Haliclona (Haliclona) oculata ( Linnaeus, 1759 )

Van Soest, Rob W. M., 2025, Typification of Porifera described in the 10 edition of Linnaeus’ Systema Naturae, volume II, 1759, Zootaxa 5638 (1), pp. 1-65 : 13-17

Haliclona (Haliclona) oculata ( Linnaeus, 1759)

Figs 6A–H , 7A–F

Spongia oculata Linnaeus, 1759: 1348 , sp. 2; Pallas 1766: 390, sp. 239 [also Boddaert 1768: 492 and Wilkens 1787: 227)];

Linnaeus 1767: 1298, sp. 9; Houttuyn 1772: 454, pl. CXXV fig. 4; Esper 1790: pl. I, 1794: 180 (text); Gmelin in Linnaeus

1791: 3820, sp. 9; Bosc 1802: 142 pl. 29 fig. 6; Lamarck 1814: 452 (redescription Topsent 1933: 3); Montagu 1814: 78, pl.

VI fig. 2; Lamouroux 1816: 73; Grant 1826: fig. 22. Tupha oculata ; Gray 1821: 354. Haliclona oculata ; Grant 1841: 310; Hartman 1958: 52, pls 7–10; De Weerdt 1986: 83, pl. I fig. 4, II fig. 3, V fig. 1, text-figs

1–2 (with full complement of synonyms). Halichondria oculata ; Johnston 1842: 94, pl. III figs 1–2. Chalina oculata ; Bowerbank 1864: 208, pl. XIII fig. 262; 1866: 361; 1874: pl. XVI; Topsent 1933: 3. Haliclona (Haliclona) oculata ; De Weerdt 2000: 6.

Original description:‘ Spongia tenax ramosissima erecta:ramis teretiusculis obtusis’(i.e. erect sponge with tenacious smooth and blunt-ending branches). Later editions of the Systema Naturae slightly adapted the definition, inspired by Pallas, to include that there are oscules along the branches and the consistency is softer and the colour yellowish.

Type material: So far not identified. I propose as neotype a recently collected specimen identified by De Weerdt (1986), ZMA Por. 06031 ( Figs 7A–B ), the Netherlands, Zeeland, Kanaal door Zuid-Beveland, connecting the Oosterschelde and the Westerschelde, 51.56°N 03.94°E, depth 4 m, coll. J.J. Vermeulen, November 1977 (for extensive description see also De Weerdt 1986: 86). The locality of the neotype specimen is not too far from the cited type locality ‘between Belgium and England’ ( ICZN art. 75.3.6).

Molecular sequence: COI obtained by Niels van der Windt for this study.

Remarks: Linnaeus’ original description is based on pre-1758 works. Linnaeus (1737b: 480) already gave the definition for the species cited above, followed by Van Royen (1740: 522) and Linnaeus (1753b: 1170). Van Royen cited ‘Spongiosus fucus’ of C. Bauhinus’ Pinax (1651: 369), and Boerhaave’s ‘ Spongia ramosissima oculata’ (1727: 8), but he added a note that Bauhinus’ record is confusing as different species may be indicated by this author. Linnaeus (1753b) repeated the references of Van Royen and C. Bauhinus and added that the habitat is Italia. Although the species as we know it now is not often cited from the Mediterranean, there are recent records from Tunisia ( Ben Mustapha et al. 2003) and Morocco ( Krikech et al. 2020). Still, Linnaeus’ distribution record is dubious because the images of the species in pre-1759 literature are few and some are not certainly representing the species.

Pallas (1766: 390) cited all the references of Linnaeus (1737b, 1753b, 1759) and also Van Royen and C. Bauhinus. Further references of Pallas are Parkinson (1640: 1304, fig. 3) as ‘ Spongia ramosa brittannica’ (from the S coast of Devon, Isle of Portland, here reproduced as Fig. 6A ); Boccone (1697: 258, ‘ Spongia ramosa , oculata, viridis, frutescens’, fig. p. 116); Ray (1686: 81, as ‘ Spongia ramosa’); Ruysch (1710: pl. V fig. 1, as ‘Spongioides ramosissima marina oculata’ here reproduced as Fig. 6B ); Pontoppidan (1752: 258, figs B–E, possibly drawings representing Norwegian specimens); and Ellis (1755: 80, pl. XXXII figs F, f and g, here reproduced as Fig. 6C , with convincing description (not well-illustrated) of small holes along the branches). The last reference of Pallas is Seba (1758: vol. III pl. 97 figs 5–7, convincing images of ‘ Spongia erecta ramosa mollis atque subtilis’, from Italia, Netherlands and England, one of which is here represented by Fig. 6D ). Pallas changed the definition to: ‘ Spongia ramosissima mollis tenera flavescens, ramis tertiusculis, poris sparsis prominulis’ (changing ‘tenacious’ to ‘soft and tender’, added ‘yellowish’, and adding that there are sparse elevated pores), and he also gave a more northern locality for the species: ‘Mare inter Angliam & Belgium’.

Linnaeus (1767: 1298) cited his own and all of Pallas’ references, but with definition slightly expanded from his earlier text, adding that the branches are ‘foraminulata’ assuming that he meant they have oscules. He also took over the locality as ‘Mare Brittanico’.

Houttuyn (1772: 454, pl. CXXXV fig. 4, here reproduced as Fig. 6E ) confirmed shape and habitat.

Gmelin in Linnaeus (1791: 3820) with the same definition as Linnaeus (1767) and all the above references, cited in addition Ellis & Solander (1786: 184), and at the end added that the sponge is ‘flavescens’ (yellowish), with height 5–10 inches.

Esper (1790: pl. I, here reproduced in Fig. 6F , and 1794: 180, text), next to the above references, added Gevers (1787: 530); Bosc (1802: 142), listed only Ellis (1755), Boccone (1697), and Seba (1758), citing the habitat as northern and southern seas of Europe; Lamouroux (1816: 73–74) cited most of the above listed references; Montagu (1814: 78, pl. V fig. 2, here reproduced in Fig. 6G ) and Grant (1826: fig. 22, here reproduced in Fig. 6H ) confirmed the species characters.

De Weerdt (1986: 83; 2002: 865), still assuming Pallas to be the author of the species, reported possible dried type specimens in the Natural History Museum, London from a collection dated as 1861, obtained from ‘Mr. Koenig’, reg.nr. BMNH 1841.1.13.46, labeled probably by Ridley as ‘ Halichondria oculata’ or ’ Spongia oculata’ with note ‘History to be investigated’. There are no Pallas specimens in the Natural History Museum, but the collections of Linnaeus, part of which (the Clifford collection) was obtained by the Natural History Museum, did have contributions from Iceland, Madeira, Cape of Good Hope and India made by a person named Koenig (see Jackson 1888: 21). This makes it possible that the specimens mentioned by De Weerdt could be part of the collection available to Linnaeus before 1759. However, she is not certain about the status of this material, as she used question marks, e.g. ‘TYPE?’, or ‘?Holotype’. In her list of examined material she listed a ZMUC specimen labeled Spongia dichotoma Linnaeus (1767) from Newport, Rhode Island, stating that it ‘probably was Linnaeus’ original specimen’. Since Spongia dichotoma has been assigned to be the type species of the poecilosclerid genus Antho Gray, 1867 , the implications of her remarks are unclear, but more importantly, she did not propose a formal lectotype or neotype status for any of the specimens discussed by her, whereas she did so in other cases of Haliclona species treated in her paper. The online database of the Clifford collection in the Natural History Museum (https://doi.org/10.5519/ qd.5r3rj7zr) does not have entries of the four Spongia species including Spongia oculata listed in Linnaeus’ (1737b) ‘Hortus Cliffortianus’. I conclude that the status of these dried ‘Koenig’ specimens of uncertain origin is undecided. To remove this uncertainty and also to facilitate thus far unsuccessful attempts at integration of morphological and molecular classification of the Haplosclerida , I assigned the neotype cited above.

Species diagnosis: (after De Weerdt 1986: 83, pl. I fig. 4, II fig. 3. V fig. 1, text-figs 1–2, with comprehensive synonymy). Branching from a short stalk, height up to 30 cm, branches 1–1.5 cm in diameter, rounded at the top, rounded or flattened in outline, usually with a row of slightly elevated oscules of 1–3 mm in size. Branches may fuse to become flabelliform or may be isolated along most of their length. Surface smooth, velvety. Consistency soft but elastic. Colour light brown, with yellowish, greenish or reddish tinges. Skeleton a regular ectosomal uni- and choanosomal paucispicular reticulum of short fat oxeas. Parallel primary lines are connected by single spicules, with nodal spongin. No special ectosomal tangential skeleton. A peculiar feature is the flattened holdfast with a cluster of yellow non-functional gemmules of 0.5–1 mm.

This species is the type of the genus Haliclona Grant, 1841 , one of the largest genera of the Porifera with a current estimate of 480 accepted species ( De Voogd et al. 2023). Images of the neotype specimen is provided in Figs 7A–B , to which I add three in situ images from various parts of its range, SW Netherlands, NW France, and NE coast of the USA ( Figs 7C–E ).

Distribution ( Fig. 7F ): Northeast Atlantic, Arctic and Northwest Atlantic, cf. de Voogd et al. 2023. Some doubt exists over the Siberian Arctic occurrence, as these specimens were originally assigned to Haliclona gracilis ( Miklucho-Maclay, 1870) by Koltun (1959) rather than to Haliclona oculata .

Figures

FIGURE 6. Images assigned to Haliclona (Haliclona) oculata by various authors (originally as Spongia). A. Parkinson’s ‘Spongia ramosa brittannica’ (1640: 1304, fig. 3); B. Ruysch’ ‘Spongioides ramosissima marina oculata’ (1710: pl. V fig. 1); C. Ellis’ ‘Spongia ramosa britannica Park.’ (1755: 80, pl. XXXII figs F, f and g); D. Seba’s ‘Spongia erecta ramosa mollis atque subtilis’, (1758: vol. III pl. 97 fig. 6, from the Netherlands); E. Houttuyn’s Spongia oculata ‘Oogjesspons’ (1772: 452, pl. CXXXV fig. 4); F. Esper’s Spongia oculata (1790: pl. I figs 1–2); G. Montagu’s Spongia oculata (1814: Pl. VI fig. 2); H. Grant’s Spongia oculata (1826: fig. 22).

FIGURE 7. Haliclona (Haliclona) oculata (Linnaeus, 1759 as Spongia). A. Neotype specimen ZMA Por. 06031, designated herein, from S.W. Netherlands; B. label of neotype specimen with locality data and live colour; C–E, in situ images from various localities throughout the distribution range, C. in situ image of specimen ZMA Por. 20937 from Burgsluis, S.W. Netherlands (close to the locality from which the neotype was collected, photo P. van Bragt); D. in situ image of specimen ZMA Por. 14131 from Roscoff, N.W. France (close to the locality of ZMA Por. 14116, which was sequenced by McCormack et al. 2002, photo M.J. de Kluijver); E. in situ photo of unregistered specimen from Stellwagen Bank, East coast of the U.S.A., photo courtesy of S. Fuller (cf. Valentine et al. 2001); F. distribution records of Haliclona (Haliclona) oculata (L.), dots are WPD data, squares are OBIS data (image downloaded from the World Porifera Database, cf. De Voogd et al. 2023).