASIlAR mUSCleS WITh RelATIVelY STRONg bUT ShORT fRee meSOgleAl lAmellA. BASIlAR mUSCleS Well deVelOped, WITh ShORT ANd ThIN meSOgleAl pROCeSSeS. MARgINAl SphINCTeR mUSCle dIffUSe. LONgITUdINAl mUSCleS Of TeNTACleS ANd ORAl dISC eCTOdeRmAl. FIGURE 18. External anatomy of Phymactis papillosa (Lesson, 1830) . A) Lateral view of preserved specimen. B) Detail of acrorhagi. C) Oral view of living specimen. D) lateral view of living specimen. E) Oral view of living specimen. F) Aboral view of living specimen. Acrorhagi (Ac), Pedal disc (Pd), Vesicle (V). Scale bars: 5 mm. FIGURE 19. Internal anatomy of Phymactis papillosa (Lesson, 1830) . A) Longitudinal section of the distal column. B) Cross section of the column at the level of the actinopharynx showing the cycles of mesenteries (cycles marked by numbers). C) Detail of a cross section of column showing parietobasilar muscles. D) Cross section through a tentacle showing ectodermal longitudinal musculature. E) Detail of a longitudinal section of the column showing acrorhagi and marginal sphincter muscle. F) Detail of a cross section of the column at the level of the actinopharynx showing the retractor and parietobasilar muscles. G) Longitudinal section of proximal end showing basilar muscles. Acrorhagi (Ac), Basilar muscles (Ba), Ectoderm (Ec), Endoderm (En), Fosse (Fo), Mesoglea (Me), Parietobasilar muscles (Pa), Retractor muscles (Re), Siphonoglyphs (arrows), Sphincter (Sp), Vesicle (V), Tentacle (Te). Scale bars: A, C–G, 0.1 mm; B, 0.5 mm. FIGURE 20. Cnidae of Phymactis papillosa (Lesson, 1830) . A, C, E, F, G, I) Basitrichs. D) Holotrich. J) b -mastigophore. H, K) p -mastigophores A. L) p -mastigophore B1. B) Spirocyst. COlOR ( FIg. 18 ): IN lIVINg SpeCImeNS, pedAl dISC lIghT Red. COlUmN Red WITh dARk Red VeSICleS. ACRORhAgI beIge. ORAl dISC dARk Red WITh A dARk Red blOTCh SURROUNdINg The mOUTh. TeNTACleS dARk Red WITh blACk bASe. CNIdOm ( FIg. 20 ): BASITRIChS, hOlOTRIChS, b -mASTIgOphOReS, p -mASTIgOphOReS A ANd B1, ANd SpIROCYSTS. See TAble 8 fOR SIZe ANd dISTRIbUTION. TABLE 8. Size ranges and distribution of cnidae of Phymactis papillosa (Lesson, 1830) . N, Total number of capsules measured; M, Ratio of number of specimens in which each cnida was found to number of specimens examined. Sizes in micrometers. Letters in parenthesis correspond with images in Fig. 20. (1), named as atrichs in Carlgren (1951); (2) and (3) , named as microbasic amastigophore A and microbasic p -mastigophore B, respectively by Häussermann (2004).

Tissue	Cnidae	N	M	P. papillosa	P. papillosa
Present study	Häussermann (2004)
length	width	length	width
Tentacles	Basitrichs (A)	14	3/3	18.0–27.0	1.9–3.1	13.5–26.1	1.8–3.1
Spirocysts (B)	16	3/3	18.9–28.2	1.9–2.4	17.1–21.6	1.8–2.7
Acrorhagi	Basitrichs (C)	11	3/3	32.0–66.5	2.2–5.7	70.2–88.2	4.5–9.0
Holotrichs (D)1	12	3/3	60.1–81.6	1.8–2.5	77.4–115.2	2.3–3.6
Column	Basitrichs (E)	11	3/3	16.4–22.4	2.2–2.7	13.5–21.6.	1.2–2.7
Actinopharynx	Basitrichs I (F)	11	3/3	11.9–20.6	1.8–2.5	15.3–18.9	1.8–2.7
Basitrichs II (G)	10	3/3	23.8–34.5	2.1–3.0	24.3–32.4	2.3–3.2
p -mastigophores A (H)2	0 9	3/3	20.1–26.5	3.9–4.7	23.4–26.1	4.5–5.4
Filaments	Basitrichs (I)	10	3/3	10.4–13.7	1.8–2.2	13.5–17.1	1.8–2.7
b -mastigophores (J)	11	3/3	25.3–31.2	3.1–4.0	27.0–42.3	3.6–6.3
p -mastigophores A (K)2	11	3/3	17.9–22.3	3.9–5.0	21.6–25.2	4.5–5.4
p -mastigophores B1 (L)3	10	3/3	10.3–12.7	1.8–2.4	11.7–13.5	1.4–2.3

Geographic and bathymetric distribution. Phymactis papillosa IS fOUNd AlONg The PACIfIC OCeAN fROm ChIle TO MeXICO . IN MeXICO , CARlgReN (1951) RepORTed ThIS SpeCIeS AT CeRRITOS ANd PUeRTO ESCONdIdO. IT hAS beeN RepORTed fROm The INTeRTIdAl TO A depTh Of 16 meTeRS ( HäUSSeRmANN 2004 ). ThIS IS The fIRST ReCORd Of The SpeCIeS fOR TeCOlOTe; AddITIONAllY, The SpeCIeS WAS ObSeRVed bUT NOT COlleCTed IN The INTeRTIdAl (beTWeeN 0–5 meTeRS) IN CAleRITAS ( FIg. 1 ). Remarks. Of The SIX SpeCIeS Of Phymactis ACCORdINg TO DAlY & FAUTIN (2018) , ONlY fIVe RemAINed VAlId AfTeR HäUSSeRmANN (2004) SYNONYmIZed P. clematis WITh P. papillosa IN heR RedeSCRIpTION Of The lATTeR. Phymactis papillosa IS The ONlY SpeCIeS Of The geNUS RepORTed IN The EASTeRN PACIfIC ( HäUSSeRmANN 2004 ; GOmeS et al . 2012,). IT IS deSCRIbed AS hAVINg fOUR VARIeTIeS Of COlOR: VAR. rubra (Red) , VAR. viridis (gReeN), VAR. cyanea (blUe), ANd VAR. fusca (bROWN); ONlY TWO Of TheSe VARIeTIeS ( rubra ANd viridis ) hAVe beeN ObSeRVed IN MeXICO (GUlf Of CAlIfORNIA ANd MeXICAN PACIfIC) ( HäUSSeRmANN 2004 ). We ObSeRVed ONlY Red SpeCImeNS IN LA PAZ BAY (IN TeCOlOTe ANd CAleRITA, See FIg. 1 ). AlThOUgh We fOUNd SlIghT dIffeReNCeS IN CNIdAe—mOST Of The SIZe RANgeS Of The CNIdAe RepORTed IN ThIS STUdY ARe ShORTeR ThAN ThOSe IN pReVIOUS STUdIeS (See TAble 8 )— All OTheR feATUReS AgRee Well WITh The Re-deSCRIpTION Of P . papillosa bY HäUSSeRmANN (2004) . ACCORdINg TO HäUSSeRmANN (2004) , The SIZe RANge Of CNIdAe VARIeS ACCORdINg TO The SIZe Of The SpeCImeNS IN P. papillosa , SO We CONSIdeRed The dIffeReNCeS fOUNd TO be dUe TO INTRASpeCIfIC VARIATION. AddITIONAllY, We fOUNd dIffeReNCeS AmONg The TeRmINOlOgIeS USed fOR The CNIdAe bY The dIffeReNT AUThORS (dIffeReNCeS INdICATed IN TAble 8 ). BASed ON The ImAgeS Of The CNIdAe pROVIded bY HäUSSeRmANN (2004) ANd IN lIghT Of ReCeNT UlTRASTRUCTURe STUdIeS Of The CNIdAe (e.g. ÖSTmAN 2000 ; ÖSTmAN et al . 2010A; RefT 2012 ), We ThINk ThAT The p -mASTIgOphOReS Of ThIS SpeCIeS CORReSpONd TO p -mASTIgOphOReS A ANd B1 (See RemARkS ON CNIdAe TeRmINOlOgY IN MATeRIAl ANd MeThOdS). IN LA PAZ BAY, P. papillosa IS AbUNdANT IN The INTeRTIdAl ANd IN TIde pOOlS; IT IS fOUNd ATTAChed TO ROCkS.