Bunodosoma grande ( Verrill, 1869 )

Barragán, Yamaly, Sánchez, Carlos & Rodríguez, Estefanía, 2019, First inventory of sea anemones (Cnidaria: Actiniaria) from La Paz Bay, southern Gulf of California (Mexico), Zootaxa 4559 (3), pp. 501-549 : 517-521

Bunodosoma grande ( Verrill, 1869)

( FIgS. 2 , 9–11 , TAble 5)

Cladactis grandis Verrill, 1869

Alicia grandis ( Verrill, 1869) : Haddon & Shackleton 1893

Cystiactis grandis Duerden, 1897

Eucladactis grandis ( Verrill, 1869) : Verrill 1899

Phymactis grandis ( Verrill, 1869) : Stephenson 1922

Bunodosoma grandis ( Verrill, 1869) : Carlgren 1949

(?) Bunodosoma grande ( Verrill, 1869) : Fautin et al. 2007

Non Bunodosoma grande ( Verrill, 1869) in Verrill 1870: Fautin 2013

Material eXamined. (See TAble 2).

Description. EXTeRNAl ANATOmY ( FIg. 9 ): Well deVelOped pedAl dISC, TO 37 mm dIAmeTeR IN pReSeRVed SpeCImeNS. COlUmN CYlINdRICAl TO 38 mm heIghT ANd TO 41 mm dIAmeTeR IN pReSeRVed SpeCImeNS, deNSelY COVeRed WITh VeSICleS fROm dISTAl mARgIN TO pedAl dISC. AbUNdANT eNdOCOelIC mARgINAl pROjeCTIONS, eACh WITh AN ACRORhAgUS WITh hOlOTRIChS. DISTAl VeSICleS AbUNdANT ANd jUmbled. Deep fOSSe. FUllY eXpANded ORAl dISC ANd TeNTACleS TO 51 mm dIAmeTeR, ORAl dISC TO 37 mm dIAmeTeR IN pReSeRVed SpeCImeNS. Up TO 164 TeNTACleS, heXAmeROUSlY ARRANged IN SIX CYCleS, TO 12 mm leNgTh IN pReSeRVed SpeCImeNS, SImple, STICkY, SmOOTh, All Of SImIlAR leNgTh. MeSeNTeRIAl INSeRTIONS VISIble IN pedAl dISC ANd ORAl dISC.

INTeRNAl ANATOmY ANd mICROANATOmY ( FIg. 10 ): MeSeNTeRIeS heXAmeROUSlY ARRANged IN fIVe CYCleS: fIRST TO ThIRd CYCleS peRfeCT; ReST ImpeRfeCT. All meSeNTeRIeS feRTIle eXCepT dIReCTIVeS. GONOChORIC, OOCYTeS 0.09–0.12 mm IN dIAmeTeR IN SpeCImeNS COlleCTed IN JUlY. TWO SIphONOglYphS ATTAChed TO TWO pAIRS Of dIReCTIVe meSeNTeRIeS. SAme NUmbeR Of meSeNTeRIeS dISTAllY ANd pROXImAllY. ReTRACTOR mUSCleS STRONg ANd dIffUSe, OCCUpYINg OVeR hAlf-leNgTh Of meSeNTeRIeS. PARIeTObASIlAR mUSCleS WITh STRONg ANd ThICk fRee meSOgleAl lAmellA. BASIlAR mUSCleS Well deVelOped, WITh ShORT ANd ThICk meSOgleAl pROCeSSeS. MARgINAl SphINCTeR mUSCle CIRCUmSCRIbed. LONgITUdINAl mUSCleS Of TeNTACleS ANd ORAl dISC eCTOdeRmAl.

COlOR ( FIg. 9 ): IN lIVINg SpeCImeNS, pedAl dISC pINk. COlUmN OlIVe gReeN. DISTAl VeSICleS YellOWISh OR beIge; pROXImAl VeSICleS pINk, OlIVe OR pURple. ACRORhAgI beIge. ORAl dISC OlIVe OR bROWN. TeNTACleS bROWN OR ReddISh WITh beIge SpOTS IN ORAl SIde.

CNIdOm ( FIg. 11 ): BASITRIChS, hOlOTRIChS, b -mASTIgOphOReS, p -mASTIgOphOReS A ANd B1, ANd SpIROCYSTS. See TAble 5 fOR SIZe ANd dISTRIbUTION.

Geographic and bathymetric distribution. Bunodosoma grande IS fOUNd AlONg The PACIfIC OCeAN fROm PeRU, PANAmA, NICARAgUA ANd ECUAdOR beTWeeN 1–20 meTeRS ( VeRRIll 1869, 1870; FAUTIN et al. 2007). IT hAS beeN AlSO RepORTed IN COSTA RICA bY ACUñA et al. (2013), hOWeVeR We CONSIdeRed ThIS ReCORd dUbIOUS (See RemARkS belOW). IN AddITION, CONTRARY TO FAUTIN (2013) TheRe IS NOT meNTION Of B. grande OCCURRINg IN LA PAZ (MeXICO) bY VeRRIll (1870). ThIS IS The fIRST RepORT fOR MeXICO. IN LA PAZ BAY, IT WAS fOUNd IN PUNTA COYOTe NORTh 1, IN The INTeRTIdAl ZONe beTWeeN 1–5 meTeRS.

Remarks. VeRRIll (1869) deSCRIbed Bunodosoma grande fROm PeRU, PANAmA ANd NICARAgUA AS A NeW SpeCIeS bASed ONlY ON eXTeRNAl ChARACTeRISTICS. CARlgReN (1950) ReeXAmINed The mATeRIAl ANd INClUded dATA Of The SIZe RANgeS Of The CNIdAe; he STATed ThAT B. grande dOeS NOT ReSemble ANY OTheR SpeCIeS WIThIN The geNUS OR ANY SpeCIeS fROm The GUlf Of CAlIfORNIA. HOWeVeR, ShORTlY AfTeR, CARlgReN (1951) ReVISed The SpeCImeNS fROm The GUlf Of CAlIfORNIA ANd ReAlIZed TheRe WAS A SpeCIeS ReSemblINg B. grande ; he deSCRIbed IT AS A NeW SpeCIeS, B. californicum ( CARlgReN 1951). The ReCORd Of B. grande fROm ACUñA et al. (2013) IN The PACIfIC COAST Of COSTA RICA IS NOT RelIAble beCAUSe The AUThORS dId NOT pROVIde ANY deSCRIpTION, ImAgeS Of ANATOmY OR CNIdAe dATA. HOWeVeR, beCAUSe ThIS SpeCIeS hAS beeN SO fAR RepORTed fURTheR NORTh (MeXICO, pReSeNT STUdY) ANd SOUTh (ECUAdOR) AlONg The PACIfIC COAST, The SpeCIeS IS pRObAblY AlSO dISTRIbUTed IN COSTA RICA. NeVeRTheleSS, The RepORT bY ACUñA et al. (2013) hAS TO be CONfIRmed WITh RelIAble dATA fROm COSTA RICA, OR fROm ReeXAmINATION Of The SpeCImeNS TheY STUdIed.

TABLE 5. Si z e ran g es and distri bu tion of cnidae of Bunodosoma grande (V errill, 1 869). N, Total n u m b er of caps u les meas u red; M, Ratio of n u m b er of specimens in which each cnida was fo u nd to n u m b er of specimens examined. Si z es in micrometers. Letters in parenthesis correspond with ima g es in Fi g. 11. (1), C ate g ory named as atrichs in C arl g ren (1 950).

Tiss u e C nidae N M B. grande B. grande B. grande

P resent st u dy Fa u tin et al. C arl g ren (1 950)

(2 0 0 7)

len g th width len g th width len g th width

Tentacles Basitrichs I (A) 13 2 / 2 13. 6– 1 6 .3 1. 9– 2. 9 0 8. 5– 1 6 .2 1.2 – 3.2

Basitrichs II (B) 1 0 2 / 2 2 7 .2 – 31. 0 2. 5– 2. 8 1 9. 9– 32. 5 2.3 –4. 0 22. 6– 2 5. 4 3. 0– 3. 0

Spirocysts (C) 23 2 / 2 1 9. 7– 3 7. 5 1. 9–4. 0 1 4. 4– 3 8. 0 1. 8–4 .3

Acrorha g i H olotrichs (D) (1) 11 2 / 2 3 8. 0–47. 5 4 .2 –4. 8 3 5 .2 –45 .1 4 .2 –5. 0 C ol u mn Basitrichs I (E) 1 0 2 / 2 1 4. 0 – 2 0 .2 1. 9– 2. 6 1 4 .1 – 22.1 1. 8– 3. 8

Basitrichs II (F) 1 0 2 / 2 2 5. 0– 31.3 3.2 – 4. 4 22. 9– 3 0 .2 2. 4–4. 4

Actinopharynx Basitrichs I (G) 11 2 / 2 11. 8– 1 8. 4 1. 6– 2.1 0 8. 8– 1 4 .1 1.2 – 2. 0

Basitrichs II (H) 11 2 / 2 22. 9– 3 0 .1 2. 4– 3. 4 1 6. 6– 2 9. 7 2. 0–4. 4 2 6 .1 – 2 9. 6 3. 5–4. 0

p - masti g ophores A (I) 0 9 2 / 2 1 9. 6– 22. 8 4. 6–5 .3

Filaments Basitrichs I (J) 11 2 / 2 13. 0– 1 7. 8 1. 8– 2.2 11.2 – 1 7. 5 1. 7– 2. 9 13. 4– 1 7. 0 2. 8– 2. 8 Basitrichs II (K) 0 9 2 / 2 2 7. 4– 33. 5 2. 4– 3. 9 2 0. 4– 3 6 .3 2.2 –5. 5

b - masti g ophores (L) 0 7 2 / 2 3 0 .1 –4 3. 5 4. 8–6 .2

p - masti g ophores A (M) 1 0 2 / 2 2 0. 4– 2 5 .2 4. 5–5. 5 2 0 .1 – 32.3 4 .1 –5. 7 21.1 – 2 5. 4 4 .2 –4. 5

p - masti g ophores B1 (N) 1 4 2 / 2 1 4. 7– 1 7 .1 3.2 –4. 0

IN The pReSeNT STUdY, We ObSeRVed SeVeRAl dIffeReNCeS beTWeeN B. californicum ANd B. grande : 1) VeSICleS IN The COlUmN Of B. californicum ARe dISTINCTlY ARRANged IN lONgITUdINAl ROWS WheReAS IN B. grande The VeSICleS ARe mORe AbUNdANT, ANd eVeN IN RelAXed SpeCImeNS IT IS VeRY dIffICUlT TO dISTINgUISh Them AS ARRANged IN lONgITUdINAl ROWS ( FIgS. 6A, B ANd 9A, B , ReSpeCTIVelY); 2) dISTAl VeSICleS fORmINg The mARgINAl pROjeCTIONS Of B. californicum ARe NOT AS AbUNdANT AS IN B. grande , IN WhICh VeSICleS ARe SO AbUNdANT ThAT TheY COUld be SeeN AS A dISTAl COllAR ( FIgS. 9A, B ); TheSe TWO dIffeReNCeS AllOW dISTINgUIShINg bOTh SpeCIeS IN The fIeld; ANd 3) pARIeTObASIlAR mUSCleS IN B. californicum ARe WeAkeR ANd ThINNeR ThAN ThOSe IN B. grande , WhICh ARe STRONgeR ANd ThICkeR ( FIgS. 7F ANd 10F , ReSpeCTIVelY). CARlgReN (1950) meNTIONed NO CNIdAe IN The COlUmN, NO SpIROCYSTS IN The TeNTACleS, NO p - mASTIgOphOReS A IN The ACTINOphARYNX, ANd NO b -mASTIgOphOReS IN fIlAmeNTS. AlSO, The AUThOR USed The TeRm ATRIChS WhICh fITS VeRY Well WITh The SIZe Of hOlOTRIChS. ThIS lACk Of TYpeS Of CNIdAe IS dUe TO The bAd pReSeRVATION Of The SpeCImeNS The AUThOR ObSeRVed. HOWeVeR, The SIZeS RANgeS Of The CNIdAe he pROVIded fITS VeRY Well WITh The pReSeNT STUdY. The CNIdAe RepORTed fROm The SpeCImeNS STUdIed bY FAUTIN et al. (2007) IN The GAlApAgOS ISlANdS ARe SlIghTlY dIffeReNT fROm The ONeS RepORTed heRe. TheY fOUNd NO NemATOCYSTS IN The mARgINAl pROjeCTIONS, NO b -mASTIgOphOReS IN The fIlAmeNT, ANd ONlY lARgeR p -mASTIgOphOReS A IN The fIlAmeNTS (See TAble 5). AlThOUgh TheSe dIffeReNCeS COUld be dUe TO INTRASpeCIfIC VARIATION, We RATheR TAke ThIS ReCORd WITh CAUTION UNTIl IT CAN be fURTheR CONfIRmed.

Figures

FIGURE 2. Living specimens of A) Anthopleura dowii. B) Bunodosoma californicum. C) Bunodosoma grande. D) Isoaulactinia hespervolita. E) Phymactis papillosa. F) Phialoba steinbecki. G) Alicia beebei. H) Calliactis polypus.

FIGURE 6. External anatomy of Bunodosoma californicum Carlgren, 1951. A–B) Lateral view of preserved and living specimen, respectively. C) Detail of acrorhagi. D) Oral view of living specimen. E) Detail of distal column. Acrorhagi (Ac), Oral disc (Od), Pedal disc (Pd), Vesicle (V). Scale bars: 5 mm.

FIGURE 7. Internal anatomy of Bunodosoma californicum Carlgren, 1951. 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 longitudinal section of the column showing acrorhagi. D) Longitudinal section of proximal end showing basilar muscles. E) Cross section through a tentacle showing ectodermal longitudinal musculature. F) Detail of a cross section of the column at the level of the actinopharynx showing the retractor and parietobasilar muscles. Acrorhagi (Ac), Basilar muscles (Ba), Ectoderm (Ec), Endoderm (En), Fosse (Fo), Marginal projections (Mp), Mesoglea (Me), Parietobasilar muscles (Pa), Retractor muscles (Re), Siphonoglyphs (arrows), Sphincter (Sp), Vesicle (V). Scale bars: A, C–F, 0.1 mm; B, 0.5 mm.

FIGURE 9. External anatomy of Bunodosoma grande (Verrill, 1869). A–B) Lateral view of preserved and living specimen, respectively. C) Oral view of living specimen. D) Detail of acrorhagi. E) Specimen in situ. Acrorhagi (Ac), Oral disc (Od), Vesicle (V). Scale bar: 5 mm.

FIGURE 10. Internal anatomy of Bunodosoma grande (Verrill, 1869). 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) Cross section through a tentacle showing ectodermal longitudinal musculature. D) Longitudinal section of proximal end showing basilar muscles. E) Cross section of column showing gametogenic tissue. F) Cross section of column showing parietobasilar muscles. G) Detail of a cross section of the column at the level of the actinopharynx showing the retractor and parietobasilar muscles. Acrorhagi (Ac), Basilar muscles (Ba), Ectoderm (Ec), Endoderm (En), Fosse (Fo), Marginal projections (Mp), Mesoglea (Me), Oocytes (O) Parietobasilar muscles (Pa), Retractor muscles (Re), Siphonoglyphs (arrows), Sphincter (Sp), Vesicle (V). Scale bars: A, C–F, 0.1 mm; B, 0.5 mm.

FIGURE 11. Cnidae of Bunodosoma grande (Verrill, 1869). A, B, E, F, G, H, J, K) Basitrichs. D) Holotrichs. L) b- mastigophore. I, M) p-mastigophore A. N) p-mastigophore B1. C) Spirocyst.