Axinella infundibuliformis ( 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 : 38-42

Axinella infundibuliformis ( Linnaeus, 1759)

Figs 21A–J , 23 , 24

Spongia infundibuliformis Linnaeus, 1759: 1348 , sp. 9; Linnaeus 1767: 1296, sp. 3 (in part);? Houttuyn 1772: 432; Gmelin in Linnaeus 1791: 3818, sp. 3 (in part); Müller 1776: 256;? Olivi 1792: 262; Esper 1791a: pl. XI, 1794: 205 (text, in part); Bosc 1802: 140 (in part); Gray 1821: 359.

Spongia crateriformis Pallas, 1766: 386 , sp. 233 (also Boddaert 1767: 486 and Wilkens 1787: 222) (not: Spongia foliascens Pallas, 1766: 395 , sp. 246).

Spongia caliciformis Esper, 1797: 203 (text), pl. LVII (not: sensu Lamarck 1814: 432 and 1816: 365, as calyciformis = Axinella arctica according to Topsent 1932).

Spongia pocillum ; Lamouroux 1816: 45 (not: Müller 1776: 256 = Grantia compressa )

Halichondria infundibuliformis ; Fleming 1828: 524; Johnston 1842: 105, pl. VI fig. 3.

Isodictya infundibuliformis ; Bowerbank 1866: 317; 1874: 137, pl. LIV (not: Tanita & Hoshino 1989: 131, pl. 13 fig. 4, text-fig. 81 = Axinella sp. ).

Isodictya pocillum Bowerbank, 1866: 305 ; 1874: 135, pl. LIII figs 4–6.

Tragosia infundibuliformis ; Gray 1867: 513.

Axinella infundibuliformis ; Burton 1959: 48; Ackers et al. 1992: 121; Alvarez & Hooper 2002: 730.

Original description: ‘ Spongia turbinata cava’ (i.e. a hollow funnel-shaped sponge).

Type material: Unknown. I propose as neotype BELUM Mc 4438, United Kingdom, Scotland, Firth of Lorn, Loch Caolisport, 2.3 nm SW of Point of Knap, 55.853383°N 5.7182°W, depth 34.5 m, 24 June 2008, coll. B. Picton). The collecting locality is sufficiently close to Norwegian waters considered to be the type locality .

Molecular sequences: neotype sequences were published by Morrow et al. (2012).

Remarks: The original description was copied from Linnaeus (1753b), where there are two references, C. Bauhinus’ Pinax (1623: 368) described as ‘ Spongia forma infundibuli’ (i.e. sponge with funnel form) and Clusius’ Exoticorum (1605: 125) described as ‘ Spongia elegans’. The habitat was stated in Linnaeus (1753b) to be ‘Europa australi’, which would be Southern Europe or the Mediterranean. The C. Bauhinus reference does not provide an image, but the one in Clusius has an image of his ‘ Spongia elegans’, here reproduced as Fig. 21A . Remarkably, this image was repeated in J. Bauhinus (1651: 815) as ‘ Spongia infundibula imitata’.

Pallas (1766: 386, sp. 233), followed by his translators Boddaert (1767) and Wilkens (1787), replaced Linnaeus’ name by Spongia crateriformis , without explanation, but citing Linnaeus’ (1759, ‘Sp. infundibuliformis’), and C. Bauhinus. In addition, he also cited Mercati’s (1541: 95–96, pl. 56, ‘Alcyonium primum antiquorum’, here reproduced as Fig. 21B ). In his descriptive text Pallas referred to Antigua (Caribbean island nation) and to the beach at Sussex, England, as locations from which specimens were examined by him, and he stated that the species probably originated in the ‘American Sea’. Pallas caused confusion by citing one of the Linnaeus references for Spongia infundibuliformis (Clusius’ ‘ Spongia elegans’) under his species Spongia foliascens (cf. Pallas 1766: 395, sp. 246), along with some additional references [ Petiver 1712: pl. 19 fig. 4; Rumphius 1750: 254, pl. 90 fig. 1 (here reproduced in Fig. 21C ); Hebenstreit (1743); and Worm (1755)]. Spongia foliascens (currently known as Phyllospongia foliascens ), was stated to be from the Indian Ocean.

This confusion led Linnaeus (1767: 1296–1297) to assign Pallas’ S. foliascens to the synonymy of his Spongia infundibuliformis , which is obviously erroneous. Linnaeus (1767) did not list Spongia crateriformis , but Gmelin in Linnaeus (1791: 3818) added Spongia crateriformis to the synonymy of Spongia infundibuliformis , which partially corrected Pallas’ unexplained ignorance of the name infundibuliformis in favour of his new name crateriformis , ignored by Linnaeus (1767). Still, the list of references in Gmelin in Linnaeus (1791) contained a mixture of at least two species, Spongia infundibuliformis Linnaeus and Spongia foliascens Pallas , which was recognized by later authors (e.g. Esper 1794: 207).

Linnaeus (1767) added an apparently important reference, Gunnerus (1768: 78, pl. IV fig. 5, here reproduced in Fig. 21D ), which gave for the first time a concrete locality (Norwegian Sea) to pinpoint the occurrence of the species Spongia infundibuliformis and its eventual affiliation as Axinella infundibuliformis . Linnaeus apparently had a preview of Gunnerus’ work, because the reference pre-dated the latter. The definition for the species was slightly changed to ‘ Spongia infundibuliformis turbinata flexilis’ (emphasizing the cup wall being flexible). Gmelin in Linnaeus (1791) employed the same definition and added additional references, viz. Petiver (1712: ‘ Spongia infundibuliformis’, pl. 19 fig. 6, here reproduced in Fig. 21E ). Please note that Petiver’s Pl. 19 fig. 4 (cf. above) is clearly Phyllospongia foliascens , labeled ‘ Spongia foliata aspera’ by Petiver). The last reference is J. Bauhinus’ ‘Planta marina retiformis’ (1651: 808, fig. XLVI, here reproduced in Fig. 21F ).

Esper (1791a, 1794) described and illustrated Spongia infundibuliformis (see his plate XI reproduced here as Fig. 21G ) with all the above cited references, including Pallas’ S. crateriformis and S. foliascens as synonyms, and additionally Hebenstreit’s Museum Richterianum 1743: 384, pl. XIV fig. 1 (labeled as ‘ Spongia infundibula forma C. Bauhinus’, here reproduced in Fig. 21H , looking much like the Caribbean species Niphates digitalis ). In his ‘Fortsetzungen’ (1797) Esper separated the Norwegian (and Icelandic) records from S. infundibuliformis and for these he erected a separate species Spongia caliciformis named infundibuliformis var. in his plate LVII, here reproduced in Fig. 21I . In this separation of two species from the complement of references by various authors, he failed to pick the correct species, viz. Spongia infundibuliformis (European records, some of which were initially labeled as ‘American Sea’) and Spongia foliascens (Indian Ocean records). This makes his Spongia caliciformis a clear junior synonym of S. infundibuliformis .

Olivi 1792: 262 cited the definition of Linnaeus (1767) and reported occurrence in the northern Adriatic in deep water and specimens were only a quarter the size of reported specimens. It is unclear whether this record concerns the present species.

In conclusion, it is clear that many early records of the species concern different species, of which only the Norwegian are reliably recognized as S. infundibuliformis Linnaeus, 1759 . Several early images assigned to this species concern S. foliascens . The ‘American’ and ‘Southern Europe’ specimens are either unreliably labeled specimens of unknown origin or funnel-shaped specimens of other species, e.g. Spongia lamella Schulze, 1879 , Axinella shoemakeri De Laubenfels, 1936 , Niphates digitalis ( Lamarck, 1814) or Cribrochalina infundibulum Schmidt, 1870 . This makes it necessary to choose a neotype to confirm the properties of the present species, as is done above.

Species diagnosis: (after Ackers et al. 1992: 121, editors B. Picton & D. Moss). Cup-shaped (or lamellate in case of incomplete growth), with a short stalk and flaring thin wall (3–4 mm) ending in a rounded edge, width up to 10 cm. Small oscules are scattered over the inner surface of the cup. Consistency firm. Colour buff or creamy yellow. Skeleton plumoreticulate, with brushes of spicules at the surface, the choanosome shows a vague axial condensation. Spicules are styles (260–465 x 13–15 µm) and oxeas (210–312 x 13–15 µm) in a variable proportion, tracts of primary lines mostly contain styles, interconnecting spicules are mostly oxeas. Microscleres are in the form of trichodragmata (up to 15 µm long), their occurrence is often obscured. Difficult to find.

An in situ image of the neotype is presented in Fig. 22A .

The species is the type of genus Tragosia Gray, 1867 (designation by Alvarez & Hooper 2002), currently a junior synonym of Axinella Schmidt, 1862 .

Distribution ( Fig. 23B ). Northwest Europe (Celtic Seas, North Sea, Norwegian waters, sublittoral down to 630 m), possibly the Adriatic but this needs confirmation.

Figures

FIGURE 21. Images assigned to Axinella infundibuliformis by various authors (originally as Spongia). A. Clusius’ ‘Spongia elegans’ (1605: Lib. VI, p. 125); B. Mercati’s ‘Alcyonium primum antiquorum’ (1541: 95–96, pl. 56); C. Rumphius’ ‘Spongia infundibula forma Tsjatsjoran laut’ (1750: 254, pl. XC fig. 1); D. Gunnerus’ ‘Spongia infundibuliformis’ (1768: 78, pl. IV fig. 5); E. Petiver’s ‘Spongia infundibuliformis’ (pl. 19 fig. 6); F. J. Bauhinus’ ‘Planta marina retiformis’ (1651: 808, fig. XLVI); G. Esper’s Spongia infundibuliformis / Spongia crateriformis (1791a: pl. XI); H. Hebenstreit’s ‘Spongia infundibula forma C. Bauhinus’ (1743: pl. XIV fig. 1); I. Esper’s Spongia caliciformis (as S. infundibuliformis varietas, 1797: pl. LVII).

FIGURE 22. Axinella infundibuliformis (Linnaeus 1759 as Spongia). A. in situ image of neotype BELUM Mc44238, United Kingdom, Scotland, Firth of Lorn, Loch Caolisport, 2.3 nm SW of Point of Knap, 55.853383°N 5.7182°W, depth 34.5 m, 24 June 2008, photo and collection C. Morrow and B. Picton; B. distribution of Axinella infundibuliformis (L.), dots are WPD data, squares are OBIS data (image downloaded from the World Porifera Database, cf. De Voogd et al. 2023).

FIGURE 23. Images assigned to Spongilla lacustris by various authors. (originally as Spongia). A. lectotype LH 1295.2 designated by Manconi & Pronzato 2000: 90, photo Linnean Herbarium, https://linnean-online.org/13967/#?s=0&cv=0; B. paralectotype LH 1295.1 designated by Manconi & Pronzato 2000: 90, photo Linnean Herbarium, https://linnean-online.org/ 13966/#?s=0&cv=0; C. Esper’s Spongia lacustris (1791: pl. XXIII); D. Esper’s Spongia lacustris varietas (1794: pl. XXIIIA), which may represent Spongia fluviatilis.

FIGURE 24. Spongilla lacustris. A. in situ image from Oegstgeest, The Netherlands, 52.17329°N 4.450464°E (photo N.J. de Voogd); B. Distribution of Spongilla lacustris (L.), image downloaded from the World Porifera Database, cf. De Voogd et al. 2023 (only an outline of the actual distribution).