Clathrina primordialis, (HAECKEL, 1872)

Klautau, Michelle & Valentine, Clare, 2003, Revision of the genus Clathrina (Porifera, Calcarea), Zoological Journal of the Linnean Society 139 (1), pp. 1-62 : 32-34

CLATHRINA PRIMORDIALIS ( HAECKEL, 1872)

Original name: Prosycum primordiale nomen nudum Haeckel, 1870, Ascetta primordialis Haeckel, 1872

Type locality: Lesina, Adriatic Sea.

Type: ZMB 1306 (suggested lectotype /alcohol). Naples , PMJ. Inv.Nr.Porif. 154 (type/alcohol). Lesina, Adriatic Sea .

Citations: Haeckel (1870); von Lendenfeld (1885, 1891); Carter (1886); Lackschewitsch (1886); Minchin (1896); Breitfuss (1898, 1932, 1935); Arnesen (1901); Jenkin (1908); Row (1909); Dendy & Row (1913); Ferrer-Hernandez (1918); Burton (1926, 1963); Brondsted (1931); Row & Hôzawa (1931); Arndt (1941); Tanita (1942, 1943); Borojevic (1971); Borojevic & Peixinho (1976); Klautau et al. (1994).

We received from PMJ (Inv.Nr.Porif. 154) a syntype collected from Lesina. Mixed with this specimen, which has only triactines, was a specimen of C. cerebrum .

Colour: The preserved specimen of A. (Clathrina) primordialis is white.

Description: Cormus formed of large, irregular and loosely anastomosed tubes. Skeleton comprises two kinds of equiangular and equiradiate triactines ( Fig. 26A ); some actines conical, others cylindrical. In both kinds, the tip of the actines is sharp.

Remarks: Haeckel (1870) introduced the species for the first time under the name Prosycum primordiale , from Naples, Italy. He did not give a description, simply presenting a systematic list of calcareous sponges. It was a nomen nudum. In his famous monograph of 1872 he provided a detailed morphological description of several calcareous species, including those he had previously cited in 1870, which were described as new. Consequently, the accepted date of many of these species, including C. primordialis , became 1872.

The ‘Natural System’ of classification proposed by Haeckel distinguished the genera of calcareous sponges by their spiculation. Therefore, as P. primordiale was formed only of triactines, he changed its genus to Ascetta and called it A. primordialis . In 1870 Haeckel designated Naples as the type locality. In 1872, he did not elect one type locality but instead listed several where A. primordialis could be found, concluding that this species was cosmopolitan. Nevertheless, he listed where he had found it for the first time and, strangely, he said that it was in Nizza in 1856 and then in Naples (1859) and several other localities, Lesina (Adriatic Sea) being the last one (1871).

Haeckel probably described A. primordialis as a series of different species and even genera. We can see this looking at some of his drawings and analysing some specimens identified by him. Consequently, his description was a generalization, based only on the presence of anastomosed tubes composing the cormus, and triactines with slightly conical or cylindrical actines near the centre and semifusiform at the distal part, always with a sharp tip (or tips). He elected four varieties of A. primordialis , all from Australia: var. dictyoides, var. loculosa, var. poterium and var. protogenes . Later, von Lendenfeld (1885) elevated two of those varieties to the rank of species ( A. loculosa and A. poterium ), and returned var. dictyoides ( Leucosolenia dictyoides Haeckel, 1870 ) to the specific level, as A. dictyoides . He did not mention protogenes , although Tanita (1942) considered a new species described by von Lendenfeld, 1885) ( A. procumbens ) to be a synonym of it. Dendy and Row (1913) again reinforced the position of these varieties as distinct species, putting all of them in the genus Leucosolenia . We had access to the holotype of only one of these varieties - poterium - and it is clearly an Ascaltis (based on the organization of the cormus, which is surrounded by a cortical membrane and missing a true atrium). In relation to the other ‘varieties’, we only had access to some of the specimens from Australia, and they are, indeed, distinct species.

In the original description of C. primordialis, Haeckel said that the actines of the triactines were slightly conical or cylindrical near the centre and semifusiform at the distal part, but always with sharp tips.

We also received another syntype under the name Prosycum primordiale . This came from Berlin ( ZMB 1306) and was collected in Naples. As the first mention of C. primordialis was under the name P. primordiale and the specimen was from Naples, the coincidence proved interesting. This sponge is beige when preserved and its cormus is formed of thin, irregular and loosely anastomosed tubes. Larger tubes project above the surface, their openings then functioning as oscula; they are water-collecting. The skeleton comprises equiangular and equiradiate triactines ( Fig. 26B ). Actines are conical or slightly conical, and we can separate two populations of spicules based on this difference, the spicules with conical actines being the most abundant. The tip of the actines is always sharp and the size of the spicules matches the size given by Haeckel:

We also found two other specimens in BMNH that matched this one exactly. One ( BMNH 1897.3.25.3) is from Lesina, while the other ( BMNH 1898.5.7.3) is from Naples. We did not find any specimens which we considered as morphologically similar to PMJ.Inv.Nr.Porif.154.

Considering that originally Haeckel (1870) had given Naples as the type locality of C. primordialis , we suggest that ZMB 1306 should become the lectotype. Contrary to his concept of the species’ great morphological variability, we think that it is very well defined morphologically, being recognized by the shape of the actines of both triactine populations, by the sharp tip and even by the size of the actines, which seems to be quite consistent (conical triactines: 91.9 (±5.8)/9.6 (±0.5); slightly conical triactines: 86.6 (±13.0)/11.3 (±0.7)). The specimen sent from Jena as a syntype of primordialis (Porif. 154) is probably a new species, as the size of the spicules and the presence of cylindrical actines in one of the populations of triactines are very distinctive. However, we decided not to describe it as a new species in the present article, since we only had one very old specimen to study.

The systematics of sponges with only one kind of spicule is always very difficult. Species of Clathrina whose skeletons are composed only of triactines are considered to be the most morphologically plastic and geographically widespread. Topsent (1936) considered that this high level of plasticity in clathrinas only involved triactines, and subsequently placed C. primordialis in synonymy with C. coriacea . Since then, except for C. clathrus , all other clathrinas with a skeleton composed only of triactines became C. coriacea . In the 1970s the name C. primordialis started being used again ( Borojevic, 1971; Borojevic & Peixinho, 1976). The synonymy of C. primordialis with C. coriacea was questioned when the authors compared specimens with triactines from Brazil with specimens of C. coriacea from the British Isles (locus typicus). The shape and size of the spicules were so different that they decided to call the Brazilian specimens C. primordialis . The main difference related to the tip of the actines, which had already been used by Haeckel in 1872 to distinguish specimens of C. primordialis (sharp) from C. coriacea (blunt). Klautau et al. (1994) also considered the shape of the actines to be a useful character to distinguish specimens from two sympatric populations of Clathrina from Arraial do Cabo (Rio de Janeiro). Allozyme analysis of specimens from both populations found that there was no gene flow between them, that they are reproductively isolated and thus constitute distinct species.

As the type specimens of Haeckel’s species, including C. primordialis , seemed at that time to be lost, the population with triactines with conical actines and a sharp tip was called C. primordialis . Therefore Klautau et al. (1994) suggested that the locus typicus of this species should be considered as Rio de Janeiro. Recently, however, we have been able to work with specimens originally identified by Haeckel. We believe that it is more appropriate to consider Naples as the locus typicus, which implies that the Brazilian population should be considered as a new species ( C. conifera ). The distribution of C. primordialis is thus restricted to the Mediterranean and the Adriatic Seas.

Figures

Figure 26. Clathrina primordialis. A, two categories of triactines. B, triactines of ZMB 1306. Scale bar = 100 Mm.