Shallow Suberitida (Porifera, Demospongiae) from Peru Author Cóndor-Luján, Báslavi Universidad Científica del Sur, Facultad de Ciencias Veterinarias y Biológicas, Carrera de Biología Marina, Antigua Panamericana Sur Km 19, Villa El Salvador, Lima, Peru Author Arteaga, Alvaro 0000-0001-6001-8358 Universidad Científica del Sur, Facultad de Ciencias Veterinarias y Biológicas, Carrera de Biología Marina, Antigua Panamericana Sur Km 19, Villa El Salvador, Lima, Peru & aarteagabengoa 14 @ gmail. com; https: // orcid. org / 0000 - 0001 - 6001 - 8358 aarteagabengoa14@gmail.com Author Polo, Christian 0000-0001-6775-8156 Universidad Científica del Sur, Facultad de Ciencias Veterinarias y Biológicas, Carrera de Biología Marina, Antigua Panamericana Sur Km 19, Villa El Salvador, Lima, Peru & cpolobio @ gmail. com; https: // orcid. org / 0000 - 0001 - 6775 - 8156 cpolobio@gmail.com Author Arroyo, Yessenia 0000-0003-2477-7079 Universidad Científica del Sur, Facultad de Ciencias Veterinarias y Biológicas, Carrera de Biología Marina, Antigua Panamericana Sur Km 19, Villa El Salvador, Lima, Peru & ysabel 0396 @ gmail. com; https: // orcid. org / 0000 - 0003 - 2477 - 7079 ysabel0396@gmail.com Author Willenz, Philippe 0000-0003-4127-9346 Royal Belgian Institute of Natural Sciences, Taxonomy and Phylogeny, Rue Vautier 29, B- 1000, Bruxelles, Belgium & Université Libre de Bruxelles, Laboratoire de Biologie Marine, Avenue F. D. Roosevelt, 50, B- 1050, Bruxelles, Belgium philippe. willenz @ naturalsciences. be; https: // orcid. org / 0000 - 0003 - 4127 - 9346 philippe.willenz@naturalsciences.be Author Hajdu, Eduardo Universidade Federal do Rio de Janeiro, Museu Nacional, Departamento de Invertebrados, Quinta da Boa Vista, S / N, 20940 - 040, Rio de Janeiro, RJ, Brazil text Zootaxa 2023 2023-04-17 5264 4 451 489 http://dx.doi.org/10.11646/zootaxa.5264.4.1 journal article 10.11646/zootaxa.5264.4.1 1175-5326 7836930 675A2650-4738-4DDB-8970-1FE4307F6B3C Hymeniacidon perlevis ( Montagu, 1814 ) ( Figure 4 , Figure 5 , Table 4 ) Synonyms: (East Pacific, only) Hymeniacidon sinapium de Laubenfels, 1930: 26 , Desqueyroux-Faúndez and van Soest 1997: 432 , Hofknecht 1978: 54 . Hymeniacidon perlevis ( Montagu, 1814 ) : Turner 2020: 542 ; Harbo et al. 2021: 280 , Fig. 3 . Material examined. Twenty-two specimens. MNRJ 11252 , Chilca Islet , Pucusana , Lima ( 12°28′42.13″S 76°47′55.26″W ), 1 m depth , coll. E. Hajdu , Y. Hooker & Ph. Willenz , 20.IX.2007 . MNRJ 11288 , North of Colorada Inlet , Samanco Bay , Chimbote , Ancash ( 09°11′40.80′′S 78°23′21.00′′W ), 2.7 m depth , coll. E. Hajdu , Y. Hooker & Ph. Willenz , 24.IX.2007 . MNRJ 11430 , Mancora Beach “El Point”, Piura ( 04°06′21.00″S 81°03′31.30″W ), intertidal, coll. Y. Hooker & M. Ríos , 14.X.2007 . MNRJ 12084 , Hotel de Turistas Beach , Ilo , Moquegua ( 17°38′11.63″S 71°20′28.99″W ), intertidal, coll M. Ríos , 09.XI.2008 . MNRJ 12205 and MNRJ 12206 , La Vieja Island , Independencia Bay , Paracas , Ica ( 14°16′08.20″S 76°11′25.00″W ), 4.2 and 4.4 m depth , Y. Hooker , Ph. Willenz & F. Azevedo , 09.XII.2008 . MNRJ 12181 and MNRJ 12183 , Punta San Juan , San Juan de Marcona , Ica ( 15°22′03.23″S 75°11′18.41″W ), intertidal, coll. Y. Hooker , M. Vichez , Ph. Willenz, G. Lobo-Hajdu, F. Azevedo , 05.XII.2008 . MNRJ 12859 , Roquedal , Laguna Grande , Paracas , Ica ( 14°09′11.80″S 76°15′51.30″W ), 1.3 m depth , coll. E. Hajdu , Y. Hooker & Ph. Willenz , 12.XII.2008 . MNRJ 12886 , Islote , Atenas Beach , Paracas , Ica ( 13°49′38.71″S 76°18′07.41″W ), 1.6 m depth , coll. Y. Hooker , Ph. Willenz & N. Mostajo , 13.XII.2008 . UCSUR 07-000007 , San Lorenzo Island 1, El Callao , Lima ( 12°05′46.71″S 77°11′29.46″W ), 7 m depth , coll. L. Aguirre , XII. II.2010 . UCSUR 07-000013 , Las Ninfas Beach , Pucusana , Lima ( 12°28′49″S 76°48′3″W ), intertidal, coll. B. Moreno & A. Cuba , 27. VI .2010. UCSUR 07-000040 , Playa Hermosa , San Juan de Marcona , Ica ( 15°21′20″S 75°10′16″W ), intertidal, coll. J. Francia , 06.VIII.2018 . UCSUR 07-000043 , north side of Santa Rosa Island , Independencia Bay , Paracas , Ica ( 14°15′24″S 76°13′9″W ), subtidal, coll . R . Canales & B. Cóndor-Luján , 14.XI.2010 . UCSUR 07-000051 , Atenas Beach , Paracas , Ica ( 13°49′13.16″S 76°18′2.81″W ), 7 m depth , coll. K. Farfán , 01.II.2019 . UCSUR 07-000053 , Las Ninfas Beach , Pucusana , Lima ( 12°28′49″S 76°48′3″W ), intertidal, coll. A. Arteaga , Y.Arroyo & B. Cóndor-Luján , 19.II.2019 . UCSUR 07-000059 , Las Ninfas Beach , Pucusana , Lima ( 12°28′49″S 76°48′3″W ), intertidal, coll. M. Pita , 13.IV.2019 . UCSUR 07-000069 , UCSUR 07-000070 , UCSUR 07-000071 , UCSUR 07-000072 and UCSUR 07-000073 , Las Ninfas Beach , Pucusana , Lima ( 12°28′49″S 76°48′3″W ), intertidal, coll. C. Polo , 23.X.2019 . Description. Sponge with variable growth. Encrusting or cushion-shaped (up to 1 cm thick, Fig. 4A, B , 5A ). Bearing massive lobes (over 5 cm in height) or short digitiform fistules (up to 1 cm high, Fig. 5B ). Specimens reaching over 50 cm in maximum diameter, but more often up to 10 cm . Oscula (diam. = 1–4 mm ) on top of small elevations (average height = 6.5 mm ), with one or multiple openings ( Fig. 4A , 5B ). Surface rough, consistency slightly compressible. Colour. Orange ( Fig. 4A–B ) or yellow with a slightly orange tint ( Fig. 5A–B ) in life and dark mustard to beige in ethanol. Skeleton. Ectosomal,architecture tangential or paratangential,reticulated or resembling a parchment arrangement in tangential view ( Fig. 4C and 5C ), with subectosomal cavities (average diameter = 430 µm). Choanosomal architecture confused, with ascending tracts ending in bouquets when reaching the surface ( Fig. 4D and 5D ). FIGURE 4. Hymeniacidon perlevis ( Montagu, 1814 ) —orange. A–B, live specimens (MNRJ 12181 and MNRJ 12183); C, ectosomal skeleton in tangential ground section (UCSUR 07-000059); D skeleton architecture in transverse ground section (MNRJ 12084); E–F, styles (MNRJ 12084); G–H, details of E. Scale bars: A–B, approx. 2 cm; C, 200 µm; D, 500 µm; E–F, 100 µm; G–H, 20 µm. FIGURE 5. Hymeniacidon perlevis ( Montagu, 1814 ) —yellow. A –B, live specimens (MNRJ 12859 and MNRJ 11252); C, ectosomal skeleton in tangential ground section (UCSUR-07-000007); D, skeleton architecture in transverse ground section (MNRJ 12859); E–F, styles (MNRJ 12859); G–H, details of E. Scale bars: A, approx. 2 cm; B, 5 cm; C, 200 µm; D, 500 µm; E–F, 100 µm; G–H, 20 µm. Spicules. Megascleres. Styles to subtylostyles, smooth, straight or slightly curved, tyles small subterminal (102–443 x 2–13 µm, Fig. 4E–H and 5E–H , Table 4 ). Ecology. This species is very frequent in intertidal and shallow subtidal habitats and it can attain considerable coverage and biomass in certain localities (e.g Pucusana and northern Peru ). Specimens live attached to rocky substrates, near algae (e.g. Ulva lactuca ) and invertebrates such as anemones, gastropods, mussels, ophiuroids and cirripeds ( Gastaldi et al. 2016 ; this study). Geographic and bathymetrical distributions. Cosmopolitan. N Atlantic and Mediterranean Sea ( Montagu 1814 ; Topsent 1934 ). SW Atlantic: Argentina ( Cuartas 1985 ; Gastaldi et al. 2018 ). SE Atlantic: Namibia and South Africa ( Samaai et al. 2022 ). SW Indian: South Africa ( Samaai et al. 2022 ). NW Pacific: Japan and Korea ( Sim & Lee 2003 ; Hoshino et al. 2008 ). E Pacific: Canada , USA , Mexico and Galapagos Islands ( Laubenfels 1932 ; Hofknecht 1978 ; Desqueyroux-Faúndez & van Soest 1997 ; Turner 2020 ; Harbo et al. 2021 ). Along the Peruvian coast: from Mancora ( Piura , 04° S ) to Ilo ( Moquegua , 17° S , this study). MEOW in Peru : Guayaquil, Central Peru and Humboldtian ecoregions ( Spalding et al. 2007 ). Intertidal to subtidal ( 25 m depth, Samaai et al. 2022 ). Remarks. In the Eastern Pacific, eight valid Hymeniacidon species are known ( de Voogd et al. 2022 ): H. actites ( Ristau, 1978 ) , H. adreissiformis Dickinson, 1945 , H. calva ( Ridley, 1881 ) , H. corticata ( Thiele, 1905 ) , H. fernandezi Thiele, 1905 , H. longistylus Desqueyroux, 1972 , H. rubiginosa ( Thiele, 1905 ) and H. perlevis ( Montagu, 1814 ) . The Peruvian specimens match the descriptions of H. fernandezi , H. perlevis and H. rubiginosa (see Table 5 for details) but differ from the Chilean H. fernandezi and H. rubiginosa in the presence of subtylostyles, only described for H. perlevis ( Erpenbeck & van Soest 2002 ) . Considering this, we rather identify the Peruvian material as H. perlevis , which has been found in the NE Pacific (e.g. Turner 2020 , Harbo et al. 2021 ). TABLE 4. Spicule measurements of Hymeniacidon perlevis from Peru. Min=minimum, Max=maximum, SD=standard deviation. Number of spicules measured = 30.
Specimen Locality Length (μm) Width (μm)
Min Mean SD Max Min Mean SD Max
MNRJ 11252 Pucusana 123 283.1 94.3 411 4 8.4 2.4 12
MNRJ 11288 Chimbote 132 247.9 74.4 357 2 6.4 2.1 11
MNRJ 11430 Mancora 102 188.2 47.1 291 2 4.1 0.9 6
MNRJ 12084 Ilo 132 259.9 57.7 358 4 7.2 1.3 10
MNRJ 12205 Paracas 132 316.9 75.1 443 4 8.6 1.9 12
MNRJ 12206 Paracas 142 272.1 78.9 383 4 7.9 1.9 12
MNRJ 12181 Marcona 147 266.5 64.3 363 5 8.1 1.6 11
MNRJ 12183 Marcona 124 215.4 58.6 362 5 7.3 1.5 11
MNRJ 12859 Paracas 157 268.6 53.7 361 4 7.1 1.1 9
MNRJ 12886 Paracas 128 274.6 66.3 378 4 8.4 1.8 12
UCSUR 07-000007 San Lorenzo 150 267.1 58.8 350 3 6.7 1.7 10
UCSUR 07-000013 Pucusana 130 274.1 57.7 360 3 7.6 2.1 13
UCSUR 07-000040 Marcona 130 289.3 82.9 390 2 8.5 2.6 13
UCSUR 07-000043 Paracas 140 313.0 73.1 400 3 9.1 0.3 13
UCSUR 07-000051 Paracas 130 293.2 65.3 385 2 6.7 1.7 10
UCSUR 07-000053 Pucusana 140 283.9 67.5 370 2 6.8 1.9 10
UCSUR 07-000059 Pucusana 110 261.2 55.9 330 2 5.3 1 8
UCSUR 07-000069 Pucusana 140 288.7 75.0 390 5 8.4 2.2 13
UCSUR 07-000070 Pucusana 150 292.0 66.4 390 5 8.4 1.9 10
UCSUR 07-000071 Pucusana 150 301.3 49.3 370 3 7.6 1.4 10
UCSUR 07-000072 Pucusana 140 310.7 67.1 390 4 8.2 1.7 10
UCSUR 07-000073 Pucusana 140 294.3 52.1 360 2 6.0 1.8 10
TABLE 5. Comparative table of confirmed and alleged records of Hymeniacidon perlevis , H. fernandezi and H. rubiginosa , including locality and habitat, external morphology, skeleton organisation, and style size.
Species Locality and habitat External morphology Ectosome Choanosome Styles (length x width in µm)
Hymeniacidon perlevis
Montagu (1814) Devon Coast, England Apparently attached to a rock Yellow (alive) and light brown (dry) Numerous obtuse papillae on surface Scattered moderate-sized pores - - -
de Laubenfels (1932) , as H. sinapium Newport Bay, California Intertidal, abundant on rocks, in calm and warm waters Bright yellow with orange tints (alive) Encrusting to massive Oscula on raised processes Surface smooth with scattered low conuli Thin with subdermal brushes and lacunae Confused with tracts, mostly directed verti- cally and with points of the styles up 340 x 9
Hofknecht (1978) , as H. sinapium Gulf of California Inter- tidal and subtidal Abundant on rocks Bright orange (alive) and dark mustard (alcohol) Encrusting to massive Surface smooth with ridges Oscula over small elevations - - Slightly curved 350 x 10
Cuartas (1985) , as H. sanguinea Mar del Plata, Argentina Intertidal Associated with mussels Reddish orange (alive) Encrusting to massive with grooved surface Firm and homogeneous consistency Paratangential with subder- mal brushes Tracts ascending to surface Choanocyte chambers around inhalant canals Ectosomal: 240 x 5. Choanosomal (main): with two means in length 410 x 10 and 450 x 10
Desqueyroux-Faúndez & van Soest (1997) , as H. sinapium Galapagos Islands. Intertidal On barnacles Orange (formalin and ethanol) Thinly encrusting to low cushions Soft and surface smooth No apparent oscula Tangential, easily detacha- ble with subdermal lacunae Confused with large open spaces Tracts randomly oriented towards the surface Curved 148–237–312 x 4
......continued on the next page TABLE 5. (Continued)
Species Locality and habitat External morphology Ectosome Choanosome Styles (length x width in µm)
Erpenbeck & van Soest (2002) Tenby, Wales Intertidal to shallow subtidal On rocks, sand and Laminaria holdfasts Orange to blood-red (alive) and orange- brown (dry) Encrusting with short papillae and small digitations Consistency fragile and crumbly Paratangential with spicule brushes Confused with wispy bun- dles of spicules following a wavy course to surface Slightly curved often with subterminal tylotes 152–475 x 3–12
Gastaldi et al. (2018) San Antonio Bay, Ar- gentina Lower intertidal to upper subtidal Hard ledges, pebbles, and sandy/muddy bot- toms Orange-yellow, yellow, orange (alive) and dark orange-brown (ethanol) Cushion-shaped bearing irregular projec- tions Surface grooved Consistency soft Paratangential not so easily detachable Subectosomal canals Confused with longitu- dinal tracts ascending to surface Straight or slightly curved, some with bi- or trilobed bases 110–404 x 5–16
Turner (2020) California On rocks and sediments Yellow or yellowish-orange (alive) Irregular projections Paratangential or tangential with styles forming meshes on the surface Confused with tracts Curved and few with subterminal tyle 116–415 x 2–12
Harbo et al. (2021) Ladysmith Harbour, Brilliant orange with reddish tint (alive) Tangential with vague pali- Confused with plumose Moderately curved
Canada On ropes and hard substrate Fistular processes (10–20 cm) with apical oscula Surface smooth, micropapillate, micro- hispid sades of styles that penetrate tracts. the surface Larger canals Canals some with subtermi- nal tyle 135–399 x 3.5–13
Samaai et al. (2022) From Port Nolloth (Namibia) to Kosi Bay (South Africa) Intertidal On rocks Different orange patterns (alive) Thin or thickly encrusting to cushion Surface smooth and tuberculate or with digitate processes Oscula 0.5 – 1.5 mm Compressible. Tangential Confused with ill-defined tracts near to surface, without bouquets Large canals Scarce spongin Smooth, straight or slightly curved, thickest centrally 155–337 x 7
......continued on the next page TABLE 5. (Continued)
Species Locality and habitat External morphology Ectosome Choanosome Styles (length x width in µm)
this study Peru: 04–17 °S. Inter- tidal and subtidal Orange or yellowish (alive) and dark mustard to beige (ethanol) Reticulated (paratangential) or parchment (tangential) Confused with ascend- ing tracts forming or not Straight or slightly curved
On rocks Encrusting (thin to thick) Oscula on small elevations Subectosomal cavities bouquets Small subterminal tyle
(Slightly) rough and compressible 102–442.7 x 2–13
Hymeniacidon fernandezi
Thiele (1905) Juan Fernandez Island, Chile On Antipathes Vermillion (alive) and pale yellowish (preserved) Encrusting with wrinkles on surface Isolated oscula Meshes with spicules paral- lel to the surface (tangential) Presence of cavities 200–340 x 3–10
Burton (1934) Port Albemarle, Falkland Islands Depth: 18–100 m Subspherical sponge Grooved surface Subectosomal canals Ascending tracts
Desqueyroux (1972) Juan Fernandez Islands, Chile Depth: 10 m Yellowish-white (ethanol) Irregular, rugose and rough surface - Irregular network of tracts and independent spicules parallel to surface 300–400 x 7–10
Hymeniacidon rubiginosa
Thiele (1905) Iquique, Chile On rocks Reddish (alive) and yellowish (preserved) Shallow crusts with small elevations (1 mm), with small openings Paratangential Meshes. Confused Subectosomal cavities 325 x 7
Desqueyroux (1972) Iquique, Chile Depth: 0–970 m On rocks Gray-white encrusting Irregular and rugose surface Oscula on small elevations Paratangential with subecto- somal brushes Tracts with independent spicules around 300–420 x 4.7–16
Cuartas (1991) Mar del Plata, Argentina Intertidal Pink (alive) encrusting with grooved surface Oscula on the surface and small eleva- tions Elastic consistency Discontinuous tangential tracts Tracts of styles 270–390 x 5–9
It is important to mention that due to their high morphological similarities in colour (red to yellowish), encrusting growth form with oscula frequently located on small elevations, confused choanosomal skeleton arrangement, and styles measuring 200–400 µm x 3–16 µm, H. fernandezi and H. rubiginosa have been suggested as synonyms of H. perlevis ( de Laubenfels 1932 ; Erpenbeck & van Soest 2002 ; Burton 1932 ; Desqueyroux-Faúndez & van Soest 1997 ; Turner 2020 , Table 5 ) and it would not be surprising to confirm the synonymy of these three species in future studies.